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Seagrasses of Costa Rica: from the mighty Caribbean to the dynamic meadows of the Eastern Tropical Pacific/Pastos marinos de Costa Rica: del majestuoso Caribe a las praderas dinamicas del Pacifico Este Tropical.

Costa Rica is a small country in Central America with 51 000 [km.sup.2] total land area (Cortes & Wehrtmann, 2009). The general conditions for seagrass meadows vary greatly between the Caribbean and Pacific coasts, despite both coasts being only 120-285 km apart. The Pacific coast is very irregular, with a length of 1 254 km, has a ~3 m tidal range, and hosts a wide variety of habitats (Cortes, 2016a). Meanwhile, the Caribbean coast a length of 212 km with more regular geomorphology, and a much smaller tidal range of approximately 0.5 m (Cortes, 2016b). Costa Rica hosts 3.5 % of the worlds' marine biodiversity (Wehrtmann, Cortes, & Echeverria-Saenz, 2009), which includes the species associated with seagrass meadows on both coasts (Cortes & Salas, 2009).

Seagrass meadows are important marine habitats that are found along coastlines around the world (Hemminga & Duarte, 2000). Seagrasses are an ecological group of flowering plants that thrive in saline to brackish waters, and there are between 60 and 72 species of seagrass worldwide (Den Hartog & Kuo, 2006; Short et al., 2011). These plants are the foundation species of diverse coastal ecosystems in shallow waters, with a multitude of associated organisms, providing a plethora of ecosystem services, such as nursery habitat, nutrient recycling, carbon sequestration, sediment deposition, and coastal protection (Nordlund, Koch, Barbier, & Creed, 2016).

Seagrass meadows are declining worldwide (Waycott et al., 2009), including the Caribbean region (Van Tussenbroek et al., 2014). This decline has been strongly linked to anthropogenic activity, mainly nutrient enrichment, increased sedimentation, and light reduction (Orth, Carruthers, Dennison, Duarte, Fourqurean, Heck, et al., 2006; Waycott et al., 2009). Conservation and management initiatives that aim to preserve the functions and presence of seagrass meadows need to have a strong basis on scientific knowledge, including the location and spatial area of seagrass meadows. Presence of seagrass meadows has been acknowledged for many years in Costa Rica; particularly in the Caribbean (Cortes & Salas, 2009). Here, we present recent additions to the reported species and locations of seagrasses for Costa Rica. We analyze previous research efforts on Costa Rican seagrass meadows, and identify key areas for further study.

MATERIALS AND METHODS

Information for this study was gathered by searching google scholar for "Costa Rica + seagrass", "Eastern Tropical Pacific + seagrass", and in Spanish "pastos marinos + Costa Rica", "fanerogamas + Costa Rica". Local library databases were also searched for these terms, and for university thesis, as was the Revista de Biologia Tropical online search engine (available at: revistas.ucr.ac.cr). References related to seagrass or seagrass habitat research in Costa Rican coasts were selected for further analyses.

Multiple herbarium collections were searched for Costa Rican seagrass specimens using the different genus and species as keywords. Herbarium specimens from the Herbario Nacional de Costa Rica (CR) and Herbario de la Escuela de Biologia, Universidad de Costa Rica (USJ) were photographed and analysed in person. Specimens from the Herbario Nacional de Mexico, Universidad Nacional Autonoma de Mexico (MEXU), were searched digitally online and detailed high resolution photographs of each specimen were assessed (datosabiertos. unam.mx). The information from the specimens at the Missouri Botanical Garden Herbarium (MO) was searched on their online Tropicos database (tropicos.org). Information was also included in this analysis from new herbarium samples collected in recent years that have been recently deposited at CR, USJ and the State Herbarium of South Australia (AD) and have yet to be assigned herbarium identification numbers.

Informal interviews were conducted with local contacts that responded positively when asked if they had ever seen seagrasses on the Pacific coast of Costa Rica. Interviewees included researchers, field technicians and research assistants at the Centro de Investigacion en Ciencias del Mar y Limnologia (CIMAR) at the Universidad de Costa Rica (UCR), students at the Escuela de Biologia, UCR, Non-Government Organization workers, conservation area government workers, divers and local coastal residents. During these interviews, they were asked to identify the species from a plate with pictures of various local seagrass species or to describe details of the spotted plants including the location and date of the sighting. A map was used to identify key locations whenever possible. Maps with identified seagrass locations or names of locations provided during interviews and herbarium samples with only location name were then used to extract approximate coordinates for each meadow using Google Earth (7.1.7.2606). Additional localities have been observed directly by the authors.

