Course:GEOS303/2023/Puerto Rico

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Main ecosystems of Puerto Rico and surrounding area
Walter-Leith climate diagram for Puerto Rico, completed from a GEOS 303 class activity.

Puerto Rico is a tropical island located in the Caribbean that is considered to be an unincorporated U.S. territory. There are three smaller islands around the main island of Puerto Rico (Mona, Vieques, and Culebra), as well as other scattered ones. It is situated between the North Atlantic Ocean and the Caribbean Sea, at 18.15°N and 66.30°W [1]. The island is mainly covered in hilly terrain, with the highest mountain range, Cordillera Central, reaching over 900m of elevation [2]. Puerto Rico mainly has a mountainous interior leading to coastal lowlands, and then deep waters offshore [1].

The territory has a wet season that spans from May to November, with the north coast receiving the majority of the heavy rainfall and the south coast being in a rain shadow [3]. Therefore the amount of rainfall in specific regions varies based on topography, but areas on average receive between 800mm and 4,000mm per year. Meanwhile, temperatures stay around 20-25℃ due to high amounts of insolation [4].

The territory has many ecoregions and biomes due to its varying topography and climate, as well as covering both land and water areas. These include: tropical wet forest, tropical dry forest, wetlands, grasslands, mangroves, coral reefs, and seagrass beds.

Climate and biomes

Puerto Rico has two very important marine biomes: seagrass beds and coral reefs. Seagrass is found along gently sloping coastlines and thrives in shallow, salty, and brackish waters where light is abundant, typically within one to three meters of the surface[5]. Seagrass beds are one of the most productive marine ecosystems [5], and they provide many ecosystem services. They provide shelter to an array of marine mammals and invertebrates by creating a leafy underwater canopy; moreover, seagrass roots trap sediment, which reduces erosion and buffers coastlines against storms; improve water quality by absorbing nutrients from run off; and produce vast amounts of oxygen through photosynthesis [5]. Although seagrass has seen a decline globally, Puerto Rico has actually seen increased cover between 1950 and 2014, specifically in the regions of La Parguera and Caja de Muertos Island[6].

Coral reefs are some of the most diverse ecosystems in the world and are often referred to as “rainforests of the sea” because of the abundance of diversity within these habitats [7]. Puerto Rico is surrounded by over 5,000 km2 of shallow coral reef systems, which protect the coast from storms and erosion [3]. The east coast of Puerto Rico has well-established fringing reef systems, along with the south and west coasts which both house an array of coral communities[8]. Unlike the rest, the north coast has poorly developed coral reefs because of large swells and bed loads[8]. Puerto Ricans not only depend on these coral reefs for protection but also for food from fishing and income from recreational activities [3]. Puerto Rico's population density and annual number of tourists put pressure on these delicate ecosystems, which inadvertently create adverse effects[8]. The declining health of the reefs is due to coral bleaching, disease, reduced water quality, algae blooms, overfishing, and other anthropogenic activities [8].

Puerto Rico’s tropical dry and wet forests are essential terrestrial biomes that account for much of the biodiversity the island displays. The tropical dry forest biome is located along the southwestern and south-central coast of Puerto Rico, and continues in areas among the adjacent islands of Vieques, Culebra, Mona, Caja de Muertos and Desecheo [9]. The southwest coast of the island is subject to most coverage of tropical dry forest due to the rain shadow of the central mountain system [10]. On average, the dry forest has low annual precipitation, ranging from 600mm to 1000mm [9]. Furthermore, the biome only experiences a single rainy season, while the dry season lasts several months [9]. Temperatures fluctuate by 10º throughout the year, ranging from a high of 31ºC in summer months (June to September), to a low of 21ºC in the winter (December to March) [9]. Vegetation found in the forest has made several adaptations to cope with the dry climate, ranging from deciduous leaves, water storage structures, and wavy coating on branches, leaves and trunks [9]. The biome is an essential habitat to many species, such as The Puerto Rican Nightjar (Caprimulgus noctiferia), an endemic bird species only found in the dryer southwest areas of the forest [9].

