Inventory and taxonomic diversity of macroalgae from the coast of Jalisco, México

I. Enciso-Padilla1; E. Ríos-Jara1; M.C Esqueda-González1*

Correspondence: *. Corresponding Author: María del Carmen Esqueda González: Camino Ramón Padilla Sánchez 2100 Nextipac, Zapopan, Jalisco, México 45110. E-mail: E-mail: . Tel.: (01) 33 37 77 11 50 ext 33164 Departamento de Ecología, Centro Universitario de Ciencias Biológicas y Agropecuarias, Universidad de Guadalajara.


Abstract:

This study integrates an updated inventory based on previous and new information on the composition and distribution of the macroalgae of the Jalisco coast, obtained from three main sources: 1) Herbarium data, 2) Bibliographic information, and 3) Collection of specimens from different sites distributed along the coastline of this region performed between 2000 and 2018. All records were incorporated in 19 sites and included in a total of 263 species of three divisions, Chlorophyta (44 species), Ochrophyta (52) and Rhodophyta (167). The list of species was updated and ordered according to the registers of each site, including two new records: Solieria filiformis and Piropya raulaguilarii. Also, a high ecological rarity (incidence) was documented with the 38 % of unique species (recorded only in one locality) and 21 % of duplicate species (in two sites). The Rhodophyta had the highest rarity (29 % unique and 13 % duplicate).

The sampling effort was evaluated with species accumulation curves using non parametric estimators taking into consideration both the algae collected during the present research and all the historic records (herbarium and bibliographic data) from the intertidal, and the shallow and deeper subtidal environments of all sites. All curves tend to be asymptotic with a mean sampling efficiency of 70 %, compared to an expected maximum number of species of 475.

A taxonomic distinctness analysis using records of presence-absence of all the species, allowed to recognize the contribution of each locality to the taxonomic diversity of the coast of Jalisco. The largest number of species was found in sites of the center and northern coastline: Chamela Bay (105), Tenacatita (86), Puerto Vallarta (84) and Careyes (71). Together, the southern sites (Tenacatita, La Manzanilla, El Tamarindo, Cuastecomates, Melaque and Barra de Navidad) had the larger contribution to all taxonomic categories, from species to orders; while only Puerto Vallarta in the north, and Chamela and Careyes in the center, had a good taxonomic representation. The contribution of the coast of Jalisco to the overall inventory of macroalgae species from the tropical Mexican Pacific is 69 %, which highlights the importance of this region to the diversity of macroalgae of Mexico.

Received: 2019 January 29; Accepted: 2019 June 8

revbio. 2020 Mar 20; 6: e656
doi: 10.15741/rebvio.06.e656

Keywords: Key words: Mexican marine algae, species list, distribution, Jalisco, taxonomic distinctness..

Introduction

Phycological inventories include taxonomic, ecologic and biogeographic information of species of algae, and are indispensable to recognize the biodiversity of a given region. These inventories are the result of the analysis and the integration of new and previous information, which allows comprehensive evaluations of past and current data that constitute an excellent tool to identify short and long term changes in the ecosystems (González, 1992; Huisman et al., 1998; Giangrande, 2003). Information obtained from herbaria and the revisions of bibliographic references are critical sources of data for evaluations of the biodiversity and conservation of the macroalgal flora from a certain region. These data provide an opportunity to review the state of knowledge and to examine how this knowledge of has been built over time in terms of the number of sites sampled and the number of species recognized, and to identify where there are gaps in the current inventories as far as numbers of specimens per taxon, as well as with respect to geographical and seasonal coverage (Nelson et al., 2013). Additionally, a reliable evaluation requires the taxonomic update of the species involved in order to serve as a reference for future research in the same region and to make comparisons with other regions (Serviere-Zaragoza, 1993; Candelaria-Silva, 1996; León, 1996). For this, it is always necessary to know if the sampling effort is sufficient to have a reliable inventory taking into account the rarity of the species (Ceshia et al., 2007; Capetillo et al., 2015).

The diversity has been evaluated using the taxonomic distinctness indices (Warwick & Clarke, 1995), which allows to know the variability in the composition of the different taxonomic hierarchies including the class, order, family, genus and species levels. Taxonomic distinctness is a univariate (bio) diversity index which, in its simplest form, calculates the average “distance” between all pairs of different assemblages of species; it attempts to capture phylogenetic diversity rather than simple richness of species and is more closely linked to functional diversity; it is robust to variation in sampling effort because it utilizes only simple species lists based on presence/absence data (Clarke & Warwick, 1999). In addition, according to Mistri et al. (2000), the integration and use of historic data gathered from bibliographic revisions and scientific collections that contain only presence-absence records are often used to recognize the diversity, in contrast to other indexes because the taxonomic differentiation could use independent data from different sources and it is less reliant on the sampling effort (Heino, 2005).

The taxonomic diversity indices have been used to compare different environments (e.g. marine and freshwater) (Terlizzi et al., 2005; Heino et al., 2007), regions and latitudes (e.g. polar, temperate and tropical) (Conlan et al., 2004; Miranda et al., 2005; Nicholas & Trueman, 2005; Włodarska-Kowalczuk et al., 2005; Leonard et al., 2006), being also a useful tool for assessing diversity among different sites of a certain region. Recently, these indices were used to evaluate the taxonomic diversity of different marine groups of different regions of Mexican Pacific including stony corals (Reyes-Bonilla et al., 2013), echinoderms (Rios-Jara et al., 2013), bivalves (Esqueda-González et al., 2014) and fishes (Barjau-González et al., 2012; Herrera-Valdivia et al., 2016). However, evaluations of the structure of macroalgae communities using the taxonomic distinctness to measure the variability of the biodiversity among sites by using differences between hierarchical levels are scarce (e.g. Price et al., 2006; Tonetto et al., 2016).

The tropical Mexican Pacific is one of the regions with the greatest diversity of macroalgae species in the western America (Mendoza-González & Mateo-Cid, 1998). However, previous studies are restricted to very specific taxa (Carballo et al., 2002; Senties & Dreckmann, 2014; Lopez et al., 2017), or to adjacent subtropical regions of the Gulf of California (Reyna & Rodriguez, 1996; Serviere-Zaragoza et al., 2003; Aguilar-Rosas et al., 2010; Castañeda-Fernandez de Lara et al., 2010; Mora-Valdes & Riosmena-Rodriguez, 2016; Cossete et al., 2016).

The coast of Jalisco comprise an extensive region of approximately 350 km located in the central part of the Mexican tropical Pacific, and includes a great marine biodiversity in a variety of representative tropical habitats such as rocky and sandy beaches, coral and rocky reefs, estuaries and coastal lagoons with mangrove swamps. The coast of Jalisco has been recognized for its great biodiversity of macroalgae. The first phycological records in Jalisco were those of Dawson (1949). Subsequently, the list of species increased with a series of works performed by Dawson (1951, 1953a, 1953b, 1954, 1960, 1961a, 1961b, 1962, 1963b) and later complemented by Chávez-Barrera (1980), Pedroche & González-González (1981), Enciso-Padilla et al. (1995) and Águila-Ramírez et al. (1998). In addition, the works of Serviere-Zaragoza et al. (1993, 1998), Pedroche et al. (2005, 2008) and Rodríguez et al. (2008) incorporated new records and range extensions of several species. At that moment, a total of 77 species of marine algae were reported for the coast of Jalisco; in a later work, Mendoza-González et al. (2011) updated this inventory to 165 species, this is a 214 % increase in the number of species. Since then, there are no recent inventories that integrate the macroalgal flora of the coast of Jalisco.

The present study describes an up to date taxonomic composition of macroalgae and their distribution in 19 sites of the coast of Jalisco, with an evaluation of the taxonomic categories recorded in each locality using the average Taxonomic Distinctness index (Δ+) and its variation (Λ+) in order recognize the contribution of each locality to the taxonomic diversity of this region.

