The Bats of Matanzas

The Province of Matanzas, in western Cuba, is known for the wonderful white sands of Varadero Beach, its turquoise waters, the amazing Bellamar caves, and the Zapata Swamp, the largest "humedal" in all the Caribbean. What Matanzas is not known for, however, is for its richness in bat species. Of the 28 living species recorded for the Cuban archipelago, 26 inhabit the province of Matanzas, representing the six bat families that inhabit Cuba (1).

Leach's Single-Leaf nosed bat (Monophyllus redmani).
This species feeds mostly on pollen and plays a key role in pollination of plants.

A reason for the high diversity of bats in Matanzas may be that Cuba does not possess major geographical barriers such as very tall mountains or deserts. Instead, the island is characterized by its low-lying landscape, with hills that rarely surpass 300 m in height. As a result, bat distribution in Cuba is highly homogeneous. Similar numbers of species are found in all other of Cuba's 15 provinces. This could be a reflection of the area's most recent geological history or less collecting efforts in the rest of Cuba.

The Cuban Archipelago (GoogleEarth). 

Bats are amazing creatures, with amazing adaptations. With their skin-webbed wings, velvety fur, and sharp teeth, bats have probably cruised the Cuban skies in search of food and shelter for a least 33 million years (Eocene-Oligocene), when the island emerged and became available for colonization; although, unfortunately, we only have bat fossils from the last 20 thousand years (2).

Waterhouse's Leaf-nosed bat (Macrotus waterhousei). 

The biological diversity and uniqueness of Cuba is a result of the island’s intricate geological history and its long isolation from the mainland. Over 60% of the Cuban landscape is karstic, and nearly 80 % if the submerged platform is counted, indicating a high potential in the availability of caves, crucial shelters that allow high species richness. In fact, this has been correlated by bat researchers (Brunett and Medellin, 2001). Of the 28 known Cuban bats, 15 are strict cave -dwellers, with most others using caves opportunistically (1).

Insectivorous Waterhouse's Leaf-nosed bat (Macrotus waterhousei) in flight

Here is where Matanzas shines. Matanzas harbors today the most extensive subaerial karst region of the entire Cuban archipelago, a potentially very cave-rich region ~65,500 km² wide. Probably, no other province in Cuba has more caves available for bat roosting than Matanzas today. Moreover, this was more strikingly so 10,000 years ago, when the Gulf of Batabanó, south of the western half of Cuba, had the largest potential in the availability of caves for bat roosting anywhere in the Cuban archipelago, competing in the Caribbean only with the Bahama bank. Once the ice of the last glacial maximum melted with the warmer temperatures of the Holocene epoch, sea level rose and inundated most of the Cuban ancient karst plains, drowning about ~13,300 km² of latent cave-rich territory (3), essential for bat life in the island, and likely culling the territory of a few species. Many have postulated this as the reason for the disappearance of several bat species.

Jamaican Fruit-eating bat Artibeus jamaicensis  roosting on
the calcarenite limestone of Varadero's Ambrosio Cave. 

Matanzas has played an important role in the study of Cuban bats since at least the XIX century. Four of Cuba's bats Pteronotus parnelli, Pteronotus quadridens, Phyllonycteris poeyi and Tadarida brasiliensis (muscula), were collected and described for the first time from Matanzas, near the coffee plantation Fundador de Canímar. This feat is the work of the German naturalist, Johannes Gundlach.

Sooty Moustached-bat Pteronotus quadridens

Gundlach stopped in Cuba on his way to South America and fell in love with the island. I venture to say, he fell in love with Matanzas as well, for he took residence there for nearly the rest of his life. He settled in the lush region near the Canímar River, where he stayed with the Booth family who had plantations there. Gundlach roamed the countryside, especially the Zapata Swamp, and the Canímar River gorge where he observed and collected specimens of mollusks, reptiles, and bats.