RESULTS

Seven seagrass species are reported for Costa Rica. Six species are reported for the Caribbean, and four species for the Pacific (see taxonomic considerations for some seagrass species in the discussion). Three species are only present in the Caribbean, and one species is only reported for the Pacific coast of Costa Rica. Three species occur along both coasts (Table 1). Most species are classified as Least Concern in the IUCN Red List; except for Halophila baillonis which is listed as Vulnerable, and Halodule beaudettei is classified as Data Deficient, mainly due to uncertainties concerning its taxonomical status (Table 1).

Seagrasses occur at 31 locations, of which 16 are reported for the first time in this study (Fig. 1, Table 2). These new records consist of herbarium samples collected previously (four locations), personal communications during the informal interviews in this study (seven locations), and recent new sightings by the authors (five locations) (Table 2). Of the locations where seagrasses have been found previously (Table 2), we can confidently say, from recent observations, that seagrasses are present on the Pacific coast at Bahia Jobo, Bahia Culebra, Bahia Potrero, Isla Chora, Punta Leona, Parque Nacional Marino Ballena, and most extensively at Golfo Dulce. In the Caribbean, seagrasses have been observed recently at Isla Uvita, Parque Nacional Cahuita (hereafter PNC), Puerto Viejo, and Refugio Nacional de Vida Silvestre Gandoca-Manzanillo (hereafter RGM). Based on the current information, total seagrass area for Costa Rica is estimated as 1 328 384 [m.sup.2] (132.8 ha), most on the Pacific coast.

Seagrasses have been found at ten locations in the Caribbean, with three locations reported in this study for the first time (Table 2). Seagrass presence is limited to sheltered areas near the port city of Limon and in coral reef lagoons further south at PNC and the RGM. Seagrasses on the Caribbean coast have been found up to 8 m deep. The most studied location and with the highest number of species reported for the Caribbean coast is PNC, with four species of seagrass and a total of 37 herbarium samples, dating back to 1975. The oldest samples of seagrasses found were Thalassia testudinum at Puerto Limon from 1963 and of Ruppia maritima at Moin from 1965 (Table 1). Seagrass area for the Caribbean coast is estimated as 360 400 [m.sup.2] (36.0 ha).

Seagrasses have been found at 21 locations on the Pacific coast, 13 of them reported here for the first time (Table 2). On the Pacific coast of Costa Rica, the locations with seagrasses consist mainly of sheltered bays in the northern Pacific coast (eight locations), central Pacific coast (four locations), and the southern Pacific coast (nine locations) (Fig. 1). The most extensive meadow currently present in the Pacific coast is located at Playa Colibri in Golfo Dulce, with an area up to 900 000 [m.sup.2] and three seagrass species (Table 2). At Bahia Culebra, only isolated shoots of R. maritima have been sighted in the last ten years. Seagrasses on the Pacific coast have been found up to 8 m deep. The oldest herbarium samples refer to R. maritima collected in 1967 from Playas del Coco (Table 2). Seagrass area for the Pacific coast is estimated as 967 984 [m.sup.2] (96.8 ha).

A total of 43 studies were found that included or reported seagrasses in Costa Rica, and the number of studies has been steadily increasing over time (Fig. 2). The majority of the studies were done in meadows along the Caribbean coast, and publications on seagrasses from the Pacific coast only appeared after the year 2000 (Fig. 2). Most studies have been taxonomic reports, followed by some ecological studies and monitoring, with recent focus on associated fauna (Fig. 3). The top three most studied species are T. testudinum, followed by Syringodium filiforme and H. baillonis, and the least studied species is H. beaudettei (Fig. 3).

DISCUSSION

Seven seagrass species have been reported for Costa Rica; however, a number of taxonomic considerations have to be taken into account. Two key aspects are potential misidentifications of seagrass species and the taxonomic standing of some species (Short et al., 2011). The current species name of the clover grass is H. baillonis, though there is discrepancy among specialists whether Halophila baillonii should be the correct species used (see Kuo & Wilson, 2008 for a detailed discussion on the taxonomic standing of the clover grass). In the past, the species H. baillonis has been misidentified with Halophila decipiens (Kuo & Wilson, 2008). The only record of H. baillonis in the Caribbean of Costa Rica was by Dawson (1962), which may have been H. decipiens, illustrations were not included. In contrast, H. decipiens has been correctly reported for the Caribbean of Costa Rica, with many herbarium samples, and it is currently present at Isla Uvita. Nonetheless, H. baillonis has recently been reported in the Caribbean Sea at Belize (Short, Fernandez, Vernon, & Gaeckle, 2006), Honduras (Caviedes-Sanchez & Carrasco, 2016), and in the Atlantic coast of Brazil (Barros, Costa, & Rocha-Barreira, 2014; Magalhaes, Borges, & Pitanga, 2015); thus, the record by Dawson (1962) may have been correct, and it is also possible that this inconspicuous species may still be found along the Caribbean coasts of Costa Rica.