Puerto Rico's tropical wet forest biome is contained to one sole area, El Yunque National Forest. The forest is located in northeastern Puerto Rico and spans 28000 acres across the slopes of the Sierra de Luquillo mountains [11]. Constant solar insolation throughout the year provides for consistent air temperatures, averaging at 21ºC [4]. The forest experiences a year-round tropical climate with average temperatures of 26ºC in the summer and 20ºC in the Winter [12]. El Yunque National Forest has no distinct wet or dry season as it rains year round. Average annual precipitation is 3048mm (120 inches), with a high of 6096mm (240 inches) [13]. High daily temperatures and constant daylight length provides for ideal, year round growing conditions, allowing the forest to have high productivity and biodiversity [12]. El Yunque National Forest is composed of four different forest vegetation areas: Palo Colorado Forest, Dwarf Forest, Tabonuco Forest and Sierra Palm Forest. These four sub-regions present some differences in vegetation and species due to their location and elevation on the Sierra de Luquillo mountains. The forest as a whole is subject to high levels of biodiversity, with countless endemic and native species [12].

Visualization of where wetlands and mangroves form on Puerto Rico.

Mangroves are a vitally important ecosystem that only take up roughly 1% of the total surface area of Puerto Rico, at about roughly 8300 hectares[14]. The mangrove trees that make up these biomes sit in salt water and provide several services. The salinity of the water promotes a lot of carbon absorption and sequestration compared to other forest ecosystems. The trees in the area also provide great habitat and coverage for fish, crustaceans, and other aquatic and subaquatic species[14]. The mangrove growths bordering the edge of Puerto Rico also serves as a natural storm barrier for tropical hurricanes and storms. All 4 species of mangroves found in the Caribbean can be found in Puerto Rican mangrove as well: red mangrove (Rhizophora mangle), black mangrove (Avicennia germinans), white mangrove (Laguncularia racemosa), and button mangrove (Conocarpus erectus). However, urban encroachment and destruction of these mangrove habitats as well as eco-tourism puts these vulnerable biomes at risk[15].

The grasslands and wetlands of Puerto Rico are in a similar state to the mangrove forests, in that constant human expansion and use have caused these areas to shrink in size[16]. The wetlands are transitional zones between the drier grasslands and forests and the coastal regions, with mangrove and wetland habitats tending to overlap[17]. Wetlands can also be found in the more mountainous regions of the island, with an approximate 13,000 acres of wetland between cloud, colorado, and palm forest[17]. These higher wetlands also serve as unique ecosystems that provide habitats for rare species not found anywhere else on the island, as well as a water reservoir for urban settlements downstream[17]. The grasslands in Puerto Rico appear to be predominantly manmade, formerly made of forest but cut and burned to create space for livestock grazing and plantations. As of a 2003 Landsat survey, 41% of eastern Puerto Rico is grassland[16]. The endemic bird species of the island have been able to adapt to the creation of grasslands, while the dominant mammal species of the areas are all introduced livestock[18].

Diversity

Terrestrial Ecosystems

Puerto Rico is part of the Greater Antilles, a collection of islands in the Caribbean Sea, and one of the globally recognized biodiversity hotspots [19]. The island encompasses many different terrestrial biomes, but is dominated by wet and dry tropical forests. Meanwhile wetlands, grasslands, and mangroves support ecosystems closer to the coastline of the island. Due to its range of terrain, Puerto Rico hosts a large range of species. The majority of present-day terrestrial biodiversity of the island, along with the rest of the Greater Antilles is predicted to extend back to the permanent emergence of the Greater Antilles, 40 million years ago. The processes involved vicariance and over-water and long distance dispersal events. Vicariant events can be linked to the existence of the Greater Antilles Aves Ridge, a land bridge connecting the group of islands to continental South America during the Eocene–Oligocene[19]. The land bridge, along with other vicariance and dispersal events have allowed for Puerto Rico’s great biodiversity.

