Borikén IMMA
Size in Square Kilometres
9 380 km2
Qualifying Species and Criteria
Greater Caribbean Manatee – Trichechus manatus manatus
Criterion A; Criterion B(1); Criterion D (1)
Humpback Whale – Megaptera novaeangliae
Criterion C (1, 3)
Marine Mammal Diversity
Trichechus manatus manatus, Megaptera novaeangliae, Globicephala macrorhynchus, Stenella longirostris, Steno bredanensis, Tursiops truncatus, Physeter macrocephalus
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Summary
The Borikén IMMA consists of the coastal, neritic over the insular shelf, and oceanic waters adjacent to the shelf-edge of the island of Puerto Rico. This IMMA harbors a resident and genetically isolated population of Greater Caribbean manatees (Trichechus manatus manatus), together with a diverse ensemble of 21 cetacean species, of which six occur regularly, including humpback whales, which use the IMMA during Northern Hemisphere winter months for breeding.
Description of Qualifying Criteria
Criterion A: Species or Population Vulnerability
The Greater Caribbean manatee (Trichechus manatus manatus) (based on Mignucci-Giannoni et al. 2024), previously known as the Antillean manatee, is red-listed as Endangered based on current population estimates and a decreasing population trend (Self-Sullivan and Mignucci-Giannoni 2008, Self-Sullivan and Mignucci-Giannoni 2012, Morales-Vela et al. 2024). The primary threat to the species until the early 1990s was hunting for their meat (Mignucci-Giannoni et al. 2000, Freire and Mignucci 2022), but since then other major threats have emerged including increased mortality due to watercraft strikes, orphaning of calves, and the effects of small population size and low genetic variability (Mignucci 2010, Freire and Mignucci 2022).
Criterion B: Distribution and Abundance
Sub-criterion B1: Small and Resident Populations
The Greater Caribbean manatee population inhabiting Puerto Rico is considered a resident, small, isolated population, with no known or significant immigration or emigration of individuals from or to other areas (Hunter et al. 2012). Manatees can be found along the entire coast of Puerto Rico but are detected less often along the northern coast where seagrass beds are not as extensive and habitat is less suitable (Powell et al. 1981, Collazo et al. 2019). The highest concentrations are documented in four areas: Ceiba on the east coast, Jobos Bay between Guayama and Salinas on the southeast coast, Guayanilla and Guánica Bay on the southwest coast, and off the Guanajibo River, between Cabo Rojo and Mayagüez on the west coast (Powell et al. 1981, Rathbun et al. 1986, Freeman and Quintero 1990, Drew et al. 2012, Mignucci-Giannoni et al. 2018, Collazo et al. 2019). A recent sighting was recorded for Mona Island, and sightings in Culebra Island have become more common in recent years. Their distribution is dependent on available resources such as freshwater, seagrass, and areas that provide shelter from strong waves (Mignucci-Giannoni 1989, Drew et al. 2012).
While rescue data of orphaned calves includes all months of the year and all municipalities of Puerto Rico, including Culebra Island (Adimey et al. 2012, Caribbean Manatee Conservation Center unpubl. data), aerial survey data (Powell et al. 1981, Rathbun et al. 1985, Freeman and Quintero 1990, Mignucci-Giannoni et al. 2018, Collazo et al. 2019), indicates that the calving occurs also throughout the year, and most commonly in Salinas, Jobos Bay, Guánica, Lajas, Guayanilla Bay, and Ceiba. However, recent sightings attest to calves being raised in the Condado Lagoon, a protected sanctuary in Metropolitan San Juan (Caribbean Manatee Conservation Center, unpubl. data). Helicopter aerial surveys off the south coast (Mignucci-Giannoni et al. 2018), detailed that reproductive groups were most commonly observed in Guanica Bay, Jobos Bay, Juana Díaz, and Peñuelas.