Material and Methods

Study area

The state of Jalisco is located in the central-west area of the Mexican tropical Pacific (19°−21° N, 104°−106° W) and has an approximate coastal extension of 350 km (De la Lanza, 1991). The continental shelf is narrow due to its proximity to the convergent limits between the Cocos and the North America plates. This region exhibit particular oceanographic characteristics due to the circulation patterns originated by the influence of two large main current systems, the California current from the North with temperate waters, and the Equatorial countercurrent from the South with warm waters; these two great currents converge off the coasts of Nayarit and Jalisco and then flow away from the continental mass offshore to the west (Wyrtki, 1966). The physiography of most of the coastline of Jalisco is characterized by having bays and open and semi-closed coves, with sandy and rocky beaches with some small islands and islets interspaced with coastal lagoons and estuaries with mangrove swamps; in the central area there is a long sandy strip beach of approximately 50 km interrupted by coastal lagoons and several rocky beaches. Other representative tropical habitats are also present along the coastline such as coral and rocky reefs. This combination of marine environments produces a great heterogeneity that influences the structure and composition of the algae communities.

The information on macroalgae records of the coast of Jalisco was integrated from three sources:

Herbarium records

The macroalgal floral of five national herbaria was compiled: a) MEXU (Instituto de Biología, Universidad Nacional Autónoma de México); b) FCME, (Facultad de Ciencias, Universidad Nacional Autónoma de México); c) ENCB (Escuela Nacional de Ciencias Biológicas, Instituto Politécnico Nacional); d) GUADA (Universidad Autónoma de Guadalajara); and e) IBUG (Instituto de Botánica, Universidad de Guadalajara). A revision of material collected from the coast of Jalisco was performed in each herbarium, recording the following information: scientific name, date of collection, locality and environmental variables (intertidal level, type of substrate).

Bibliographic information

An exhaustive research of publications containing reports of species and their supra-specific categories was performed with the aim of integrating and systematizing the information on species of marine algae that have been reported in the coast of Jalisco. The research of documentary information included the revision of printed editions (books, journals, catalogs), together with the consultation of databases of both multidisciplinary and specialized electronic publications (for example ISI Web, EBSCO, ProQUEST). The use of the combination of key words (i.e. inventories, listings, distribution, macroalgae, phycoflora, algae, Jalisco) was used to refine the search depending on the language of the publication.

Collection of specimens

Sampling campaigns in 19 sites distributed along the coast of Jalisco were performed throughout all months of the year from 2000 to 2018 (19 years) (Figure 1). However, the most intensive collections were performed from March to June, during the dry season; and from September to November, during the rainy season. Collections were done along the intertidal and shallow subtidal zones (<10 m deep). Freediving (snorkel) was performed during direct collections in shallow waters (< 3 m) while SCUBA diving in deeper areas (<15 m depth). Environmental data such as wave exposure, depth, specific habitat (intertidal pool, current canal, rocky platform or cliff) and the type of substrate was associated to each specimen.


[Figure ID: f1] Figura 1.

Área de estudio y localidades de recolecta en la costa de Jalisco, México.


The algae collected were deposited in properly labeled plastic bags with seawater and then transported to the Laboratorio de Ecosistemas Marinos y Acuicultura (LEMA) of the Departmento de Ecología of the Universidad de Guadalajara (UdeG). One part of the samples was herborized and the other preserved in a 4 % solution of formaldehyde and seawater. Histological analysis was performed with the latter specimens for taxonomic determination, together with examination of external and internal morphology and reproductive characteristics with the aid of an optic and a stereoscope microscope. Taxonomic keys were used together with specialized bibliography such as Dawson (1944, 1947, 1953a, 1960, 1961a, 1961b, 1962), Taylor (1960), Abbott & Hollenberg (1976) and Norris and Johansen (1981), and the consultation of the database of the webpage AlgaeBase (2018). Herborized specimens were deposited in the Macroalgae collection of the Laboratorio de Ecología Molecular, Microbiología and Taxonómia (LEMITAX) of the Departamento de Ecología (UdeG).

Data Analysis

A database that compiles the composition and distribution of algae species from the coast of Jalisco was completed with an inventory of species obtained from herbaria records, documentary analysis and the collections from all sites. New macroalgae records were recognized using historic data and from new collections.

To perform the sampling effort and the Taxonomic Distinctness analysis, all the historic information and the new data from collections were grouped in 19 different sites of the north, center and south zones of Jalisco. These sites are more or less equidistant and representative of the coastal habitats of the Mexican tropical Pacific.

The sampling effort was evaluated considering the algae collected during sampling campaigns and those of historical records of the 19 selected locations of the coast of Jalisco. Species accumulation curves were done that were built with 10,000 iterations without replacement; the expected richness was calculated using the non-parametric estimators Chao 2, Jackknife 1 and Jackknife 2 by using the software EstimateS V8 (Colwell, 2006).

For the analysis of Taxonomic Distinctness (Clarke & Gorley, 2006) a presence-absence matrix for all the species recorded at 19 selected sites was completed; afterwards, a taxonomic aggregation matrix was integrated with six taxonomic levels: Division, Class, Order, Family, Genera and Species. These categories were weighted according to Warwick & Clarke (1995) in which w1 belong to species from the same genera; w2, species within the same family but different genera; w3, species within the same order but different family, and so on. The average taxonomic distinctness (Δ+) and its variation (Λ+) were estimated for each locality.

Results

Sampling effort

All the accumulation curves of species tended towards the asymptote, indicating a good representation of the sampling effort made in the 19 selected sites of Jalisco (Figure 2). The observed species (363) represented between 65 and 74 % of the theoretical number of expected species, with an average value of 70 %. The Jackknife 2 estimator had the highest number of expected species with 403 species and, therefore, the lowest representation (66 %), while Jackknife 1 and Chao 2 had intermediate values of 359 (73 %) and 355 (74 %) ) expected species, respectively. This due to the high variability in the total number of species recorded and strong differences in species rarity among sites.


[Figure ID: f2] Figure 2.

Accumulation curves of observed and expected macroalgae species from the coast of Jalisco, Mexico. The expected curves were estimated with the non-parametric indexes Chao 2, Jackknife 1 and Jackknife 2. The projections were constructed with 10,000 non-replacement iterations.


A high ecologic rarity (incidence) of the macroalgae was registered in Jalisco. This was evident when a more detailed revision of the observed rarity of the macroalgae species was performed, showing that an important number of species were exclusive of a single site. In fact, a total of 101 species (38 %) were unique (recorded in one site) (e.g. Asteronema breviarticulatum, Sargassum pacificum, Chondria repens, Melanothamnus bajacali, Gelidium arborescens, Amphiroa foliacea, Ulva lobata, Cladophora sericea) and, 56 (21 %) were duplicate species (recorded in two sites) (e.g. Dictyota pinnata, Ectocarpus simulans, Gracilaria tepocensis, Ulva californica, Cladophora prolifera, Bryopsis corticulans, Codium giraffa, Chlorodesmis hildebrandtii). The highest ecological rarity of the species was in the Rhodophyta (29% unique and 13 % duplicate). Also, the highest number of both unique and duplicate species was recorded in Chamela (42), Tenacatita (33), El Tamarindo (27 and Puerto Vallarta (24). In the remaining sites ecological rarity was between 1 and 21 species, with the exception of Punta Pérula with no rare species recorded.

Richness and distribution of species in the coast of Jalisco

The integration of the information gathered from three sources (revisión of herbaria material, documentary information and collections) shows a species richness of 263 macroalgae species for the coast of Jalisco, integrated in 109 genera and 61 families. The list of species is shown in Table 1, the taxonomic arrangement follows that from Silva et al. (1996) and complemented by that proposed by Wynne (1986, 2017).

Table 1.

Systematic list of macroalgae species and their distribution in 19 sites of the coast of Jalisco. 1=Puerto Vallarta, 2=Yelapa, 3=Chimo, 4=Bahía Corrales, 5=Playitas, 6=Cabo Corrientes, 7=Mayto, 8=Tehuamixtle, 9= Chalacatepec, 10=Punta Pérula, 11=Chamela, 12=Isla Cocinas, 13=Careyes, 14= Tenacatita, 15=La Manzanilla, 16=El Tamarindo, 17=Cuastecomates, 18=Melaque, 19= Barra Navidad. * unique species; ** duplicates species; + new record.