Another case of misidentification was Halophila engelmannii, reported for the Pacific coast of Costa Rica from a single specimen at the herbarium of the Universidad Nacional Autonoma de Mexico (UNAM) (MEXU-1249952) (Crow, 2003b). High definition photographic material for this specimen was carefully studied by us and we consider the correct species to be H. baillonis. The specimen has four short leaves per foliar shoot with oval apices and defined petioles, similar to H. baillonis (Kuo & Den Hartog, 2001; Van Tussenbroek, Santos, Wong, Van Dijk, & Waycott, 2010). In addition, H. engelmanni has not been reported along the Pacific coast elsewhere (Phillips & Menez, 1988; Green & Short, 2003).

The taxonomy of Halodule needs to be clarified. H. beaudettei is considered by some to be a synonym of Halodule wrightii, as it has been argued that leaf-tip morphology is insufficient to define the taxonomic status of the species within this genus (Phillips, 1967). The specimens of H. beaudettei, reported here for the first time for the Pacific coast, present an acute median tooth differing from samples of H. wrightii from the Caribbean coast (Kuo & Den Hartog, 2001; Van Tussenbroek et al., 2010). Genetic analyses on H. wrightii from the Caribbean and H. beaudettei from the Pacific of Costa Rica are currently under way.

Ruppia maritima has been reported for both coasts; however, its verified current presence is extremely limited. In the Caribbean, there is only one herbarium sample from 1965 supporting its presence but there have been no other sightings of this species. On the Pacific coast, there are multiple herbarium samples of this species; yet the only reported meadow disappeared in the mid-1990s; with only some isolated shoots noted in the northern and central Pacific coast in the last 15 years (pers. obs. JCN & pers. com. Eleazar Ruiz). In Mexico, specimen of this species previously identified as R. maritima were considered to belong to a new species Ruppia mexicana (Den Hartog, Van Tussenbroek, Wong, Mercado-Ruaro, & Marquez-Guzman, 2016), and therefore careful examination of specimens from Costa Rica should clarify their taxonomic status. A final cautionary note on species in Costa Rica: the newly introduced Halophila stipulacea in the Caribbean has not been found in Costa Rica. Considering its rapid expansion throughout the Caribbean (Willette et al., 2014; Van Tussenbroek et al., 2016) it is most likely just a matter of time before this species will be found on the Caribbean coast of Costa Rica.

Different seagrass species occupy the Pacific and Caribbean coasts of Costa Rica with distinct habits in each coast. In the Eastern Tropical Pacific (ETP), seagrasses belong to the genera Halophila, Halodule and Ruppia (Cortes, 2001; Green & Short, 2003; Short, Carruthers, Dennison, & Waycott, 2007). These seagrasses are mainly small pioneer or opportunistic species, which can also be ephemeral (Orth et al., 2006; Kendrick et al., 2012). In the Caribbean, seagrass meadows are mostly dominated by the large species T. testudinum, the faster growing S. filiforme, and opportunistic ephemeral H. wrightii (Orth et al., 2006; Van Tussenbroek et al., 2010; Kendrick et al., 2012). In the Pacific, seagrasses develop in deeper or more turbid waters than in the Caribbean, where they are found in shallow coastal or reef lagoons. Even so, both coasts of Costa Rica belong to the Tropical Atlantic seagrass bioregion (Short et al., 2007).

The number of seagrass locations is higher in the Pacific than the Caribbean; yet Caribbean locations are mostly large meadows (Cortes, Soto, & Jimenez, 1994; Cortes & Jimenez, 2003), while in the Pacific the largest meadow is found in the southern Pacific coast with many smaller patches. Seagrass presence in the Caribbean is restricted to a sheltered island habitat in the central coast and coral reef lagoons in the southern areas of the Caribbean coast (Cortes & Guzman, 1985; Cortes, 2016b). The northern Caribbean coast of Costa Rica is rectilinear, has high wave energy and precipitation leading to higher river effluent than the southern Caribbean, likely hindering seagrass presence there (Cortes, 2016b). Given limited research, current estimates of seagrass coverage in Costa Rica are considered to be an underestimation of actual seagrass area.

On the Pacific coast of Costa Rica, following seagrass loss in the northern Pacific coast in the mid-1990s (Cortes, 2001), there were no other known extant seagrass meadows, until 2009 and 2010 when seagrasses were reported in the southern Pacific coast (Bessesen & Saborio-R., 2012; Samper-Villarreal, Bourg, Sibaja-Cordero, & Cortes, 2014; Sarmento de Carvalho, 2013; Sarmento, Chaves, Retamosa, Ruepert, Jimenez, & Blanco, 2016). In this area, seagrasses have been continuously sighted since, and here we report an additional five new locations. The new sightings may indicate recent colonization by seagrasses in this sheltered fjord-like estuary; however, residents recall the presence of seagrasses at some of these sites for decades.