The El Yunque National Forest lies on the eastern side of the island, and is a prime spot for biodiversity. Despite having an area just short of 30,000 acres, it provides for over a thousand plant species, including 250 types of trees and over 100 ferns [20]. Apart from the El Yunque, Puerto Rico has a unique layout of its forests due to anthropogenic sources. The island has formed a mosaic of forest patches and corridors, which are able to sustain a range of biodiversity in the region [21].

The island also hosts many groups, including bats, reptiles, birds, as well as a significant species of frog. The thirteen species of bats on Puerto Rico are the only native mammal remaining on the island, yet are relatively uncommon [22]. Frugivorous bats such as these play a key role in pollination, dispersal, and insect control in tropical forests [23]. Every other mammal that exists on Puerto Rico today have been brought by humans to the island, including domesticated animals and rats from early European settlers [24].

One of the most notable species on Puerto Rico is the E. Coqui, a type of frog native to the area. Also called the common Coqui, it is known to be the most abundant frog species on the island at around 200,000 per hectare [25].

Land-use changes have threatened the biodiversity of Puerto Rico. From Spanish colonialism in the early 1500s to the present-day U.S occupation and industrialization of the island, land-use changes have altered greatly [26]. Exploitation of the soil and deforestation and mismanagement of the forests to make room for agriculture has historically altered the biomes and biodiversity of the island. However, industry and land-use changes within the last few decades has led to a decline in agricultural land, leading to increased cover of grasslands, shrubland woodland and forest cover, as well as greater urbanization [26].

Marine Ecosystems

Puerto Rico’s surrounding waters are filled with different species, and the country's species richness ranks among the highest within the Caribbean. Notably, the island waters contain a much higher number of mussel species compared to the rest of the Antilles and the Caribbean[27], as well as featuring three of the five bioluminescent bays in the world: Mosquito Bay, Laguna Grande, and La Parguera[28]. Numerous species types can be found within Puerto Rico’s marine ecosystems, including primitive sponge invertebrates, scleractinian corals, mollusks, crustaceous amphipods, and echinoderms. Of these phylum classes, Puerto Rico has been reported to house 40 different species of sponges, 72 hard corals, 1078 mollusks, 25 amphipods, and 121 echinoderms, totalling 1336 species within five ecoregions. With a total coastline length of 501 kilometres, this calculates to 262 species per 100km[27].

Seagrass beds are abundant and valuable to the diversity of Puerto Rico’s marine ecosystem, found all across shallow waters surrounding both the main and smaller islands. Various seagrass species are supported by clear, oligotrophic, or oxygen-rich, waters and stretch far until barriers are reached[29]. Seagrass species include, from most to least common, Turtle grass, Manatee grass, Shoal grass (colonizer species), Paddle grass, and Widgeon grass. Seagrass meadows, coral reefs, and mangrove forests experience tight biotic interactions and are thus extremely interconnected, stabilizing the greater ecosystem of Puerto Rico[30]. Said seagrass cover contributes to sediment stabilization, and provides feed and shelter for both birds and marine organisms, in turn, supporting higher productivity rates and increasing the biodiversity of the area[29].

A snapshot of a coral reef found off the coast of Puerto Rico

The coral reefs of Puerto Rico take up roughly 3,400 square kilometres around the main island and the surrounding smaller islands. For inshore reefs, algae coverage ranges from 4-49%, while offshore reefs are in generally better condition. However, all coral zones suffered during the 2005 coral bleaching event in the region[31]. The reef services 242 different reef fish species such as parrotfish, groupers, grunts, snappers, and barracudas, and is commonly commercially fished and subject to overexploitation[31]. The species that frequent the coral reefs also travel in between the mangroves that line the edges of the islands, the seagrass beds, and make trips past the edge of the continental shelf and into deeper waters[32]. The most diverse coral location was around El Palo reef, while the highest species richness area occurred at a shelf edge between El Palo and Margarita reef[32].