DIVISION CHLOROPHYTA
 CLASE ULVOPHYCEAE
 ORDEN BRYOPSIDALES
  Familia Bryopsidaceae
  Bryopsis corticulans Setchell 1903 (11, 14) **
  Bryopsis galapaguensis W.L.Taylor 1945 (14,16) **
  Bryopsis hypnoides J.V. Lamouroux 1809 (6, 9, 11, 18)
  Bryopsis pennata var. minor J. Agardh 1887 (1, 9, 11, 12, 14, 16, 19) **
  =Bryopsis pennatula J. Agardh 1847
  Derbesia marina (Lyngbye) Solier 1846 (1, 11, 13)
  Familia Caulerpaceae
  Caulerpa chemnitzia (Esper) J.V.Lamouroux 1809 (11, 13, 14, 15, 16, 17, 19)
  =Caulerpa peltata J.V. Lamouroux 1809
  Caulerpa sertularioides (S.G. Gmelin) Howe 1916 (7, 8, 12, 13, 14)
  Familia Codiaceae
  Codium dichotomun (Hudson) S. F. Gray 1821 (9, 11, 12, 13, 16)
  Codium giraffa Silva 1979 (9, 11) **
  Codium isabelae W.R. Taylor 1945 (1, 11, 15, 16) **
  = Codium santamariae W.R Taylor 1945
  Codium picturatum Pedroche et P.C. Silva 1996 (1, 14, 15)
  Codium setchelli N.L. Gardner 1919 (11, 12, 15)
  Codium simulans Setchell et N.L. Gardner 1924 (1, 2, 16, 17) **
  =Codium cuneatum Setchell et N.L. Gardner 1924
  Familia Halimedaceae
  Halimeda discoidea Decaisne 1842 (7, 8, 9, 10, 11, 12, 13, 14, 16)
  Familia Udoteaceae
  Chlorodesmis hildebrandtii A. Geep et E. Geep 1911 (11, 14) **
 ORDEN CLADOPHORALES
  Familia Cladophoraceae
  Chaetomorpha antennina (Bory) Kützing 1847 (1, 2, 3, 4, 6, 7, 8, 10, 11, 12, 13, 14, 15, 16, 17, 18, 19)
  =Chaetomorpha media (C. Agardh) Kützing 1849
 vChaetomorpha gracilis Kützing 1845 (19) *
  Chaetomorpha linum (Müller) Kützing 1845 (1, 16, 18)
  Cladophora albida (Hudson) Kützing 1843 (1) *
  Cladophora coelothrix Kützing 1843 (11, 19) *
  =Cladophora repens Harvey1849
  Cladophora laetiverens (Dillwyn) Kützing 1843 (11, 13, 16)
  Cladophora microcladioides Collins 1909 (1, 16) **
  Cladophora prolifera (Roth) Kützing 1843 (1, 14) **
  Cladophora sericea (Hudson) Kützing 1843 (19) *
  Cladophora vagabunda (Linneaus) van den Hoek 1963 (16, 19) **
  Rhizoclonium riparium (Roth) Harvey 1849 (16, 19) **
  =Rhizoclonium kerneri Stockmayer 1890
  ORDEN DASYCLADALES
  Familia Polyphysaceae
  Parvocaulis parvulus (Solms-Laubach) S.Berger, Fettweiss, Gleissberg, Liddle, U.Richter, Sawitzky et Zuccarello 2003 (11, 13, 14)
  =Polyphysa parvula (Solms-Laub) Schnetter et Bula-Meyer 1982
  =Polyphysa parvula (Solms-Laub) Schnetter et Bula-Meyer 1982
 ORDEN SIPHONOCLADALES
  Familia Siphonocladaceae
  Boodlea composita (Harvey) Brand 1904 (15) *
  Phyllodictyon anastomosans (Harvey) Kraft et Wynne 1996 (11) *
  ORDEN ULVALES
  Familia Ulvaceae
  Ulva californica Wille 1909 (7, 11) **
  Ulva clathrata (Roth) C.Agardh 1811 (16, 17, 19) **
  =Enteromorpha crinita Nees 1820
  Ulva compressa Forsskål 1753 (11, 12, 14, 15, 18)
  =Enteromorpha compressa (Linneaus) Greville 1830
  Ulva expansa (Setchell) Setchell et N.L. Gardner 1920 (4, 5, 7, 8, 13, 16)
  Ulva fasciata Delile 1914 (1, 6) **
  Ulva flexuosa Wulfen 1803 (4, 5, 7, 8, 11, 13, 16, 17, 19)
  =Enteromorpha flexuosa (Wulfen) J. Agardh 1883
  Ulva intestinalis Linnaeus 1753 (1, 8, 9, 13, 15, 16, 19)
  =Enteromorpha intestinalis (Linneaus) Nees 1820
  Ulva lactuca Linnaeus 1753 (1, 2, 4, 5, 8, 9, 11, 12, 13, 14, 16, 17, 18)
  Ulva linza Linnaeus 1753 (1, 6) **
  =Enteromorpha linza (Linneaus) J. Agardh 1883
  Ulva lobata (Kützing) Setchell et N.L. Gardner 1920 (13) *
  Ulva prolifera O.F.Müller 1778 (1, 6, 11, 16)
  =Enteromorpha prolifera (Müller) J. Agardh 1883
  =Enteromorpha salina Kützing 1845
  Ulva rigida C. Agardh 1823 (11, 13, 14, 15, 17, 18, 19)
  Ulva taeniata (Setchell) Setchell et N.L. Gardner 1920 (6, 8, 11, 13, 14, 16, 18)
  =Ulva dactylifera Setchell et N.L. Gardner 1920
  Familia Ulvellaceae
  Phaeophila dendroides (P. Crouan et H. Crouan) Batters 1902 (11) *
  Ulvella viridis (Reinke) R.Nielsen, C.J.O’Kelly et B.Wysor in R.Nielsen et al. 2013 (19)*
  =Acrochaete viridis (Reinke) R.Nielsen 1979
DIVISION OCHROPHYTA
 CLASE PHAEOPHYCEAE
  ORDEN DICTYOTALES
  Familia Dictyotaceae
  Dictyopteris delicatula J.V. Lamouroux 1809 (14, 15, 16)
  Dictyota bartayresiana J.V. Lamouroux 1809 (10, 13, 14)
  Dictyota ciliolata Kützing 1859 (10, 11, 13)
  Dictyota crenulata J. Agardh 1847 (1, 2, 3, 11, 13, 14, 15, 18)
  Dictyota dichotoma (Hudson) J.V. Lamouroux 1809 (8, 10, 12, 16)
  Dictyota dichotoma var. intricata (C. Agardh) Greville 1830 (13, 14, 17)
  Dictyota implexa (Desfontaines) J.V.Lamouroux 1809 (1, 10, 13, 14, 16)
  =Dictyota divaricata J.V. Lamouroux 1809
  Dictyota flabellata (Collins) Setchell et N.L. Gardner 1924 (17, 18) **
  Dictyota friabilis Setchell 1926 (4, 11, 17)
  =Dictyota pfaffii Schnetter 1972
  Dictyota pinnata (E.Y.Dawson) I.Hörnig, R.Schnetter et Prud’homme van Reine 1993 (11, 13, 17)
  =Dilophus pinnatus E.Y. Dawson 1950
  Lobophora variegata (J.V. Lamouroux) Womersley ex Oliveira 1977 (13, 15)
  **
  Padina caulescens (Thivy) W.R. Taylor 1945 (1, 9, 10, 11, 12, 13, 14, 15, 16, 19)
  Padina concrescens Thivy 1945 (8, 11) **
  Padina crispata (Thivy) W.R. Taylor 1945 (1, 2, 4, 9, 11, 13, 15, 17, 18)
  Padina durvillaei Bory 1827 (1, 2, 3, 4, 6, 11, 13, 14, 16)
  Padina gymnospora (Kützing) Sonder 1871 (1, 2, 4, 9, 10, 11, 12, 13, 14, 15, 16, 17, 18, 19)
  =Padina vickersiae Hoyt 1920
  Padina mexicana E.Y. Dawson 1944 (1, 5, 7, 12, 14)
  Padina pavonica J.V. Lamouroux 1816 (8, 17,18)
  Zonaria farlowii Setchell et N.L. Gardner 1924 (2, 10, 11, 12, 13, 17, 18)
 ORDEN ECTOCARPALES
  Familia Chordariaceae
  Compsonema serpens Setchell et N.L. Gardner 1922 (1) *
  Hecatonema streblonematoides (Setchell et N.L.Gardner) Loiseaux 1970 (1, 11, 15)
 v=Compsonema secundum Setchell et N.L. Gardner 1922
  Hecatonema terminale (Kützing) Kylin 1937 (14) *