Albumen print of Johannes Gundlach (XIX century)

It is through the work of the proliferous Johannes Gundlach and Gilberto Silva Taboada that I came to love bats. In 1992, my parents gave me Silva Taboada's Los Murcielagos de Cuba (The Bats of Cuba), which to my delight had a great introduction to the life of Gundlach and his bat research.

Two-thousand-year-old fossils of Jamaican Fruit-eating bat (above)
and the ultra rare Cuban pallid bat Antrozous koopmani (below). 

Under the auspice of Gundlach and Silva, I studied the bats living in the roof of our schools and nearby caves, amassing a large set of information, with other colleagues, on the bat diversity in the city and nearby caves. This information resulted in over 100 new fossil and modern bat-collecting localities, several publications, and first records for the province of Matanzas.
For example, we (Ricardo Viera and I) reported the new records of the rare and extinct Common vampire bat Desmodus rotundus, Cuban fruit-eating bat Artibeus anthonyi, Peter’s ghost-faced bat Mormoops megalophylla, Greater funnel-eared bat Natalus primus, and Koopman’s pallid bat Antrozous koopmani. In addition, to new records of living Cuban lesser funnel-eared bat Chilonatalus macer, Cuban yellow bat Lasiurus insularis and Pfeiffer’s red bat Lasiurus pfeifferi , and including remote localities in the Zapata Swamp as in the urban Varadero (see publications here, and Viera's here).

A male Jamaican Fruit-eating bat Artibeus jamaicensis
from Palenque Hill Cave, Mayabeque. 

Currently, we are finishing a gazetteer on all the known fossil and modern bat localities in the province that can be useful towards entropy modeling for species distribution in the archipelago. We hope to collaborate with all those interested.

More so, the research continues. Some of our findings have been corroborated by Proyecto CUBABAT under the direction of Melissa Connelly, with the collaboration of colleagues in Matanzas. They have recently reported, and photographed, the Cuban fig-eating bat Phyllops falcatus in Varadero, so far only reported there from fossil remains (see citations above), and the Cuban lesser funnel-eared bat Chilonatalus macer, and Pfeiffer’s red bat Lasiurus pfeifferi (M. Connelly, pers. comm.) This project has a great potential, for it disseminates important information on the ecological importance of bats. Additionally,  through research, they collect useful data crucial for bat conservation in not only Matanzas but also all of Cuba and the Greater Antilles.

We wish them success!


Acknowledgements

I thank once more, my friend and mentor Dr. Adrian Tejedor for his support and guidance. And once again for helping unravel my torturous prose. Thank you profe. I also thank Ricardo A. Viera, Lazaro Vinola, Leonel Perez, Canido Santana, and Joel Monzon for the information provided and years of trecking up and down the caves of Matanzas in search of bats and fossils.

Sources

1. Silva-Taboada, G. 1979. Los Murciélagos de Cuba. Editorial Academia, La Habana. 424pp.

2. Iturralde-Vinent, M. see his geological literature regarding Matanzas on Biblioteca Digital Cubana de Geociencias.

3. Atlas Nacional de Cuba 1969-1985.

Jiménez, O., M. M. Condis, and E. García. 2005. Vertebrados post-glaciales en un residuario fósil de Tyto alba scopoli (Aves: Tytonidae) en el occidente de Cuba. Revista Mexicana de Mastozoología, 9:84-111.

Orihuela, J. 2011. Skull variation of the vampire bat Desmodus rotundus (Chiroptera: Phyllostomidae): Taxonomic implications for the Cuban fossil vampire bat Desmodus puntajudensis. Chiroptera Neotropical 17(1): 963-976.

Orihuela, J. 2012. Late Holocene fauna from a cave deposit in Western Cuba: post-Columbian occurrence of the vampire bat Desmodus rotundus (Phyllostomidae: Desmodontinae). Caribbean Journal of Science, 46 (2): 297-313.