Seagrass research in Costa Rica has been limited, but has been steadily increasing. Forty-three studies on seagrasses were found, most of which were either taxonomic reports or site descriptions. Presence of seagrasses has also been reported in studies on other organisms such as marine turtles (Sarmento de Carvalho, 2013; Chacon-Chaverri, Martinez-Cascante, Rojas, & Fonseca, 2015a, 2015b; Sarmento et al., 2016), corals (Fernandez & Alvarado, 2004), and fish (Myers, Wagner, & Vaughan, 2011). Seagrasses in Costa Rica are subtidal, and are only exposed during the maximum low tides. The absence of large exposed intertidal seagrass beds which can be easily accessed at low tide, common in the Indo-Pacific or temperate regions (Moore & Short, 2007; Van Tussenbroek et al., 2007), may be an important reason for limited research effort on Pacific seagrasses. The search for seagrass populations in the Pacific is more challenging than in the Caribbean. Pacific seagrass species are smaller, more ephemeral, and occur in deeper or more turbid waters, with tidal ranges 3 m or higher (Lizano, 2006); while Caribbean meadow species are larger and grow in clearer and calmer waters, with a tidal range of 0.3-0.5 m (Cortes, 1998; Lizano, 2006). In the Caribbean, there has been one continuous monitoring program in place since 1999, the Caribbean Coastal Marine Productivity Program known as CARICOMP (Fonseca, Nielsen, & Cortes, 2007; Cortes et al., 2010); which accounts for all the monitoring publications focusing on plant biomass, productivity and turnover rates. Studies on grazing and non-charismatic organisms associated with seagrass meadows are very limited, with only one study focused on macroalgal epiphytes in the Caribbean (Samper-Villarreal, Bernecker, & Wehrtmann, 2008), and one study mentioning limited grazing and associated macrofauna in the southern Pacific (Samper-Villarreal et al., 2014). Seagrasses in the southern Pacific coast are known as a resting, breeding and foraging area for green sea turtles and they are important locations along migratory routes (Bessesen & Saborio-R., 2012; Sarmento de Carvalho, 2013; Chacon-Chaverri et al., 2015a, 2015b). Given the Vulnerable status of H. baillonis according to the IUCN red list (Short et al., 2011) and the importance of these meadows for sea turtles, conservation and management initiatives of these meadows is warranted.

Seagrasses in Costa Rica are subjected to multiple disturbances, and further research on seagrass meadows in Costa Rica is urgently needed to establish management and conversation initiatives. Loss of seagrasses in the northern Pacific coast followed a severe storm resulting in light deprivation from increased and prolonged water turbidity, and uprooting of the plants (Cortes, 2001). These seagrass meadows have shown no signs of recovery to date. In 1991, the Limon Earthquake caused coastal uplifting of the Caribbean coast (Cortes, Soto, Jimenez, & Astorga, 1992; Cortes et al., 1994; Cortes & Jimenez, 2003), which resulted in the subaerial exposure and subsequent die-off of more than 40 000 [m.sup.2] of seagrass (Cortes et al., 1992; Cortes et al., 1994). Another impact was whitening and shedding of T. testudinum leaves in 2005 following heavy storms at Manzanillo; but leaf regrowth shortly resumed afterwards and no permanent damage to the meadows was reported (Krupp, 2006; Krupp et al., 2009). The long-term monitoring program in Caribbean meadows has revealed a decline over time, most likely due to increased turbidity (Van Tussenbroek et al., 2014). In November 2016 Level 4 hurricane Otto passed through the country, with high wave and wind energy and increased precipitation. Other threats are local, such as increased turbidity from land alteration, increased nutrient runoff, and anchoring on seagrass meadows (Cortes & Risk, 1985; Orth et al., 2006; Cortes et al., 2010). Further research, monitoring, management and conservation strategies are needed for Costa Rican seagrasses, with a stronger focus needed on research looking at associated fauna and flora, and incorporating ecosystem resilience and services.

ACKNOWLEDGEMENTS

This research was funded by the Vicerrectoria de Investigacion at the University of Costa Rica (project 808-B6-170). International academic exchange linked to this work was supported by the Oficina de Asuntos Internacionales y Cooperacion Externa (OAICE), Universidad de Costa Rica, and Centro de Estudios Mexicanos, Universidad de Costa Rica-Universidad Nacional Autonoma de Mexico. We would like to thank all those that contributed to seagrass research and herbarium maintenance in previous years. Our thanks to the many individuals who shared their knowledge on seagrass sightings with us.