In Puerto Rico the average minimum population estimate for manatees is 386 (range 312-535, standard deviation 89) based on surveys conducted from 2010-2014 (Collazo et al. 2019). Ten percent of the manatees counted during these surveys were calves. One additional survey was recently flown in April 2021 with a direct count of 109 manatees, 11 of those calves (Mignucci-Giannoni 2022), but the estimate from this survey has not yet been calculated. Radiotelemetry data from 35 manatees between 1992 and 2005 (Slone et al. 2006), and between 2010-2017 (Hernández-Lara and Mignucci-Giannoni 2024a,b) in Puerto Rico show short distance movements (~3-15 km) to take advantage of food, water, and shelter resources (Hernández-Lara and Mignucci-Giannoni 2024a).
Mignucci-Giannoni and Beck (1998) detailed the diet of the Greater Caribbean manatee in Puerto Rico, mainly consisting of seagrasses, turtle grass (Thalassia testidinum), manatee grass (Syringodium filiforme), and shoal grass (Halodule wrightii). Some ingestion of green alga (Ulva lactuca), mangrove, and the seaweed Caulerpa prolifera was also documented (Mignucci-Giannoni and Beck 1998). Aerial surveys (Rathbun et al. 1985, Mignucci-Giannoni et al. 2018, Collazo et al. 2019) and telemetry studies (Slone et al. 2006), have revealed important feeding areas for manatees along the east and south coast of the island, including Vieques Island, Ceiba and Naguabo, Patillas, Jobos Bay, Salinas Bay, Santa Isabel, Guayanilla, Guánica, and Cabo Rojo, which has been used by Drew et al. (2012) to recommend boundaries for protected areas in Puerto Rico.
Criterion C: Key Life Cycle Activities
Sub-criterion C1: Reproductive Areas
Humpback whales (Megaptera novaeangliae) are known utilize the coasts of this IMMA, particularly the northern and western coast, during the winter season (November to May, peak season is mid-February to mid-March) (Matilla and Clapham 1989, Mignucci-Giannoni 1989, Burks and Swartz 2000, Sanders et al. 2005, MacKay 2015, MacKay et al. 2016). Females arrive early in the season to give birth in protected waters off Aguada and Aguadilla, and males sing and court females in the Borinquen Bank off Aguadilla, Isabela, off Rincón, Desecheo Island, and banks west of Cabo Rojo and Mayagüez (bajoa de Sico, El Seco, and Sierra) in the Mona Passage (Sanders et al. 2005, MacKay et al 2016). Songs are also readily heard and recorded off Culebra Island (P. Knapp, pers. comm.).
Sub-criterion C3: Migration Routes
While some whales use the IMMA as their primary breeding ground for calving and nursing, others use it as a stop-over on their migration to other North Atlantic humpback whale breeding areas in the Caribbean. Their residency peaks from mid-February to mid-March) (Matilla and Clapham 1989, Mignucci-Giannoni 1989, Burks and Swartz 2000, Sanders et al. 2005, MacKay 2015, MacKay et al. 2016). Their general movement seems to be from east to west, probably headed to Borinquen Bank off Aguadilla and Isabela and to Rincón and Desecheo Island, before moving along to Samaná Bay, Silver and Navidad banks in the Dominican Republic, and later back to their northward migration to the summer feeding grounds. Of 558 individually identified humpback whales in Puerto 43.4% where matched in Eastern Canada (Newfoundland, Labrador, and Gulf of St. Lawrence), 35.7% were matched in the Dominican Republic, 8.4% in the Gulf of Maine, 6.5% off Bermuda, 3.4% off Greenland, 1.5% off Iceland, and 1.1% in Eastern North Atlantic (Martin et al. 1984, NAHWC unpubl. data 2018). Intra-Caribbean sightings of humpback whales has also been recorded between Puerto Rico and Guadeloupe, Anguilla, and Dominica islands in the Lesser Antilles (Stevick et al. 1999, MacKay 2015, MacKay et al. 2019).