Orihuela, J., and A. Tejedor. 2012. Peter's ghost-faced bat Mormoops megalophylla (Chiroptera: Mormoopidae) from a pre-Columbian archaeological deposit in Cuba. Acta Chiropterologica 14(1): 63-72.

Orihuela, J., R. Viera, and L. Vinola. 2017. New bat records based on modern and fossil remains from the province of Matanzas, Cuba.

Suárez, W. 2005. Taxonomic Status of the Cuban Vampire Bat (Chiroptera: Phyllostomidae: Desmodontinae: Desmodus). Caribbean Journal of Science 41 (4):761-767.

Viera, R. A. 2004. Aportes a la Quiropterofauna nacional. 1861: Revista de Espeleologia y Arqueologia, Matanzas, 5 (1): 21-23.

Woloszyn, B.W., and N.A. Mayo. 1974. Postglacial remains of a vampire bat (Chiroptera: Desmodus) from Cuba. Acta Zool.Cracoviensia 19:253-265.

The living cave floor: intro to cave faunas

E. A. Martell, one of the first professional cave explorers, expressed in 1894 that "caverns and abysses are natural laboratories ready for numerous and curious researchers".

Those who visit caves frequently notice that some caves are alive! Walk into a cave room and with a swipe of the lantern you see the cave floor move! - Seems like something out of a horror film, but caves are truly alive.

Cave roaches of the family Polyphagidae, scavengers of the guano cave floors. This is an example of a guanophile troglobites. 

Caves are alive because they are teeming with life. From the microscopic viruses and bacteria to the much more complex bats, and even humans, caves provide a subterranean environment that is both beautiful and mysterious, resulting in the specialization of extraordinary organisms.

American roach Periplaneta americana (Blattidae) another common cave scavenger. 

The subterranean world is a complex environment. In caves, light enters unevenly. The deepest rooms, those farthest away from the entrance are the darkest if no other source of light exists. But even in such absence of light, organisms proliferate and evolve. These cave organisms can be defined or grouped depending on their cave habitat and preferences. Each has its own niche.

Troglobites are organisms that are strict cave dwellers, and complete their life cycles deep inside caves. Troglophiles are occasional cave dwellers. They do not complete their life cycle inside the cave, but they could if needed or preferred. They can be facultative cave inhabitants.

Trogloxenes, on the other hand, are similar to troglophiles in that they too are facultative cave dwellers, but their life cycles always require they leave the cave, surfacing to eat or reproduce. In this case, most organisms that people associate with cave dwelling are readily trogloxenes. Bats, birds like oil birds, owls, swallows, some reptiles, amphibians, and several species of insects, are all trogloxenes.

Cavernicolan glowworm in a cave in the Dominican Republic. Photographs courtesy of Adrian Tejedor. This is the larval state of a trogloxene arthropod. It is bioluminescent and truly interesting to observe. 

Other cave organisms are strictly aquatic cave dwellers. These are called stygobites, for they can only live in cave ponds or lakes. The blind fish and arthropods that live in vadose water lakes found within deep parts of a cave system represent this group. These organisms are very specialized. They have adapted to the dark aquatic environments and have lost their eyesights, color pigmentation, or eyes altogether.

There are organisms that can be considered accidentals because they enter caves by falling in, or because they are momentarily attracted by food or shelter inside. These include several mammals such as rodents and carnivores, birds, reptiles, amphibians, and many arthropods. Troglodyte is a term reserved for humans who use caves as a home or shelter. Humans have been using caves since the paleolithic, nearly a million years ago!

A carcass of a Cuban fruit-eating bat Artibeus jamaicensis eaten by
cave roaches of the family Polyphagidae: scavengers or the cave floor.
Cueva de Los Nesofontes, Mayabeque, Cuba 2003.

Caves not only provide secluded microclimates but microhabitats that are richly ecologically stratified and interconnected. An example is illustrated by numerous species of different bacteria and arthropods, that as an adaptation, live off the bat droppings (or guano) deposited in cave floors. These are known as guanophiles and are often troglobitic arthropods.