  Streblonema penetrale Setchell et N.L. Gardner 1922 (18) *
  Familia Acinetosporaceae
  Feldmannia mitchelliae (Harvey) H.-S.Kim 2010 (13, 15, 17, 19)
  =Hincksia mitchelliae (Harvey) P.C.Silva 1987
  =Giffordia mitchelliae (Harvey) Hamel 1939
  Familia Ectocarpaceae
  Ectocarpus parvus (Saunders) Hollenberg 1971 (1, 11, 18)
  Ectocarpus rallsiae Vickers 1905 (13) *
  =Hincksia rallsiae (Vickers) P.C.Silva 1987
  Ectocarpus siliculosus (Dillwyn) Lyngbye 1819 (11, 14) **
  Ectocarpus simulans Setchell et N.L. Gardner 1922 (1, 11) **
  Hincksia saundersii (Setchell et N.L. Gardner) P.C.Silva in P.C. Silva, E.G. Meñez et Moe 1987 (16) *
  =Giffordia saundersii (Setchell et N.L. Gardner) Hollenberg et Abbott 1966
 ORDEN FUCALES
  Familia Sargassaceae
  Sargassum agardhianum Farlow 1889 (1) *
  Sargassum alternato-pinnatum Yamada 1942 (1, 2) **
  =Sargasum asymmetricum E.Y. Dawson 1944
  Sargassum brandegeei Setchell et N.L. Gardner 1924 (11) *
  Sargassum horridum Setchell et N.L. Gardner 1924 (11) *
  Sargassum howellii Setchell 1937 (1, 2, 3, 4, 7, 8, 10, 11, 12, 14, 15, 16, 17, 18, 19)
  Sargassum liebmanii J. Agardh 1847 (1, 2, 3, 4, 6, 8, 10, 11, 12, 13, 14, 15, 16, 17, 18, 19)
  Sargassum pacificum Bory 1828 (1) *
 ORDEN RALFSIALES
  Familiy Hapalospongidiaceae
  Hapalospongidion gelatinosum Saunders 1899 (18) *
  Familia Neoralfsiaceae
  Neoralfsia hancockii (E.Y.Dawson) D.Leon-Alvarez et M.L.Nunez-Reséndiz in Leon-Alvarez et al. 2014 (13, 14, 15)
  = Ralfsia hancockii E.Y.Dawson 1944 Familia Ralfsiaceae
  Ralfsia confusa Hollenberg 1969 (1, 4, 11, 14)
  Ralfsia fungiformis (Gunner) Setchell et N.L. Gardner 1924 (11, 18) **
  Ralfsia hesperia Setchell et N.L. Gardner 1924 (1, 4, 11, 18)
  Ralfsia pacifica Hollenberg 1944 (1, 17, 18)
 ORDEN SCYTOSIPHONALES
  Familia Chnoosporaceae
  Chnoospora minima (Hering) Papenfuss 1956 (1, 2, 3, 4, 9, 10,11, 12, 13, 14, 15, 16, 17, 18)
  =Chnoospora pacifica J. Agardh 1847
  Familia Scytosiphonaceae
  Colpomenia sinuosa (Roth) Derbés et Solier 1856 (1, 16) **
  Petalonia fascia (O.F.Müller) Kuntze 1898 (14) *
  =Ralfsia californica Setchell et N.L. Gardner 1924
 ORDEN SCYTOTHAMNALES
  Familia Bachelotiaceae
  Bachelotia antillarum (Grunow) Gerloff 1959 (14) *
  Familia Asteronemataceae
  Asteronema breviarticulatum (J. Agardh) Oriques et Bouzon 2000 (11, 15, 17, 18, 19)*
  = Ectocarpus breviarticulatus J. Agardh 1847 (11)*
 ORDEN SPHACELARIALES
  Familia Lithodermateceae
  Pseudolithoderma nigrum Hollenberg 1969 (17) *
  Familia Sphacelariaceae
  Sphacelaria californica Sauvageau 1925 (1) *
  Sphacelaria tribuloides Meneghini 1840 (16) *
  =Sphacelaria mexicana W.R.Taylor 1945
  Sphacelaria novae-hollandiae Sonder 1845 (11, 13) **
  Sphacelaria rigidula Kützing 1843 (1, 2, 11, 19)
DIVISON RHODOPHYTA
 CLASE BANGIOPHYCEAE
  ORDEN BANGIALES
  Familia Bangiaceae
  Bangia fuscopurpurea (Dillwyn) Lyngbye 1819 (11, 14) **
  Piropya raulaguilarii (5)* +
CLASE FLORIDEOPHYCEAE
 ORDEN ACROCHAETIALES
  Familia Acrochaetiaceae
  Acrochaetium hancockii (E.Y.Dawson) Papenfuss 1945 (11, 15) **
  Acrochaetium microscopicum (Nägeli ex Kützing) Nägeli 1858 (14, 15) **
 ORDEN AHNFELTIALES
  Familia Ahnfeltiaceae
  Ahnfeltia plicata (Hudson) Fries 1835 (6, 12, 13, 18)
  Ahnfeltia svensonii W.R. Taylor 1945 (1, 2, 6, 11, 13, 14, 15, 17, 18)
 ORDEN BONNEMAISONIALES
  Familia Bonnemaisoniaceae
  Asparagopsis taxiformis (Delile) Trevis 1845 (11, 14) **
 ORDEN CERAMIALES
  Familia Callithamniaceae
  Aglaothamnion endovagum (Setchell et N.L.Gardner) I.A.Abbott 1972 (11) *
  =Callithamnion endovagum Setchell et N.L.Gardner 1924
  Callithamnion rupiculum C.L.Anderson 1894 (11) *
  Familia Ceramiaceae
  Antithamnionella breviramosa (E.Y. Dawson) Womersley et Bailey 1970 (1) *
  Antithamnionella elegans (Berthold) J.H.Price et D.M.John 1986 (14, 19) **
  Centroceras clavulatum (C. Agardh) Montagne 1846 (1, 2, 4, 5, 6, 7, 8, 10, 11, 12, 13, 14, 15, 16, 17, 18)
  Ceramium affine Setchell et N.L. Gardner 1930 (14) *
  Ceramium camouii E.Y Dawson 1944 (19) *
  Ceramium caudatum Setchell et N.L. Gardner 1924 (14, 16) **
  Ceramium hamatispinum E.Y. Dawson 1950 (13, 15, 19)
  Ceramium zacae Setchell et N.L. Gardner 1937 (15, 16, 19)
  Gayliella flaccida (Harvey ex Kützing) T.O.Cho et L.J.McIvor 2008 (1, 4, 11, 13, 14, 15, 16, 17, 18, 19)
  =Ceramium flaccidum (Kützing) Ardissone 1871
  Gayliella fimbriata (Setchell & N.L. Gardner) T.O.Cho et S.M.Boo 2008 (11, 14, 15, 18)
  =Ceramium fimbriatum Setchell et N.L. Gardner 1924
  Gayliella taylorii (E.Y. Dawson) T.O.Cho et S.M.Boo 2008 (5, 8, 11, 12, 13, 14, 15)
  =Ceramium taylorii E.Y. Dawson 1950
  Familia Dasyaceae
  Dasya sinicola var. abyssicola (E.Y. Dawson) E.Y. Dawson 1963 (11) *
  Familia Delesseriaceae
  Taenioma perpusillum (J. Agardh) J. Agardh 1848 (11, 15) **
  Familia Rhodomelaceae
  Bostrychia moritziana (Sonder ex Kützing) J.Agardh 1863 (19) *
  Chondria acrorhizophora Setchell et N.L.Gardner 1924 (2, 6, 15, 19)
  =Chondria californica (Collins) Kylin 1941
  Chondria arcuata Hollenberg 1945 (1, 2, 16)
  Chondria repens Borgesen1924 (15) *
  Herposiphonia littoralis Hollenberg 1970 (11, 16) **
  Herposiphonia secunda (C.Agardh) Ambronn 1880 (16, 17) **
  Herposiphonia subdisticha Okamura1899 (14) *
  Herposiphonia tenella (C. Agardh) Ambronn 1880 (1, 2, 11, 16)
  Herposiphonia verticillata (Harvey) Kylin 1925 (1) *
  Laurencia clarionensis Setchell et N.L. Gardner 1937 (15) *
  Laurencia decidua E.Y. Dawson 1954 (11, 13) **
  Laurencia gardnerii Hollenberg 1943 (17) *
  Laurencia hancockii E.Y.Dawson 1944 (14, 16) **
  Laurencia humilis Setchell et N.L Gardner 1930 (14, 16) **