Criterion D: Special Attributes
Sub-criterion D1: Distinctiveness
The Greater Caribbean manatee population in Puerto Rico is distinctive enough genetically, based on mitochondrial and nuclear DNA, considered to be marginal, isolated, and with an evolutionary connection with the now extinct population of the Lesser Antilles (Vianna et al. 2006, Hunter et al. 2012, Álvarez-Alemán 2019).
From mitochondrial genetic data (Hunter et al. 2012), two distinct haplotype manatee subpopulations have been detected: Haplotype A01 manatees (30.1%), inhabiting mostly the north and north-eastern coast of the island, and Haplotype B01 animals (65.5%), inhabiting mostly the south and south-western coast of the island. A01 and B01 haplotypes meet in the mid-west coast and mid-east coast, where they seem to intermingle for reproduction to enhance genetic variability (Hunter et al. 2012). Haplotype A01 is shared with manatees in Florida, Cuba, and Dominican Republic, and haplotype B01 is only shared with manatees from the Dominican Republic (Vianna et al. 2006, Álvarez-Alemán 2019, Álvarez-Alemán et al. 2022). Based on this, Hunter et al. (2012), recommended that the Puerto Rico manatee population be managed as a distinct population, compared to the Florida subspecies.
Sub-criterion D2: Diversity
Puerto Rico hosts not only endangered Greater Caribbean manatees, an additional six species are commonly and regularly observed, including humpback whales between the months of November and May of each year, short-finned pilot whales (Globicephala macrorhynchus), spinner dolphins (Stenella longirostris), rough-toothed dolphins (Steno bredanensis), and common bottlenose dolphins (Tursiops truncatus) and less frequently, but with some regularity, sperm whales (Physeter macrocephalus) (Mignucci-Giannoni 1989, Rodríguez-Ferrer et al. 2018).
A further 16 cetacean species are occasionally sighted, but their presence is well documented by less regular sightings and stranding events (Barragán et al. 2023; Bolaños-Jiménez et al. 2014, 2021, 2023; Burks and Swartz 2000; Caballero-Gaitán et al. 2012; Cardona-Maldonado and Mignucci-Giannoni 1999; Carrasquillo-Casado et al. 2002; Erdman 1970; Ewing and Mignucci-Giannoni 2003; Lettrich et al. 2023; MacKay 2015; MacKay et al. 2016; MacKay and Bacon 2019; Mattila and Clapham 1989; Merten and Rodríguez-Ferrer 2014; Mignucci-Giannoni 1988, 1989, 1996, 1998; Mignucci-Giannoni et al. 1998a,b, 1999a,b., 2000, 2003, 2009; Pérez-Zayas et al. 2002; Rodríguez-Ferrer et al. 2018, 2020; Rodríguez-López and Mignucci-Giannoni 1999; Rosario-Delestre et al. 1999; Sanders et al. 2005; Swartz et al. 2001; Tellez et al. 2014).
Supporting Information
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Rodríguez-Ferrer G, Appeldoorn RS, Schizas NV. 2017. Abundance of the common bottlenose dolphin Tursiops truncatus (Montagu 1821) (Mammalia: Artiodactyla) off the south and west coast of Puerto Rico. LEB 4(4):242-268.
Rodríguez-Ferrer G, Cruz-Motta JJ, Schizas NV, Appeldoorn RS. 2020. Modelling distribution of the common bottlenose dolphin, Tursiops truncatus off the southwest coast of Puerto Rico. Journal of Marine Systems, 103371.
Rodríguez-Ferrer G, Appeldoorn RS, Mignucci-Giannoni AA, Rinaldi R, Schizas NV. 2024. The presence of two distinct mitochondrial lineages in the bottlenose dolphin (Tursiops truncatus) in Puerto Rico and their affinities with previously reported lineages. Mammalian Biology 104:499-512.