Plants growing from seeds brought in the excrement
of fruit bats A. jamaicensis. The Dominican Republic, 2004.

Trogloxenes often serve as the food of troglobites. Many are scavengers that feed on decomposing matter brought in by other organisms, such as seeds brought in by fruit bats, or the dying bats themselves, as is the case of the Cuban fruit bat Artibeus jamaicensis carcass devoured by cave-floor roaches in a cave in Cuba shown above.
Many of these organisms are agents of bone accumulations and dispersion within caves, a micro-field within taphonomy that I am most interested in. See my previous ruminations on this previous post.

Decomposed and guano-buried funnel-eared bat Natalus major eaten by small cave-floor ticks genus Pantricola (?). Los Haitises, the Dominican Republic, 2004.

A wide variety of organisms has evolved in caves. These are characterized by intricate specializations such as staged reproduction cycles, feeding mechanisms, or sensitivities to light that allows them to exist only in the intricate microclimates-ecosystems found in the subterranean world. Such diversity and these special characteristics have been naturally selected over millennia depending on the distribution of these organisms and the niches they inhabit withing the geography of a cave.

Cave lake in the interior of Feather's Cave, Matanzas, Cuba.This lake used to have blind fish of the genus Lucifuga. 

Most of troglobites and trogloxenes are relics of the past. Some of the aquatic stygofauna are descendants of freshwater, intertidal, or marine organisms that became trapped in a cave pools and lakes, becoming isolated there. Most of them evolved from exterior faunas that adapted to the cave conditions after isolation.

Cuban boa Chilabrothus angulifer on the hunt for fruit bats. Nesofonte's Cave, Mayabeque, Cuba. This species is a trogloxene that uses caves as a refuge, for feeding and reproduction. These reptiles have adapted to hang from the walls and catch flying bats. 

Exterior faunas enter caves with inflowing rivers, floodwaters, and groundwaters that invade, actively or passively, cave environments. Others become trapped or fall in accidentally as mentioned before. More so, streams and rivers can carry inside sediment and plant debris often carrying organisms from the exterior into the cave. These organisms then become underground colonizers, evolving in isolation from their exterior populations. The cave itself being the barrier to their gene flow.

Cave gours or pools from Bellamar's Cave, in Matanzas. Some of the green and red films that can be seen represent cyanobacteria that live off the chemicals of the rocks and the dim light that is artificially provided by the tour lamps (they are photosynthetic, thus needing light). 

But caves are sensitive environments and even the most minute changes can alter their microhabitats. Alterations to these delicate subterranean environments include hoards of people using caves as a tourist attraction, or "collecting" their endemic faunas or crystal formations, often downgrading their natural beauty. Or affecting their natural flora and fauna, thus affecting the natural cycles that occur within caves. Cave microenvironments are also sensitive to overall climate change, and rising sea levels. The former have the potential to drown existing habitats, and thus pushing many cave endemics to extinction or habitat fragmentation.

The isolation of subterranean environments makes cave faunas interesting subjects to the study of evolutionary pathways, genetics, biogeography, and even virology (-the Ebola and Zaire viruses seem to have evolved in African cave environments-). Some biologists consider caves even as islands, isolated natural laboratories for evolution and natural selection. 

I think Martell was right. And like him, and many other generations of cave scientists, also known as speleologists, we continue to be attracted to caves and their intricacies. We study and divulge the beauty and complexity of caves and cave systems with the hope of understanding such environments and ecosystems better, hopefully helping us protect and understand their faunas more effectively. 

I hope this post has been informative of other interesting aspects of caves since I have already covered bats, hot caves, and cave formation in my previous posts. This post is in honor of those who dedicate their time, and often health, to study caves and unraveling the mysteries of the subterranean world.

Stay tuned for more future ruminations on caves and cave faunas!