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Risk, M., Murillo, M., & Cortes, J. (1980). Observaciones biologicas preliminares sobre el arrecife coralino en el Parque Nacional Cahuita, Costa Rica. Revista de Biologia Tropical, 28, 361-382.

Samper-Villarreal, J., Bernecker, A., & Wehrtmann, I. S. (2008). Inventory of macroalgal epiphytes on the seagrass Thalassia testudinum (Hydrocharitaceae) in Parque Nacional Cahuita, Caribbean coast of Costa Rica. Revista de Biologia Tropical, 56(4), 163-174.

Samper-Villarreal, J., Bourg, A., Sibaja-Cordero, J. A., & Cortes, J. (2014). Presence of a Halophila baillonii Asch.(Hydrocharitaceae) seagrass meadow and associated macrofauna on the Pacific Coast of Costa Rica. Pacific Science, 68(3), 435-444.

Sarmento de Carvalho, M. M. (2013). Analisis descriptivo de las condiciones ambientales y estado de salud de la tortuga negra (Chelonia mydas agassizii) en un sitio de Forrajeo en el Golfo Dulce, Costa Rica (Master's thesis). Heredia, Costa Rica: Universidad Nacional.

Sarmento, M., Chaves, A., Retamosa, M., Ruepert, C., Jimenez, R., & Blanco, K. (2016). Analisis descriptivo de las condiciones de conservacion de un sitio de forrajeo de la tortuga verde del Pacifico (Chelonia mydas agassizii, Testudines: Chelonioidea) en el Golfo Dulce, Costa Rica; se toma como base el estado de salud de la tortuga y el hallazgo de plaguicidas en el ambiente. Revista Ciencias Veterinarias, 33(2), 81-96.

Short, F. T., Fernandez, E., Vernon, A., & Gaeckle, J. (2006). Occurrence of Halophila baillonii meadows in Belize, Central America. Aquatic Botany, 85(3), 249-251.

Short, F. T., Carruthers, T., Dennison, W., & Waycott, M. (2007). Global seagrass distribution and diversity: A bioregional model. Journal of Experimental Marine Biology and Ecology, 350(1), 3-20.

Short, F. T., Polidoro, B., Livingstone, S. R., Carpenter, K. E., Bandeira, S., Bujang, J. S., ... Zieman, J. C. (2011). Extinction risk assessment of the world's seagrass species. Biological Conservation, 144(7), 1961-1971.

Van Tussenbroek, B., Vonk, J. A., Stapel, J., Erftemeijer, P. L., Middelburg, J. J., & Zieman, J. C. (2007). The biology of Thalassia: paradigms and recent advances in research. In A. Larkum, R. J. Orth, & C. M. Duarte (Eds.). Seagrasses: Biology, Ecology and Conservation (pp. 409-439). The Netherlands: Springer.

Van Tussenbroek, B., Santos, M. G. B., Wong, J. G. R., Van Dijk, J. K., & Waycott, M. (2010). A Guide to the Tropical Seagrasses of the Western Atlantic. Mexico D.F.: Universidad Nacional Autonoma de Mexico.

Van Tussenbroek, B., Cortes, J., Collin, R., Fonseca, A. C., Gayle, P. M., Guzman, H. M., Jacome, G. E., ... Weil, E. (2014). Caribbean-wide, long-term study of seagrass beds reveals local variations, shifts in community structure and occasional collapse. PloS one, 9(3), e90600.

Van Tussenbroek, B., Van Katwijk, M., Bouma, T., van der Heide, T., Govers, L., & Leuven, R. (2016). Non-native seagrass Halophila stipulacea forms dense mats under eutrophic conditions in the Caribbean. Journal of Sea Research, 115, 1-5.

Waycott, M., Duarte, C. M., Carruthers, T. J. B., Orth, R. J., Dennison, W. C., Olyarnik, S., ... Hughes, A. R. (2009). Accelerating loss of seagrasses across the globe threatens coastal ecosystems. Proceedings of the National Academy of Sciences, 106(30), 12377-12381.

Wehrtmann, I. S., Cortes, J., & Echeverria-Saenz, S. (2009). Marine biodiversity of Costa Rica: perspectives and conclusions. In I. S. Wehrtmann & J. Cortes (Eds.). Marine Biodiversity of Costa Rica, Central America (pp. 521-533). Berlin: Springer + Business Media B.V.

Wellington, G. (1974). The benthic flora of Punta Cahuita: an annotated list of species with new additions to the Costa Rican Atlantic flora. Brenesia, 3, 19-30.