Rodríguez-Ferrer G, Reyes R, Hammerman NM, García-Hernández JE. 2018. Cetacean sightings in Puerto Rican waters: including the first underwater photographic documentation of a minke whale (Balaenoptera acutorostrata). Latin American Journal of Aquatic Mammals 13(1-2):26-36.
Rodríguez-López MA, Mignucci-Giannoni AA. 1999. A stranded pygmy killer whale (Feresa attenuata) in Puerto Rico. Aquatic Mammals 25(2):119-121.
Rosario-Delestre RJ, Rodríguez-López MA, Mignucci-Giannoni AA, Mead JG. 1999. New records of beaked whales (Mesoplodon spp.) for the Caribbean. Caribbean Journal of Science 35(1-2):144-148.
Sanders IM, Barrios-Santiago JC, Appeldoorn. 2005. Distribution and relative abundance of humpback whales off western Puerto Rico during 1995-1997. Caribbean Journal of Science. 41. 101-107.
Self-Sullivan C, Mignucci-Giannoni AA. 2008. Trichechus manatus ssp. manatus. In: IUCN 2008 IUCN Red List of Threatened Species. Version 2014.2. <www.iucnredlist.org>. Downloaded on 28 October 2014
Self-Sullivan C, Mignucci-Giannoni AA. 2012. West Indian manatees (Trichechus manatus) in the Wider Caribbean Region. In Hines E, Reynolds J, Aragones L, Mignucci Giannoni AA, Marmontel M (eds) Sirenian Conservation: Issues and strategies in developing countries. Gainesvile, Florida: University Press of Florida, pp. 36-46.
Slone, D. H., J. P. Reid, R. K. Bonde, S. M. Butler, B. M. Stith. 2006. Summary of West Indian manatee (Trichechus manatus) tracking by USGS-FISC Sirenia Project in Puerto Rico. With additional information on aerial surveys, carcass recovery and genetics research. United States Geological Survey’s Florida Integrated Science Center, Gainesville.
Stevick PT, Carlson CA, Balcomb KC. 1999. A note on the migratory destinations of humpback whales from the eastern Caribbean. Journal of Cetacean Research and Management, 1, 251–254.
Swartz SL, Martinez A, Stamates J, Burks C, Mignucci-Giannoni AA. 2001. Acoustic and visual survey of cetaceans in the waters of Puerto Rico and the Virgin Islands: February-March 2001. NOAA Technical Memorandum NMFS-SEFSC-463, 62p.
Tellez R, Mignucci-Giannoni AA, Cabellero SJ. 2014. Initial description of short-finned pilot whale (Globicephala macrorhynchus) genetic diversity from the Caribbean. Biochemical Systematics and Ecology 56:196-201.
Vianna JA, Bonde RK, Caballero S, Giraldo JP, Lima RP, Clark A, Marmontel M, Morales-Vela B, de Sousa MJ, Parr L, Rodríguez-Lopez MA, Mignucci-Giannoni AA, Powell JA, Santos FR. 2006. Phylogeography, phylogeny and hybridization in trichechid sirenians: Implications for manatee conservation. Molecular Ecology 15:433–447.
Williams EH, Mignucci-Giannoni AA, Bunkley-Williams L, Bonde RK, Self-Sullivan C, Preen A, Cockcroft VG. 2003. Echeneid-sirenian associations with information on remora diet. Journal of Fish Biology 63:1176-1183.
Wyrosdick H, Chapman A, Mignucci-Giannoni AA, Rivera-Pérez CI, Bonde RK. 2018. Internal parasites of the two subspecies of the West Indian manatee Trichechus manatus. Diseases of Aquatic Organisms 130:145-152.
Wyrosdick HM, Gerhold R, Su C, Mignucci-Giannoni AA, Bonde RK, Chapman A, Rivera-Pérez CI, Martínez J, Miller DL. 2017. Investigating seagrass in Toxoplasma gondii transmission in Florida (Trichechus manatus latirostris) and Antillean (T. m. manatus) manatees. Diseases of Aquatic Organisms 127: 65-69.
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