There have been advancements in studying Puerto Rico’s ecosystems past its rich terrestrial forests and coastline and into the seafloor as the Puerto Rico trench is the deepest in all of the Atlantic Ocean. Recently, a new and rare species was discovered, described as the isopod genus Acanthocope (Munnopsidae). This trench is over 8500 meters deep and is the subduction zone of the North American plate going under the Caribbean tectonic plate [33]. It is expected to support more species that have yet to be identified, vastly contrasting the organisms found in Puerto Rico’s shallow waters.

Endemism

Puerto Rico is home to many faunal species, some of which can only be found on the Puerto Rican islands. There are a total of 354 bird species in Puerto Rico, 16 of which are endemic[34]. Some notable species are the green mango (Anthracothorax viridis), the Puerto Rican emerald (Chlorostilbon maugaeu), and the Puerto Rican vireo (Vireo latimeri)[35]. There are 25 amphibian species; 19 are native to Puerto Rico, and 16 are endemic[35]. This includes the melodius coqui (Eleutherodactylus wightmanae) and the locust coqui (Eleutherodactylus locustus)[35]. Then there are 56 reptile species, out of which there are 42 endemic terrestrial species, including the island's largest snake, the Puerto Rican boa (Epicrates inornatus)[35]. Within this category, there are 28 endemic lizard species, such as the Mona Island iguana (Cyclura stejnegeri) and the arboreal giant anole (Anolis cuvieri)[35].

Human influences

History

Puerto Rico was originally inhabited by hunter-gatherer native populations. Up until the 15th century, there had been zero contact with others outside of those living in the Caribbean region, but this changed with the arrival of Christopher Columbus and the Spanish colonists [36]. The island was named Puerto Rico (“Rich Port”), and within two decades the native populations had been placed under colonial rule and forced to work in farming and the gold mines. The Spanish Crown eventually decreed that the natives be treated the same as Spaniards, but the damage had been done and the population had been decimated by disease and abuse[37]. Meanwhile, the Spanish had started importing slaves from Africa to staff the farms beginning to establish, with several towns popping up due to the highly sheltered bays serving as harbours[36]. The island would remain under Spanish rule and while fending off pirates, British, Dutch, and French attackers the island would cultivate a primarily coffee based economy, mostly trading with the United States[36].

Sugarcane plantation in 1946

Up until the 18th century, the main crops included tobacco, cacao, rice, and other miscellaneous plants. Around the annexation of Puerto Rico into the United States in 1898, there began a shift from just normal subsistence farming to exploitation of the rapidly growing sugarcane market[37], with the previously dominant coffee crop cultivated by the Spanish replaced with the highly valuable sugar cash crop. However, with the abolition of slavery in 1873, the slavery model plantation system was starting to decline. The US government instituted Charles Allen to govern the territory, who immediately economically pushed for the sugar trade[38]. Approximately 45% of farmable land on the island was converted into monoculture sugar plantations[38]. The clearing of Puerto Rican forest and land for plantation use continued as more sugar, tobacco, and coffee were planted for economic gain. In 1941, Puerto Rico legislature enacted a land reform act to specifically cull the power of the monoculture sugar industry, which had taken over the island so completely that it was described as “a mountain range entirely surrounded by sugar cane”[39]. This land reform not only decreased the extent of monoculture practices but also led to farmers being dispossessed from their subsistence-based farmland [40].

New forests began to emerge once the extent of plantations and other small farms decreased, allowing natural reclamation of new ecosystems and biodiversity growth. This reforestation is not at all homogenous across the territory – research has shown there are very few areas of continuous forest, and instead predominantly scattered forest fragments [41]. Afforestation tended to occur on former coffee-producing land, some upland tobacco and sugar farms, but mainly on land that was for small-scale subsistence farming [40]. Both the historic and current use of natural resources has shaped the composition of Puerto Rico’s forests, including the previously mentioned plantations, as well as current-day agroforestry, charcoal production, and logging [42]. Species found today in these forests differ drastically from pre-colonial forests, and include a large percentage of alien tree species [41]. Urbanization and growth of settlements has also been a major driver of land use change in Puerto Rico, with smaller urbanized areas found dispersed throughout forest fragments around the main island [41]. There has also been a high degree of urban sprawl originating from the city of San Juan, largely due to transportation infrastructure and residential growth [43].