  Laurencia lajolla E.Y. Dawson 1958 (11) *
  Melanothamnus bajacali (Hollenberg) Díaz-Tapia et Maggs in Díaz-Tapia et al. 2017 (16) *
  =Polysiphonia bajacali Hollenberg 1961
  Melanothamnus johnstonii (Setchell et N.L.Gardner) Díaz-Tapia et Maggs in Díaz-Tapia et al. 2017 (14, 16) **
  =Polysiphonia johnstonii Setchell et N.L. Gardner 1924
  Melanothamnus savatieri (Hariot) Díaz-Tapia et Maggs in Díaz-Tapia et al. 2017 (15) *
  =Polysiphonia japonica var. savatieri (Hariot) H.Y.Yoon 1986
  Melanothamnus simplex (Hollenberg) Díaz-Tapia et Maggs 2017 (1, 14, 15, 17, 18, 19)
  =Neosiphonia simplex (Hollenberg) Y.P.Lee 2008
  =Polysiphonia simplex Hollenberg 1942
  Melanothamnus sphaerocarpus (Borgesen) Díaz-Tapia et Maggs in Díaz-Tapia et al. 2017 (13) *
  =Neosiphonia sphaerocarpa (Borgesen) M.S.Kim et I.K.Lee 1999
  Murrayellopsis dawsonii Post 1962 (14, 16) **
  Osmundea sinicola (Setchell et N.L. Gardner) K.W.Nam 1944 (13) *
  Polysiphonia confusa Hollenberg 1961 (14, 16) **
  Polysiphonia hendryi N.L. Gardner 1927 (16) *
  Polysiphonia mollis Hooker et Harvey 1947 (19) *
  Polysiphonia pacifica Hollenberg 1942 (16, 17, 18)
  Polysiphonia sertularioides (Grateloup) J. Agardh 1863 (14) *
  Polysiphonia subtilissima Montagne 1840 (19) *
  Symphyocladia dendroidea (Montagne) Savoie et G.W.Saunders 2016 (14, 16) **
  =Pterosiphonia dendroidea (Montagne) Falkenberg 1901
  Tayloriella dictyurus (J. Agardh) Kylin 1941 (1, 2, 3, 4, 6, 11, 13, 14, 15, 16, 17)
  Familia Wrangeliaceae
  Anotrichium tenue (C.Agardh) Nägeli 1862 (11) *
  Pleonosporium globuliferum Levring 1941 (11) *
 ORDEN COLACONEMATALES
  Familia Colaconemataceae
  Colaconema daviesii (Dillwyn) Stegenga 1985 (1, 2, 15, 19)
  =Acrochaetium daviesii (Dillwyn) Nägeli 1862
  Colaconema hypneae (Borgesen) A.A.Santos et C.W.N.Moura 2010 (14, 15, 19)
  =Acrochaetium hypneae (Borgesen) Borgesen 1915
  Colaconema savianum (Meneghini) R.Nielsen 1994 (11, 13, 19)
  =Acrochaetium savianum (Meneghini) Nägeli 1862
  Grania pectinata (Kylin) Athanasiadis 2016 (14) *
  =Colaconema pectinatum (Kylin) J.T.Harper et G.W.Saunders 2002
 ORDEN CORALLINALES
  Familia Corallinaceae
  Amphiroa beauvoisii J.V.Lamouroux 1816 (1, 2, 3, 11, 12, 13, 14, 15, 16, 17, 18)
  =Amphiroa drouetii E.Y.Dawson 1953
  =Amphiroa mexicana W.R. Taylor 1945
  Amphiroa dimorpha Lemoine 1929 (1, 2, 3, 5, 7, 8, 9, 11, 12, 13, 14, 16, 17, 18)
  Amphiroa foliacea J.V. Lamouroux 1824 (3) *
  Amphiroa franciscana W.R. Taylor 1945 (16) *
  Amphiroa misakiensis Yendo 1902 (1, 11, 13, 14, 16, 17, 18)
  = Amphiroa brevianceps E.Y.Dawson 1953 Amphiroa polymorpha Lemoine 1929 (1) *
  Amphiroa rigida J.V. Lamouroux 1816 (1, 2, 16)
  Amphiroa subcylindrica E.Y. Dawson 1946 (18) *
  Amphiroa taylorii E.Y. Dawson 1934 (2, 16) **
  Amphiroa valonioides Yendo 1902 (1, 13, 18)
  Bossiella californica (Decaisne) Silva 1957 (17) *
  Crusticorallina muricata (Foslie) P.W.Gabrielson, Martone, K.R.Hind et C.P.Jensen in Hind et al. 2016 (2) *
  = Lithophyllum lichenare Mason 1953
  Fosliella paschalis (Lemoine) Setchell et N.L. Gardner 1930 (2) *
  Hydrolithon farinosum (J.V. Lamouroux) Penrose et Chamberlain 1993 (1, 2, 11, 14)
  =Fosliella farinosa (J.V. Lamouroux) Howe 1920
  Jania adhaerens J.V. Lamouroux 1816 (1, 2, 4, 10, 11, 12, 13, 14, 16, 17, 19)
  Jania capillacea Harvey 1853 (18) *
  Jania longiarthra E.Y. Dawson 1949 (11, 16) **
  Jania pacifica Areschoug 1852 (1, 2, 3, 4, 11, 13, 14, 15, 16, 18, 19)
  =Jania mexicana W.R. Taylor 1945
  = Titanoderma corallinae (P.Crouan et H.Crouan) Woelkerling, Y.M. Chamberlain et P.C.Silva 1985
  Lithophyllum decussatum (J.Ellis et Solander) Philippi 1837 (17) *
  Lithophyllum dispar (Foslie) Foslie 1909 (1, 15, 17)
  =Tenarea dispar (Foslie) Adey 1970 Lithophyllum imitans Foslie 1909 (1, 2, 18)
  Lithophyllum proboscideum (Foslie) Foslie 1900 (18) *
  Lithophyllum stictaeforme (Areschoug) Hauck 1877 (11) *
  Melobesia polystromatica E.Y. Dawson 1954 (14) *
  Melobesia membranacea (Esper) J.V. Lamouroux1812 (11, 14) **
  Neogoniolithon setchellii (Foslie) Adey 1970 (1) *
  Neogoniolithon trichotomum (Heydrich) Setchell et N.L. Gardner 1943 (1, 14) **
  Familia Hydrolithaceae
  Pneophyllum fragile Kützing 1843 (13) *
  Pneophyllum nicholsii (Setchell et L.R. Mason) P.C.Silva et P.W.Gabrielson 2004 (1, 2, 6)
  = Heteroderma nicholsii Setchell et N.L. Gardner 1943
  Familia Lithophyllaceae
  Titanoderma pustulatum (J.V.Lamouroux) Nägeli in Nägeli et Cramer 1858 (14, 15) **
  =Titanoderma pustulatum var. confine (P.Crouan et H.Crouan) Y.M.Chamberlain 1991
  Familia Lithothamniaceae
  Phymatolithon lenormandii (Areschoug) Adey 1966 (15) *
  Familia Porolithaceae
  Dawsoniolithon conicum (E.Y.Dawson) Caragnano, Foetisch, Maneveldt et Payri in Caragnano et al 2018 (14, 15) **
  =Pneophyllum conicum (E.Y. Dawson) Keats,Y.M.Chamberlain et M.Baba 1997
  Harveylithon rupestre (Foslie) A.Rösler, Perfectti, V.Peña et J.C.Braga 2016 (11) *
  =Hydrolithon rupestre (Foslie) Penrose 1996
  Harveylithon samoënse (Foslie) A.Rösler, Perfectti, V.Peña et J.C.Braga 2016 (13) *
  =Hydrolithon samoënse (Foslie) Keats et Y.M.Chamberlain 1994
  Porolithon onkodes (Heydrich) Foslie 1909 (14) *
  =Hydrolithon onkodes (Heydrich) Penrose et Woelkerling 1992
  Familia Spongitaceae
  Spongites decipiens (Foslie) Y.M. Chamberlain 1993 (1, 14, 18, 15)
  =Hydrolithon decipiens (Foslie) Adey 1970
  =Lithophyllum decipiens (Foslie) Foslie 1900
 ORDEN HALYMENIALES
  Familia Halymeniaceae
  Grateloupia clarionensis (Setchell et N.L. Gardner) S.Kawaguchi & H.W.Wang 2001 (14, 19) **
  Grateloupia doryphora (Montagne) Howe 1914 (4, 6, 11, 13, 14, 16)
  = Grateloupia multiphylla E.Y. Dawson 1954