Wetzer, R., & Bruce, N. L. (1999). A new genus and species of sphaeromatid isopod (Crustacea) from Atlantic Costa Rica. Proceedings of the Biological Society of Washington, 112, 368-380.

Willette, D. A., Chalifour, J., Debrot, A. D., Engel, M. S., Miller, J., Oxenford, H. A., ... Vedie, F. (2014). Continued expansion of the trans-Atlantic invasive marine angiosperm Halophila stipulacea in the Eastern Caribbean. Aquatic Botany, 112, 98-102.

Jimena Samper-Villarreal (1), Brigitta I. van Tussenbroek (3) & Jorge Cortes (1,2)

(1.) Centro de Investigacion en Ciencias del Mar y Limnologia (CIMAR), Ciudad de la Investigacion, Universidad de Costa Rica, San Pedro, 11501-2060 San Jose, Costa Rica; jimena.sampervillarreal@ucr.ac.cr, jorge.cortes@ucr.ac.cr

(2.) Escuela de Biologia, Universidad de Costa Rica, San Pedro, 11501-2060 San Jose, Costa Rica.

(3.) Unidad Academica Sistemas Arrecifales-Puerto Morelos, Instituto de Ciencias del Mar y Limnologia, Universidad Nacional Autonoma de Mexico, Mexico; vantuss@cmarl.unam.mx

Received 18-I-2017. Corrected 12-VI-2017. Accepted 03-I-2018.

Caption: Fig. 1. Seagrass presence on the Pacific and Caribbean coasts of Costa Rica, Central America. Numbers refer to individual seagrass locations (see Table 2). Seagrass categories: "Seagrass Loss" refers to a meadow where seagrasses were lost and have not yet recovered. "Recently Present" refers to meadows which have seen at that location within the current decade. "Reported Seagrass" refers for locations for which there are only older herbarium samples or personal communications.

Caption: Fig. 2. Number of publications on Costa Rican seagrasses per coast by time period up to December 2016.

Caption: Fig. 3. Percentage of studies on seagrasses of Costa Rica up to December 2016 by (A) type of study, n = 43; and (B) species, n = 37 (percentage indicates the number of the total publications in which that species was mentioned).
TABLE 1
Seagrass species reported for each coast of Costa Rica. IUCN Red List
and population trends from Short et al. (2011).
Pop = Population Trend Worldwide

Species                 Reports in                  Reports in the
                        the Caribbean               Pacific

Hydrocharitaceae
Thalassia testudinum    Dawson, 1962; Den Hartog,   None
IUCN: Least Concern     1970; Wellington, 1974;
Pop: Stable             Risk, Murillo, & Cortes,
                        1980; Gomez, 1984; Cortes
                        & Guzman, 1985; Cortes &
                        Risk, 1985; Phillips &
                        Menez, 1988; Cortes et
                        al., 1992; Davidse,
                        Sousa, & Chater, 1994;
                        Cortes & Jimenez, 2003;
                        Crow, 2003b; Green &
                        Short, 2003; Cortes &
                        Salas, 2009; Cortes et
                        al., 2010; Krupp, 2006;
                        Krupp, Cortes, & Wolff,
                        2009; Nielsen Munoz,
                        2006, 2007; Paynter,
                        Cortes, & Engels, 2001;
                        Van Tussenbroek et al.,
                        2010; Van Tussenbroek et
                        al., 2014;

Halophila decipiens     Gomez, 1984; Phillips &     None
IUCN: Least Concern     Menez, 1988; Davidse et
Pop: Stable             al., 1994; Crow, 2003b;
                        Green & Short, 2003;
                        Krupp, 2006; Nielsen
                        Munoz, 2006, 2007;
                        Nielsen-Munoz & Cortes,
                        2008; Cortes & Salas,
                        2009; Van Tussenbroek et
                        al., 2010

Halophila baillonis     Dawson, 1962; Wellington,   Gomez, 1984;
IUCN: Vulnerable        1974; Gomez, 1984;          Phillips & Menez,
Pop: Decreasing         Phillips & Menez, 1988;     1988; Cortes, 2001;
                        Crow, 2003b                 Green & Short,
                                                    2003; Krupp, 2006;
                                                    Nielsen Munoz,
                                                    2006, 2007; Cortes
                                                    & Salas, 2009; Van
                                                    Tussenbroek et al.,
                                                    2010 Bessesen &
                                                    Saborio-R., 2012;
                                                    Sarmento de
                                                    Carvalho, 2013;
                                                    Samper-Villarreal
                                                    et al., 2014
Cymodoceaceae
Syringodium filiforme   Dawson, 1962; Wellington,   None
IUCN: Least Concern     1974; Gomez, 1984; Cortes
Pop: Stable             & Guzman, 1985; Phillips
                        & Menez, 1988; Cortes et
                        al., 1992; Davidse et
                        al., 1994; Cortes &
                        Jimenez, 2003; Green &
                        Short, 2003; Hammel,
                        2003; Cortes & Salas,
                        2009; Krupp, 2006;
                        Nielsen Munoz, 2006,
                        2007; Krupp et al., 2009;
                        Van Tussenbroek et al.,
                        2010