Dominican Republic: A Story of Caves and Bats

"In the island, which I have said before was called Hispaniola, there are very lofty and beautiful mountains, great farms, groves and fields, most fertile both for cultivation and for pasturage, and well adapted for constructing buildings. The convenience of the harbors in this island, and the excellence of the rivers, in volume and salubrity, surpass human belief, unless one should see them"

Letter of Christopher Columbus to King Ferdinand and Queen Isabella of Spain, 1492



The heights of Pico Duarte (3098 m), and the Cordillera Central. Behind, the Chain de la Selle or Sierra Baoruco.

November 2004 found me on the island of Hispaniola. To my great pleasure and experience, I was more than very excited to go. His research concentrated on the study of a peculiar group of bats called natalids for his doctoral dissertation (see results here). That study entitled surveying and studying living populations of these bats in their natural habitats, and visiting the island of Hispaniola was essential.

Practically straight out of the plane, we were scouting for areas to set our mistnets and observe our first bats. On that first night, near the quintessential city of Santo Domingo, we captured a female fig-eating bat Phyllops falcatus (haitiensis), which was weighted, measured, and released. The efforts were rewarded by the company of researchers Adrian Tejedor, Kevin Murray and Nelson Marcano.



Fig-eating bat Phyllops falcatus (haitiensis) near Santo Domingo, Dominican Republic.

Next day, and after many hours of bureaucratic roundabouts, we set out west, across the mountains of the Sierra de Neiba and on to the Valley of Neiba on our way to Barahona. We were looking for a fisherman town called Los Patos, and a set of caves perched in the mountains of Barahona. Several other scientists had marked this location as a site of interest for bat researchers (Miller, 1916-1929), and we were following their footsteps.



End hills of the Sierra Baoruco, in the small fishermen town of Los Patos, near Barahona.

The caves 1 and 2 of Los Patos are almost vertical, inside the belly and atop the hills from which the ocean is visible. The rocks there are limestone conglomerates, which with time and erosion rolled down and covered the beach in a thick blanket of polished pebbles. Such surface made our sleep there somewhat uncomfortable, but the view was spectacular.


Los Patos beach, near Barahona, looking towards the Caribbean Sea.



View from the mouth of Cueva de Los Patos 1, over looking the Caribbean Sea.

Inside the caves laid examples of the ancient fauna, represented by delicate fossils. The floors had guano and the walls had bats. The species we observed included ghost-faced bats Mormoops blainvillei, large fruit-eating bats Brachyphylla nana, Artibeus jamaicensis, the pollen and nectar eaters Monophyllus redmani and Phyllonycteris obtusa, plus large-eared insectivorous Macrotus waterhousei. The bat and bird faunas were exquisitely diverse.

Cueva de Los Patos 1-2. Roost of large fruit bats Brachyphylla nana (pumila) and Phyllonycteris obtusa.

Large-eared bat Macrotus waterhousei. This is the large subspecies waterhousei, which lives on Hispaniola.



However, the natalids, Natalus major and Chilonatalus micropus, the goal of the expedition almost, eluded us. Our single Ch. micropus was caught late one night, as we were putting away the mist net. Just then came this low, butterfly-like, flying bat into the net. So far, this remains the only reported Ch. micropus roost site on Hispaniola (Tejedor, 2011: 35).

Chilonatalus micropus from Los Patos Cave 2

Under the chilly effect of the mountains, we headed back to Santo Domingo. On our way through the valley of the Cordillera Central towards the south, we stopped at Bani, birthplace of Maximo Gomez (1836-1905). Gomez was a brave and dedicated General of Cuban wars for independence between 1868 and 1898, and the later Cuban-Spanish-American War. Cuban history values the great contribution from this Dominican generalissimo (see fig. below).

General Maximo Gomez, early 1900s. From Library of American History, Vol.VII.

The roads crossing through the central valley of the Cordillera Oriental to Sabana del Mar, on the south coast of Samana Bay, were very deteriorated or non-existing, rough, and dangerous. But these were filled with interesting flora and fauna that we stopped to observe.