Environmental Impacts

As a tropical island, Puerto Rico is highly susceptible to climate change causing a large impact on their ecosystem. According to The Puerto Rico Climate Change Council (PRCCC) and the NOAA National Centers for Environmental Information (NOAA), there has been a measured increase of almost 2°F since 1950 due to increased greenhouse gas emissions. This has also contributed to the almost 2°F warming of the ocean waters surrounding Puerto Rico, particularly the capital of San Juan where the effect of an urban heat island has further increased felt temperatures. They projected severe increases in extreme heat events causing droughts and water-stressed fauna and flora [44].

Additionally, there has been a monitored 0.7-inch increase in sea level rise every decade since 1961. Sea levels are projected to continue to rise and with rising sea surface temperatures there is projected to be an increase in extreme precipitation event intensity. Puerto Rico’s location being within the Caribbean hurricane belt increases their chances for tropical cyclone events. Hurricane Maria in September of 2017 was the second most devastating natural disaster for the island, categorized at level 4. Extreme heavy rainfall and storm surges caused severe damage to the island's infrastructure. Flooding and mudslides also heavily damaged the natural environment, eroding a large percentage of the coastline [44].

Anthropogenic stresses have caused major changes to Puerto Rico's environment over the past several decades. Human impacts on the environment range from deforestation, urbanization, industrial development, pollution, overfishing, improper waste management, poaching, climate change and introduction of invasive species. Many of these issues are caused by land use changes, urbanization, and population growth [45]. Both terrestrial and marine ecosystems are negatively impacted by these stressors which are causing biodiversity loss, fragmentation, soil erosion and more. Historical agricultural processes, urbanization and infrastructure development has caused significant deforestation in Puerto Rico [45].

In coastal and marine ecosystems, coral reefs face significant impacts of increased eutrophication, water contamination and sedimentation [45]. Numerous dredge disposal sites are found near coral reefs, and sediments from these sites are smothering the corals [46]. Furthermore, erosion and loss of wetlands due to coastal urban and industrial development has led to serious flooding events posing risk to local populations. Loss and fragmentation of natural barriers that would normally protect against flooding poses a threat to biodiversity as well [47]. Overfishing has also negatively impacted coastal and marine ecosystems and biodiversity. Declining fish populations pose risk to the ecosystem stability as well as affects the livelihoods of local communities [48].

Future Implications

At this time there is little scientific research done on the future implications of human influence on the ecology of Puerto Rico. This is an area that can be investigated further in future research.

Conservation

Puerto Rican Boa

Image of the Puerto Rican boa

The Puerto Rican boa is an endemic species to Puerto Rico that has been protected since the early 1970s under the U.S. Endangered Species Act due to its population decline[35]. These boas were on the endangered list for over four decades without showing many signs of recovery [49]. Information on the Puerto Rican Boa is extremely sparse, and not much is known about the species ecology, natural history, and genetics; thus, recovery efforts have not been successful in the past[49]. Throughout the last decade, conservation efforts have been put into place to protect the species in hopes of aiding in their recovery [35]. These efforts include measures to stop them from being hunted and collected, gathering new and relevant information on them, conserving their habitat, as well as spreading awareness and outreach[35]. Conservation efforts are relatively new and still continuing for the boa, with future goals to create management strategies and recovery plans for the endangered species'[35]. This is crucial, as losing this species in Puerto Rico means they are gone entirely, further adding to the loss of earth's biodiversity.