Halodule wrightii       Gomez, 1984; Phillips &     Phillips & Menez,
IUCN: Least Concern     Menez, 1988; Cortes et      1988; Green &
Pop: Increasing         al., 1992; Davidse et       Short, 2003; Van
                        al., 1994; Green & Short,   Tussenbroek et al.,
                        2003; Hammel, 2003;         2010
                        Krupp, 2006; Nielsen
                        Munoz, 2006, 2007; Cortes
                        & Salas, 2009; Van
                        Tussenbroek et al., 2010

Halodule beaudettei     None                        Sarmento de
IUCN: Data Deficient                                Carvalho, 2013
Pop: Unkown:

Ruppiaceae
Ruppia maritima         Dawson, 1962; Gomez,        Gomez, 1984;
IUCN: Least Concern     1984; Davidse et al.,       Phillips & Menez,
Pop: Stable             1994; Cortes & Salas,       1988; Davidse et
                        2009; Van Tussenbroek et    al., 1994; Cortes,
                        al., 2010                   2001; Crow, 2003a;
                                                    Nielsen Munoz,
                                                    2006, 2007; Cortes
                                                    & Salas, 2009; Van
                                                    Tussenbroek et al.,
                                                    2010

TABLE 2

Seagrass locations reported for the Caribbean and Pacific coasts of
Costa Rica, with the date sighted, species present, estimated
extension, verification evidence and cited publications.
AD = State Herbarium of South Australia; CR = Herbario Nacional
de Costa Rica; MEXU = Herbario Nacional de Mexico, Universidad
Nacional Autonoma de Mexico; JS = Jimena Samper personal code;
MO = Missouri Botanical Garden; USJ = Herbario de la Escuela de
Biologia, Universidad de Costa Rica. TBA = to be assigned.
nk = Not known

Location    Location                 Species Herbarium
number                               collections
(Fig. 1)

Pacific     Northern

1           Bahia Jobo (1)           Halophila baillonis
2           Playa Naranjo, Parque    Ruppia maritima MO-Grayum #5131;
            Nacional Santa Rosa      CR-119424
3           Bahia Huevo (2)          c.f. Ruppia maritima
4           Bahia Culebra (3)        Ruppia maritima*
                                     Halophila baillonis USJ-005824
5           Playas del Coco          Ruppia maritima USJ-7228
6           Bahia Potrero (4)        Halodule beaudettei CR-TBA
                                     Halophila baillonis CR-TBA
7           Estero de Ostional       Ruppia maritima MEXU-546253
8           Isla Chora, Samara (5)   Halophila baillonis
9           Bahia Curu (2)           Halophila baillonis
            Central
10          Reserva Biologica de     Unkown
            Punta Leona
11          Herradura                Unkown
12          Parque Nacional          Ruppia maritima
            Marino Ballena
            Southern
13          Mouth of Rio Siepre      Halophila baillonis MEXU-1249952
14          Playa Ganado, Bahia      Unkown
            Drake
15          Puerto Jimenez, Golfo    Halodule beaudettei AD-TBA,
            Dulce (6)                CR-TBA, USJ-TBA
                                     Halophila baillonis AD-TBA,
                                     CR-TBA, USJ-TBA
16          Playa Neques, Golfo      c.f. Halophila baillonis
            Dulce (7)
17          Playa Colobri, Golfo     Halodule beaudettei USJ-101555,
            Dulce (8,9)              CR-TBA, AD-TBA, CR-TBA, USJ-TBA
                                     Halodule wrightii MO-2525084
                                     Halophila baillonis USJ-101556,
                                     CR-TBA, AD-TBA
18          Rincon, Golfo            Halophila baillonis USJ-101557,
            Dulce (10)               CR-TBA
19          Piedras Blancas,         c.f. Halophila baillonis
            Golfo Dulce (7)
20          Playa Animal, Golfo      Halophila baillonis AD-TBA,
            Dulce (8)                CR-TBA, USJ-TBA
21          Golfito, Golfo           Halophila baillonis AD-TBA,
             Dulce (8)                 CR-TBA, USJ-TBA