  Grateloupia filicina (J.V. Lamouroux) C. Agardh 1822 (1, 16) **
  Grateloupia hancockii E.Y Dawson 1944 (11) *
  Grateloupia huertana Mateo-Cid, Mendoza-González et Gavio 2005 (11, 14, 15)
  Grateloupia howeii Setchell et N.L. Gardner 1924 (1, 2, 4, 11)
  Grateloupia prolongata J. Agardh 1847 (13, 16) **
  Grateloupia versicolor (J. Agardh) J. Agardh 1851 (1, 11, 13, 15, 19)
  Grateloupia sp. (5, 7, 8)
  Prionitis mexicana E.Y Dawson 1944 (11) *
  Familia Endocladiaceae
  Gloiopeltis furcata (Postels et Ruprecht) J. Agardh 1851 (17) *
  Familia Peyssonneliaceae
  Cruoriella fissurata E.Y.Dawson 1953 (14) *
  Peyssonnelia orientalis (Weber-van Bosse) Cormaci et G.Furnari 1987 (1, 6, 15)
  Peyssonnelia pacifica Kylin 1925 (15) *
 ORDEN ERYTHROPELTIDALES
  Familia Erythrotrichiaceae
  Erythrotrichia carnea (Dillwyn) J. Agardh 1883 (3, 13, 14, 15, 18)
  Erythrotrichia tetraseriata N.L. Gardner 1927 (11, 13) **
  Sahlingia subintegra (Rosenvinge) Kornmann 1989 (1, 2, 14)
  Smithora naiadum (Anderson) Hollenberg 1959 (6) *
 ORDEN GELIDIALES
  Familia Gelidiaceae
  Gelidium arborescens N.L. Gardner 1927 (16) *
  Gelidium crinale (Turner) Gaillon1825 (13) *
  Gelidium mcnabbianum (E.Y Dawson) B.Santelices 1998 (19) *
  Gelidium microdentatum E.Y Dawson 1960 (4, 6, 9, 11)
  Gelidium pusillum (Stackhouse) Le Jolis 1863 (1, 2, 6, 8, 9, 10, 11, 12, 13, 14, 17, 18)
  Gelidium sclerophyllum W.R. Taylor 1945 (1, 3, 6, 9, 13, 14, 15, 17, 18, 19)
  Gelidium sp. (5, 7, 8)
  Familia Gelidiellaceae
  Gelidiella acerosa (Forkssal) J. Feldmann et Hamel 1934 (1) *
  Gelidiella hancokii E.Y.Dawson1944 (19) *
  Parviphycus antipae (Celan) B. Santelices 2004 (15) *
  Familia Pterocladiaceae
  Pterocladiella caloglossoides (M.Howe) Santelices 1998 (1, 18) **
  =Pterocladia caloglossoides (M.Howe) E.Y. Dawson 1953
  Pterocladiella capillacea (S.G.Gmelin) Santelices et Hommersan 1997 (4) *
  =Pterocladia capillacea (S.G.Gmelin) Bornet 1876
  Pterocladiella media (E.Y. Dawson) Boo & Miller in Boo et al. 2006 (2) *
  =Pterocladia media E.Y. Dawson 1958
 ORDEN GIGARTINALES
  Familia Gigartinaceae
  Chondracanthus harveyanus (Kützing) Guiry in Hommersand, Guiry, Fredericq et Leister 1993 (6) *
  =Gigartina harveyana (Kützing) Setchell et Gardner 1933
  Chondracanthus spinosus (Kützing) Guiry in Hommersand, Guiry, Fredericq et Leister 1993 (6, 17) **
  =Gigartina asperifolia J. Agardh 1899
  =Gigartina spinosa (Kützing) Harvey 1853
  Mazzaella leptorhynchos (J.Agardh) Leister in Hommersand et al. 1993 (16, 17) **
  =Gigartina leptorhynchos J. Agardh 1885
  Mazzaella volans (C.Agardh) Fredericq in Hommersand et al 1993 (17) *
  =Gigartina volans (C. Agardh) J. Agardh 1846 Familia Hypneaceae
  Hypnea johnstonii Setchell et Gardner 1924 (1, 3, 4, 6, 11, 13, 15, 17)
  Hypnea pannosa J. Agardh 1847 (1, 9, 11, 12, 14, 16, 17, 18)
  Hypnea spinella (C. Agardh) Kützing 1847 (1, 5, 8, 11, 13, 14, 15, 16, 17, 18, 19)
  Hypnea valentiae (Turner) Montagne 1841 (17, 18) **
  Familia Solieriaceae
  Solieria filiformis (Kützing) P.W.Gabrielson 1985 (5, 7) ** +
  Wurdemannia miniata (Duby) Feldmann et Hamel 1934 (14) *
  Familia Phyllophoraceae
  Ahnfeltiopsis concinna (J.Agardh) Silva et De Cew 1992 (5, 11, 12, 13, 14, 16, 18)
  = Ahnfeltia concinna J. Agardh 1847
  Ahnfeltiopsis serenei (E.Y. Dawson) Masuda 1993 (11) *
  Besa leptophylla (J.Agardh) M.S.Calderon et K.A.Miller in Calderon et al 2016 (1, 2, 4, 6, 16)
  =Gymnogongrus leptophyllus J. Agardh 1876
  Gymnogongrus crustiformis E.Y. Dawson 1961 (16) *
  Gymnogongrus johnstonii (Setchell et N.L. Gardner) E.Y. Dawson 1954 (4, 9, 13, 14, 17, 18)
  Gymnogongrus martinensis Setchell et N.L.Gardner 1937 (11) *
  Gymnogongrus platyphyllus N.L. Gardner 1927 (16) *
 ORDEN GRACILARIALES
  Familia Gracilariaceae
  Gracilaria crispata Setchell et N.L. Gardner 1924 (2, 6, 17, 18)
  Gracilaria pachydermatica Setchell et N.L.Gardner 1924 (1) *
  Gracilaria ramisecunda E.Y.Dawson 1949 (11) *
  Gracilaria rubrimembra E.Y.Dawson 1949 (11) *
  Gracilaria spinigera E.Y. Dawson 1949 (11, 13, 16)
  Gracilaria symmetrica E.Y. Dawson 1949 (11) *
  Gracilaria tepocensis (E.Y. Dawson) E.Y. Dawson 1944 (6, 13) **
  Gracilaria textorii (Suringar) De Toni 1876 (13, 16, 17)
  Gracilariopsis costaricensis E.Y. Dawson 1964 (14) *
 ORDEN HILDENBRANDIALES
  Familia Hildenbrandiaceae
  Hildenbrandia dawsonii (Ardré) Hollenberg 1971 (6) *
  Hildenbrandia rubra (Sommerfelt) Meneghini 1841 (1, 11) **
  =Hildenbrandia prototypus Nardo 1834
 ORDEN NEMALIALES
  Familia Galaxauraceae
  Tricleocarpa cylindrica (Ellis et Solander) Huisman et Borowitzka 1990 (13) *
  Familia Liagoraceae
  Dermonema virens (J.Agardh) Pedroche et Ávila Ortíz 1996 (11, 13, 14, 15, 17, 18)
  = Dermonema frappierii (Montagne et Millardet) Borgesen 1942 Izziella orientalis (J.Agardh) Huisman et Schils 2002 (15) *
  Familia Scinaiaceae
  Scinaia confusa (Setchell) Huisman 1985 (18) *
  =Gloiophloea confusa (Setchell) Levring 1914
CLASE STYLONEMATOPHYCEAE
 ORDEN STYLONEMATALES
  Familia Stylonemataceae
  Stylonema alsidii (Zanardini) Drew 1956 (1, 11, 19)
 ORDEN RHODYMENIALES
  Familia Champiaceae
  Champia parvula (C. Agardh) Harvey 1853 (14, 15, 17, 18)
  Familia Lomentariaceae
  Ceratodictyon tenue (Setchell et N.L.Gardner) J.N.Norris 2014 (11) *
  = Gelidiopsis tenuis Setchell et N.L.Gardner 1924
  Ceratodictyon variabile (J.Agardh) R.E.Norris 1987 (1, 11, 13, 14, 15, 18)
  >=Gelidiopsis variabilis (J. Agardh) Schmitz 1895
  Familia Rhodymeniaceae
  Botryocladia uvarioides E.Y. Dawson 1944 (1) *
  Halichrysis irregularis (Kützing) A. Millar 2005 (11) *
  Rhodymenia californica Kylin 1931 (16) *