Hispaniolan giant Tarantula Phormictopus cancerides

One of our first encounters was this Hispaniolan giant Tarantula (Phormictopus cancerides), and one or two Ashy-faced owls (Tyto glaucops). While asking for directions in the town of Sabana de la Mar, we spotted a large bat flying around a light post in the main central park. We parked to take a closer look. It must have been nearly 12 am, and we were dead tired, but stunned to see a large bulldog fisher bat (Noctilio leporinus) apparently eating insects attracted by the light of the lamp post!

Noctilio leporinus on the central plaza of Sabana del Mar. The white dots are likely insects captured in the glare.

Finally, that night we arrived a natural reserve station on the Haitises Park. The Haitises are a conglomerate of natural wonders. It has a high diversity both in fauna and flora, and interesting  formations called "mogotes" or in this case known as "haitises". These are conic karst hills, like the mogotes of my previous post on Pinar del Rio, western Cuba. These, however, are formed on younger limestone, smaller, and covered with more vegetation, but similarly impressive.



Conic karts, limestone formation of the Haitises as we saw them from our boat.
Courtesy and Copyright of Adrian Tejedor.

Conic karts, limestone formation of the Haitises as we saw them from our boat.
Courtesy and Copyright of Adrian Tejedor.

This variation of karst formation or karst geomorphology (as in geological manifestations of the terrain), was formed by dissolution of the limestone over time. In the Caribbean islands similar karstic formations are present, but most profusely in Cuba, Jamaica, this region of Hispaniola, and in Puerto Rico. However, they are all distinct in their level of maturity. The oldest and thus more mature are those of Pinar del Rio in Cuba, whereas those of Hispaniola and Puerto Rico are formed on most recent rocks.

Massive limestone haitises in the bay of Samana. One can almost imagine
how C. Columbus saw the Tainos right on these beaches he was there.

To get to these rounded hills, which from afar looked like elephants half submerged in the waters of the San Lorenzo bay, we had to ride on a small boat. This boat took our party along the crannies and crevices between the massive rock domes of the Haitises. We were looking for the intricate cave systems that honeycomb these formations, so important to our research since they were to host the fauna we sought.



Railroad Cave appears from within the drowned elephants that are the karst hills of the Haitises.

One of these caves was Cueva de la Linea, or Railroad Cave, known to us from the early research of William M. Gabb, William L. Abbott, Gerrit S. Miller,  and later that of Krieger (1928-29).

Railroad Cave is near an abandoned railway track near the bay of San Lorenzo. The area is surrounded by crescent sandy beaches, marshes, and caves. The most notable caves being  Simmons's Cave, Boca del Infierno cave (the Mouth of Hell), and Railroad cave, which is known locally as Cueva del Templo (cave of the temple).

William M. Gabb explored caves around this area between 1869 and 1871, finding extensive evidence of pre-Columbian inhabiting. Exploration continued calling the attention of William L. Abbott who explored several of these caves, including Railroad cave in 1883 and then in 1916. It was the former which enticed the National Museum to send Gerrit S. Miller later that year, and then again in 1928 with H. Krieger. It was this last party which carried out serious archeological and paleontological research there (1929).

Sunlit Cueva de la Linea or Railroad cave in Samana Bay.

The Ciguayan tainos inhabited these beach caves, and their former presence is felt by their many shell heaps (Strombus pugilis) at their entrances and the unforgettable cave art in their anterooms. The shell heaps also include the bones of the animals the Tainos used for food, such as hutias, manatees, conchs and fish, and are generally called kitchen middens in the archeological jargon. One can't help but imagine what Columbus saw when he visited the bay of Samana to observe an eclipse of the moon in 1492. Then, the natives lived on the north shore of the bay.

No doubt we enjoyed this particular cave much. The pictographs and petroglyphs, like those the figure below, carved into the cave rock, depicted faces, handprints, and sketches of animals like egrets, dogs, sharks, and others. Moreover, there were large bat colonies in very hot rooms separated by small water intrusions, deep into the cave system.