Caribbean   Southern
22          Moin                     Ruppia maritima
                                     MO-1936778
23          Portete                  Thalassia testudinum
                                     Halophila decipiens USJ-038574,
                                     USJ-038575, USJ-038576
                                     Halophila baillonis
24          Piuta (2)                Thalassia testudinum
25          Isla Uvita (2,8)         Thalassia testudinum
                                     Halophila decipiens USJ-TBA
26          Puerto Limon             Thalassia testudinum CR-66687
27          Cahuita                  Thalassia testudinum MO-A Lot
                                     #1255, MEXU-366392, CR-062284,
                                     CR-274353, CR-TBA, USJ-017357,
                                     USJ-031104, USJ-031105,
                                     USJ-031106, USJ-037843, USJ-
                                     024515, CR-198189, USJ-102076,
                                     USJ-92778, USJ-92779
                                     Syringodium filiforme MO-A Lot
                                     #1258, MEXU-366595, CR-50.020,
                                     CR-138191, CR-167852, USJ-017312,
                                     USJ-037843, USJ-104913,
                                      USJ-104919, USJ-104920,
                                     USJ-104921, USJ-92780
                                     Halodule wrightii MO- 2525084,
                                     MEXU-366577, MEXU-366593, AD-TBA,
                                     USJ-TBA Halophila decipiens
                                     MO-684710, MEXU-366594,
                                     CR-138146, USJ-88569, USJ-85511
28          Puerto Viejo             Thalassia testudinum CR-87408,
                                      CR-257304
29          Cocles                   Thalassia testudinum
30          Manzanillo               Thalassia testudinum CR-273678
                                     Syringodium filiforme CR-273679
31          Punta Mona               Thalassia testudinum CR-93991

Location      Extension         Water           References
number        ([m.sup.2])       column depth
(Fig. 1)

Pacific

1             60 000            0.5-2 m         This study
2             nk                nk              Crow, 2003a
3             nk                1-8 m           This study
4             <100-5 000        < 2 m           Cortes, 2001;
                                                 Crow, 2003b
5             nk                nk              This study
6             nk                6 m             This study

7             nk                0 m             This study
8             ~ 100             3 m             This study
9             nk                1-8 m           This study
10            nk                0-3.5 m         Myers et
                                                 al., 2011
11            nk                nk              Cortes, 2001
12            nk                nk              This study
13            nk                0.01-0.2 m      Cortes, 2001
14            nk                nk              BIOMARCC-SINAC-GIZ,
                                                2012
15            ~ 900              2-3 m          This study
16            nk                nk              This study
17            (> 400) 900 000   5-6 m           Bessesen & Saborio-R.,
                                                2012; Sarmento de
                                                Carvalho, 2013;
                                                Sarmento et
                                                al., 2016
18            884               4-5 m           Samper-Villarreal
                                                 et al., 2014
19            nk                nk              This study
20            ~ 100             4-5 m           This study
21            ~ 1 000           < 1 m           This study

Caribbean
22            nk                nk              Dawson, 1962
23            nk                1 m             Dawson, 1962; Cortes &
                                                Guzman, 1985; Cortes &
                                                Jimenez, 2003; Crow,
                                                 2003b
24            nk                < 2 m           This study
25            400               8 m             Cortes & Guzman, 1985
26            nk                nk              Dawson, 1962
27           200 000            0-7 m           Dawson, 1962;
                                                Wellington, 1974;
                                                Risk et al., 1980;
                                                Cortes & Guzman, 1985;
                                                Cortes & Risk, 1985;
                                                Cortes, 1998; Cortes &
                                                Jimenez, 2003; Crow,
                                                2003b; Fonseca et al.,
                                                2007; Wetzer & Bruce,
                                                1999
28            nk                nk              This study
29            Small patches     nk              Fernandez & Alvarado,
                                                2004
30            160 000           1-5 m           Cortes & Guzman, 1985;
                                                Chacon, McLarney,
                                                Ampie, & Venegas, 1996;
                                                Krupp et al., 2009
31            nk                nk              This study

Notes: (1.) Pers. com. and specimen sampled by M. Heidemeyer.
(2.) Pers. com. Eleazar Ruiz, 2016. (3.) This meadow disappeared in
the mid-1990s; there is photographic evidence. (4.) Unpublished data
JSV & pers. com. Mauricio Mendez, Javier Espinach, Gustavo Rojas-Ortega
sighted 2015 with photographic evidence, 2016. (5.) Pers. com. and
photographs from Giovanni Bassey, Alvaro Segura y Mauricio Mendez,
2016. (6.) Unpublished data JSV & JCN & pers. com. Juan Diego Pacheco
& Andres Beita, 2016. (7.) Pers. com. Helena Molina, 2016.
(8.) Unpublished data JSV & JC. (9.) Juan Esteban Barquero In Prep.
(10.) Seen again in 2012 by JC. * = only some isolated shoots in the
last 15 yrs., pers. obs. JC.
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Date:Apr 1, 2018
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