Three divisions of macroalgae were recorded, Rhodophyta (red algae), Ochrophyta (brown algae) and Chlorophyta (green algae); the red algae had the highest number of all taxonomic levels from Class to species (Table 2).

Table 2.

Number of species, genera, families, orders and classes of the three divisions of macroalgae recorded in the Jalisco coast.


Division Species Genera Families Orders Classes
Rhodophyta 167 72 37 17 4
Ochrophyta 52 22 14 7 1
Chlorophyta 44 17 10 5 1

The distribution of the species was very variable along the coast of Jalisco. The largest number of records was found in Puerto Vallarta (84) in the North, Chamela (104) and, Careyes (71) in the center, and, Tenacatita (86) and, El Tamarindo (74) in the South (Table 3). However, the areas with lower richness (14-27 species), are also from Chimo to Tehuamixtle in the North, and also from Chalacatepec to Isla Cocinas (except for Chamela) in the center. The southern sites had intermediate (54-62) to high (71-86) species richness.

Two species, Jania tenella and Chaetomorpha antennina had the widest distribution along the coast of Jalisco; these species were recorded in 17 sites, followed by Sargasum liebmanii and, Centroceras clavulatum (16 sites). Fifteen species were present along the coastline (10 to 15 sites): Ulva lactuca, Halimeda discoidea, Chnoospora minima, Padina caulescens, Padina gymnospora, Sargassum howellii, Hypnea spinella, Tayloriella dictyurus, Gelidium pusillum, Gayliella flaccida, Gelidium sclerophyllum, Amphiroa beauvoisii, Amphiroa dimorpha, Jania adhaerens and, Jania pacifica. However, the macroalgae of Jalisco are characterized by their restricted distribution due to ecological rarity, these include 101 species (38 %) of 38 different families recorded in only one site (unique species), and, 56 species (21 %) from 28 families recorded in two sites (duplicate species). The species Ectocarpus siliculosus, Cladophoropsis sundanensis and, Phyllodictyon anastomosans, among others, have been recognized as annual algae according to their life cycle, this means that they frequently live only during the most favorable season of the year (Mate-Cid & Mendoza-González, 2012). This may one of the reasons why these species has a low incidence (ecological rarity) in the coast of Jalisco.

Taxonomic Distinctness

The values of the average taxonomic distinctness (∆+) of all sites fell within the probability channel (e.g. within the confidence intervals of 95 %, p>0.05), with the exception of Bahía Chamela (Figure 3). These sites had a greater contribution to the mean taxonomic diversity of macroalgae of the coast of Jalisco. The sites that contributed the most were Cuastecomates, La Manzanilla, El Tamarindo and, Tenacatita. On the other hand, values of the variation of taxonomic distinctness (Λ+) showed that most sites are significantly representative of the macroalgae assemblage, only four sites, Punta Pérula, Chimo, Yelapa and, El Tamarindo, fell out of the 95 % probability channel. This indicates that most of the sites along the coast of Jalisco, have a good representation of the different taxonomic categories of macroalgae (Figure 3; Table 3).


[Figure ID: f3] Figure 3.

Average taxonomic distinctness ( +) and the variation of the taxonomic distinctness (Λ +) of the macroalgae from the coast of Jalisco, Mexico. The solid line shows the 95% confidence intervals and the dashed line the values of + & Λ +. The statistical significance of + and Λ + was assessed using 1,000 permutations. See Table 3 for symbology.


Table 3.

Number of macroalgae taxa recorded in 19 sites of the coast of Jalisco, Mexico. sites are arranged from north to south.


Site / Taxa Species Genera Families Order Classes Division
1. Puerto Vallarta 84 45 34 21 4 3
2. Yelapa 39 29 20 16 3 3
3. Chimo 15 11 9 9 3 3
4. Bahía Corrales 26 16 14 11 3 3
5. Playitas 14 11 10 8 4 3
6. Cabo Corrientes 27 19 17 13 3 3
7. Mayto 14 12 11 10 3 3
8. Tehuamixtle 21 13 11 10 3 3
9. Chalacatepec 18 11 10 7 3 3
10. Punta Pérula 16 10 8 8 3 3
11. Chamela 104 53 40 27 5 3
12. Isla Cocinas 26 18 15 11 3 3
13. Careyes 71 39 30 21 4 3
14. Tenacatita 86 57 39 23 5 3
15. La Manzanilla 62 41 32 21 4 3
16. El Tamarindo 74 36 22 15 4 3
17. Cuastecomates 54 32 23 19 4 3
18. Melaque 55 32 26 19 4 3
19. Barra Navidad 44 26 20 16 4 3

New records of species

The exploration of new sites as well as the increase of sampling effort at sites with single or very few references, allowed to identify two new records for the coast of Jalisco, Solieria filiformis (Kützing) of the Family Solieriaceae, collected from October to May in Playitas and Mayto; and, Piropya raulaguilarii, a unique species of the Family Bangiaceae, collected only in April in Playitas. Both macroalgae were found in the low intertidal of rocky beaches.

Discussion

The completion of an updated inventory of the phycoflora from the coast of Jalisco, integrated from different sources of information, allowed to increase the number of species to 263; this is more than 100 % compared to the last inventory made by Pedroche et al. (2005, 2008), who reported 152 species for the coast of Jalisco. It is important to point out that the most recent inventory was made by Mendoza-González et al. (2011), who recorded the same number of species (152) only in the southern coast of Jalisco. Although many records were found in several specific reports and herbarium, any of them represent true compilations or complete inventories for the entire coast of Jalisco. However, all these were taken into account as part of the documented phycoflora of this region and included in our inventory. The only two new records reported here were found during our own samplings performed in the coast of Jalisco. It is also important to note that with this inventory the species richness from the coast of Jalisco is greater than that found in other adjacent areas, such as the coast of Nayarit with 184 species (Mateo-Cid & Mendoza-González, 1992; Serviere et al., 1993), located in the northern limit of the Mexican tropical Pacific; and the coast of Colima with 125 (Mateo-Cid & Mendoza- González 1991), located towards the south of Jalisco; this indicates the great contribution of the coast of Jalisco to the biodiversity of macroalgae. The species of Jalisco represents 60 % of the total number of species (444) compiled by Pedroche & Sentíes (2003) for the Tropical Mexican Pacific. Finally, comparisons with other recent inventories from the Gulf of California (63 in Norris, 2010; and 124 in Mora-Valdez & Riosmena-Rodriguez, 2016), suggest that the inventory of 263 species here presented has a good representation of the diversity of algae of the Mexican Pacific.

The Rhodophyta dominated the taxonomic diversity of all macroalgae recorded in Jalisco, with the highest number of species (167) and all taxa levels from Class to Genus. These algae grow successfully as epizoic, epiphytic or epilithic on the surface of animals, other algae or rocks with a higher coverage and diversity in tropical marine environments (Mendoza-González et al., 2011). Other studies performed in the Tropical Mexican Pacific have recorded a lower diversity of red algae: Mateo-Cid & Mendoza-González (1992) reported 69 species for the southern coast of Nayarit; Mateo-Cid & Mendoza- González (2002), 93 species for the coast of Guerrero; Mateo-Cid & Mendoza- González (2003), 87 species from Oaxaca; and Dreckmann et al. (2006), only 29 species from Chiapas.