Puerto Rican Parrot

Image of the Puerto Rican parrot

Puerto Rico is home to an endangered species of parrot, commonly referred to as Iguaca or simply the Puerto Rican parrot. At the beginning of the 16th century there were rumoured to be close to a million of this species living in the territory, but their populations have dwindled due to habitat loss from human activity and extreme weather events[50]. This parrot is known to naturally reside in both the El Yunque National Forest and the Río Abajo State Forest, and more recently, the species was introduced to the Maricao Commonwealth Forest[51]. In 1938, less than thirty parrots remained on the island which prompted the creation of the Puerto Rican Parrot Recovery Plan, involving government-based and other organizations from both the US and local groups[50]. The project aims to “...conserve, protect and manage wild and captive parrots in order to downlist the species from endangered to threatened”[51]. Two aviaries that focus on breeding and supporting existing populations have helped the Iguaca recover, mainly through a metapopulation approach to conservation – creating multiple, more distinct populations of the species[52]. As of March 2022, there are around 250 wild parrots living in Puerto Rico’s forests thanks to conservation efforts[53].

Image of a Ravenia urbanii flower

Endemic Tree

The Botanic Gardens Conservation International organization developed the BGCI’s tree Conservation Programme to facilitate the preservation of the two Puerto Rican endemic tree species Garcinia portoricensis and Ravenia urbanii. Endemic species to the island are adapted to hurricane occurrence, however, these two flora are threatened species that were greatly impacted by Hurricane Maria and Hurricane Irma, both occurring in 2017. The 2021 project was conducted through field surveys, specimen collection, propagating materials, reinforcing wild sites and a hurricane-resistant nursery[54] With partnerships and funding, said protected sites were established and propagation of seedlings was successfully transferred. Eye On The Rainforest, a local NGO and the main partner for this project, cultivated gardens for said seedlings and supports scientific research, botanical care and educational outreach for the islands endemic species [55] Currently, growing of the young trees within nurseries are still being carried out before they are able to be sustainability introduced back into wild sites, due to such, both species are still considered under-threat.

Image of a Puerto Rican crested toad

The Puerto Rican Crested Toad

The Puerto Rican Crested Toad (Peltophryne lemur) is the only toad species endemic to Puerto Rico. The species was first described in 1868, and has since faced numerous threats leading to its critical endangerment today. The species inhabits dry and semi-arid climates and burrows in openings in limestone karst [56]. Invasive species and habitat loss are the main threats that have led to the significant decline in the toads populations [57]. The toad was thought to have been extinct from 1931 to 1967, before populations were rediscovered in northern Puerto Rico [56]. To keep the species from going extinct, in 1984 the American Association of Zoos and aquariums developed a species survival plan (SSP). Objectives of the plan included research, education and outreach, protection of existing habitats, creation of new habitats and to have at least six self-sustaining populations in the wild [57]. Part of the plan included the reintroduction of more than 3,400 Puerto Rican Crested Toads between 1982 and 1992. Since then, over 300,000 tadpoles have been reintroduced. In 2019, Potter Park Zoo sent 2,000 tadpoles to Puerto Rico to continue conservation efforts [58]. Invasive species and habitat loss continue to threaten the species, as well as inundation of breeding ponds during storms [59]. Although recovery efforts are extensive and still occurring, involving various conversation strategies part of the SSP, the Puerto Rican Crested Toad remains critically endangered [59].

A patch of mangroves growing in Lajas, Puerto Rico

Mangroves and their services

Mangrove trees and their presence are a critical component of the biosphere along the coast of Puerto Rico. Not only do they serve as habitats for local terrestrial and aquatic wildlife, they also act as a natural soil fortification and storm barrier for the island[14]. However, years of human influence has destroyed mangrove forests for uses such as agricultural expansion, pollution disposal, and urban development [60]. Restoration efforts began in earnest in 2016 spearheaded by the NGO Para la Naturaleza [61], and operations have started scaling up since Hurricane Maria in 2017 left many riverbanks and coastlines exposed and crumbling. This has also threatened human settlements due to the erosion of the ground and shoreline [60]. The restoration efforts aim to return Puerto Rico to an “ecological culture” that values the ecosystem services provided by a healthy ecosystem, as well as educating future generations about the importance of these natural barriers and how fragile they are in the face of human exploitation[61].

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Seekiefer (Pinus halepensis) 9months-fromtop.jpg
This Tropical Ecology Resource was created by Course:GEOS303.


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