The analysis of the distribution of the macroalgae along the northern, center and southern coast of Jalisco showed some areas with different taxonomic diversity. The sites with higher number of species are protected rocky beaches within bays which are semi-protected from waves and currents, with small barriers or patches of coral reefs. The structural complexity of the rocky and coralline substrate is a key factor for de recruitment and growth of the algal populations. The crevices, hollows and other irregularities of the hard substrate create microenvironments for the settlement of propagules and provide protection to depredation and physical perturbation (Lüning, 1990; Dudley & D’Antonio, 1991). On the other hand, an extensive area of low species richness of macroalgae located in the northern part from Chimo to Tehuamixtle, is characterized by long sandy beaches intercalated by few rocky beaches. Similarly, some sites of the central zone with few species have few suitable places (for example, rocky substrates) for the development of the phycoflora.

Many species of the subtidal environment are found in mixed substrates of rocks, skeletons of dead coral and sand, mainly in shallow areas (<20 m). Although the distribution of the macroalgae showed a greater abundance and number of species in locations with rocky reefs and scattered coral colonies, there were also macroalgae associated with other substrates, such as boulders, pebbles, gravel and sand. For example, some species of the families Cladophoraceae (Cladophora spp), Caulerpaceae (Caulerpa chemnitzia, C. sertularioides) and Halimedaceae (Halimeda discoidea) were common on sandy substrates and gravel. The rocky and coral reefs have the highest biodiversity of invertebrates (scleractina, gorgonians, polychaete worms, echinoderms, molluscs) and fish, with a significant presence of associated macroalgae.

In the coast of Jalisco, there are two protected marine areas: “Chamela-Cuixmala” and “Islas de la Bahía de Chamela”, both in the central region, where four sites of the present study are included (Punta Pérula, Chamela, Isla Cocinas and Careyes). Both areas have the main objective of preserving their biodiversity and ecological balance (CONANP, 2008), and they offer benefits to the inhabitants of the region, generating activities such as fishing and recreation, including diving, which are relevant activities due to their natural scenic beauty.

The historical records of the different groups of organisms are difficult to interpret when they are analyzed in terms of abundance, since they are affected by factors such as the sampling method and the sampling effort; sample size, records of environmental and ecological variables (Clarke & Warwick, 1998, Leonard et al., 2006, Ceshia et al., 2007). The biodiversity measured with the taxonomic distinctness (Δ + and Λ +), has been successfully used in other marine taxonomic groups with information based on the presence/absence of Echinodermata species (: (Ríos-Jara, et al. 2013);); bivalve molluscs (Esqueda-González, et al., 2014); macroalgae, macroinvertebrates and fish (García et al., 2014). The records on the specific richness of macroalgae obtained from the different sources used in our study allowed the identification of four sites that contributed the most to the taxonomic diversity of the marine algae of Jalisco (Figure 3); they were within the probability channel of 95 % of the average taxonomic distinction (Δ +). These four sites (Cuastecomates, La Manzanilla, El Tamarindo and Tenacatita) are located in the southern coastal area and have been historically documented as those with the greatest contribution to the diversity of macroalgae in the region.

The high number of unique species recorded contributed to a great extent, to the phycofloristic mosaic observed in the present study, representing 38% of the total. A characteristic of this group of unique species is that most of them are filamentous organisms, less visible and with a reduced habit size, such as the organisms of Orders Ceramiales, Rhodymeniales, Ectocarpales and Cladophorales (Table 1). These attributes, along with the characteristics of each species, could have been the reason why they were recorded only once. A significant number of these species have been reported as epiphytes of other algae such as Erythrotrichia carnea, Bangia fuscopurpurea, Acrochaetium hancockii, Colaconema daviesii, Asteronema breviarticulatum; therefore, their observation and record could have been occasional and conditioned to the collection and careful observation of the host algae, which in most cases were foliar algae such as Sargassum, or flabelated such as Padina.

It is important to note that the sites in which the greatest numbers of unique species were recorded in the southern coast; historically, this is the region with the largest number of phycological studies. This could be an important factor to increase the records of unique species; since this was not observed in the sites of the northern coast which have been recently explored (Tehuamixtle, Mayto, Playitas). However, it is precisely in this region where two new records for the phycoflora of Jalisco were found: Solieria filiformis and Piropya raulaguilarii, which shows that this must be a high priority region for future studies on marine flora and fauna.

The present study demonstrates that the phycoflora of the coast of Jalisco is diverse and it also contains characteristic elements of the Mexican tropical Pacific. However, more studies are needed for a better understanding of the structure of the macroalgae assemblages of Jalisco, with estimations of coverage and, finally, the biomass of species with a potential exploitation. The study of the different sites along the coast of Jalisco was of great relevance to integrate a more complete taxonomic inventory of the macroalgae, which is essential to identify strategies for the conservation and use of these ecosystems. At this time, this has great relevance because there are several proposals of tourism development for the coast of Jalisco. Finally, the exploitation of macroalgae with different purposes (for example, food, pharmacology) is under development in Mexico, with the species of shallow intertidal and subtidal environments being the main potential resources due to their accessibility and abundance (Carrillo-Domínguez et al., 2002; Hernández-Carmona et al., 2012). In the Mexican tropical Pacific, some common species, for example, Sargassum liebmannii (Ochrophyta), Caulerpa sertularioides (Chlorophyta) and Spyridia filamentosa (Rhodophyta), have shown good biological activity and potential use in food and pharmacology (Radulovich et al., 2015; Garcia-Granados et al., 2016; Landa-Cansigno et al., 2017). However, any proposal for its use must consider evaluations of exploitable biomass to guarantee the sustainable use of these resources.

Conclusions

Based on the updated inventory integrated from new and previous information, a total of 263 species of macroalgae from 19 sites of the coast of Jalisco, Mexico, including new records of species, were recognized. Macroalgae of Jalisco are characterized by a high ecological rarity (incidence), with 38 % of unique species (recorded only in one site) and 21 % of duplicated species (in two site). This inventory highlights the contribution of this region to the diversity of macroalgae of the Mexican tropical Pacific with at least 60 % of the species commonly found in subtidal, intertidal and shallow environments. The Jalisco coast is one of the regions with the highest species richness of the Eastern tropical Pacific (Pedroche et al., 2005, 2008), including Central America (Pedroche & Sentíes, 2003a, Pedroche & Sentíes, 2003b, Pedroche et al., 2003).


1.

fn1Cite this paper: Enciso-Padilla, I., Ríos-Jara, E., Esqueda-González, M.C. (2019). Inventory and taxonomic diversity of macroalgae from the coast of Jalisco, México. Revista Bio Ciencias 6, e656. doi: https://doi.org/10.15741/revbio.06.e656

Acknowledgments

The first author thanks the Consejo Nacional de Ciencia y Tecnología (CONACyT) and the University of Guadalajara for the financial support provided during the current research, also thanks all the students of Biology (Universidad de Guadalajara) who contributed during the last 15 years with collection of samples and the creation of the Phycology Herbarium of the Departamento de Ecología of the Universidad de Guadalajara.

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Revista Bio Ciencias, Año 13, vol. 9,  Enero 2022. Sistema de Publicación Continua editada por la Universidad Autónoma de Nayarit. Ciudad de la Cultura “Amado Nervo”,  Col. Centro,  C.P.: 63000, Tepic, Nayarit, México. Teléfono: (01) 311 211 8800, ext. 8922. E-mail: revistabiociencias@gmail.com, revistabiociencias@yahoo.com.mx, http://revistabiociencias.uan.mx. Editor responsable: Dra. Karina J. G. Díaz Resendiz. No. de Reserva de derechos al uso exclusivo 04-2010-101509412600-203, ISSN 2007-3380, ambos otorgados por el Instituto Nacional de Derechos de Autor. Responsable de la última actualización de este número Dra. Karina J. G. Díaz Resendiz Secretaria de Investigación y Posgrado, edificio Centro Multidisciplinario de Investigación Científica (CEMIC) 03 de la Universidad Autónoma de Nayarit. La opinión expresada en los artículos firmados es responsabilidad del autor. Se autoriza la reproducción total o parcial de los contenidos e imágenes, siempre y cuando se cite la fuente y no sea con fines de lucro.

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