Ciguayan Taino petroglyph at Cueva de la Linea, Samana.
One of the many human artistic representations of the area.

Moving inside these hot rooms was uncomfortable because the extreme temperature and smell of bat urine made breathing difficult. Often we had to stop and hold on to the wet walls to catch our breath before moving on forward. In the center of these rooms, there were accumulations of bat excrements and all kind of invertebrate fauna that feeds on deceased bats and the guano on the floor. The same guano that is often mined as a natural fertilizer.



Natalus major in its roost, Cueva de Cristian, Hato Mayor.



A colony of the sought after Natalus major, inside a well-vented room Cueva de Cristian, Hato Mayor.

But what was the purpose of all this?

Fieldwork is not an easy task and is well accompanied by multiple difficulties that researchers must endure reaching their goals. From sleeping on cave floors infested with ticks and roaches, to having no food or commodities, to being attacked by the native fauna (people included).

It is sad that much destruction occurs well within the boundaries of several national parks and other areas. Illegal burning, cutting, and cave guano extraction threatens and disturbs the natural fauna. This includes the nests of the Palm crows, the endangered Ridgway's hawk, Hispaniola amazon parrots, natalid bats, and a myriad of plant life; living organisms in general, but especially those that are endangered or vulnerable already. Many of the well-forested areas are cleared for avocado, coconut, and plantain plantations or tourism. Therefore, it is important that we document the existing flora and fauna so that we can establish sensitive plans of protection, so that the wonderful areas are not lost to posterity, and that other may enjoy its natural wonders in the same way that we have.

Samana peninsula and San Lorenzo bay seen from atop one of the Haitises.
A scenery reminder of the natural wonders that must be protected from complete human destruction.
Once these are gone, they are gone forever.

The experience of research, not just traveling to exotic places to see interesting organisms, but with the hope of discovering something new, is very rewarding. In the end, our efforts are towards a better understanding of the natural environment that surrounds us all.

We think these environments and their organisms are worth preserving, but one blog cannot capture the natural complexity and beauty of these amazing islands. The world would surely be a dull place without these magnificent ecosystems. We should strive to protect them, instead of destroying them.


Cited Literature

Krieger, H. W. 1929. Archeological and historical investigations in Samana, Dominican Republic. US National Museum Bulletin, 147.

Tejedor, Adrian. 2011. Systematics of funnel-eared bats (Chiroptera: Natalidae). Bulletin of the American Museum of Natural History, 353.

Cave Fieldwork in Cuba: 2002 - 2004

The early years of the new millennium were very productive for my research in Cuba. Our explorations, between 1995 and 2002, turned up very interesting fossils, representing rare and extinct faunas. The fruits of these explorations included two new records for Cuba, pertaining to two of the Caribbean's rarest fossil bats: the common vampire bat Desmodus rotundus and Peter's ghost-faced bat Mormoops megalophylla (articles can be downloaded here).

Fig. 1: An articulated skeleton of Mormoops blainvillei at Cueva de la Pluma (Cave of the Feather) in northern Matanzas, Cuba. This species is almost identical but smaller, than Mormoops megalophylla of the continent.

Our fieldwork began in the foothills of the Alturas Habana-Matanzas, a chain of lowland limestone hills that run W to E on the northern coast of Cuba. Uplift and exposure of these limestones during the last 5 million years has given origin to a vast cave region laden with fossil deposits, dating to the last glacial and inter-glacial period.

Fig. 2: Palenque Hill, a lowland limestone hill of the Havana-Matanzas range.
This is a karstic relict of uplifted Miocene seafloor that surrounded this region.

Within this region, we focused on a 200-meter high hill named Palenque (fig. 2-3). The hills of Palenque served as a shelter to runaway aborigines and slaves throughout the colonial period. A "palenque" is the name given to such a hideout. We selected this region mainly because of unstudied caves discovered in the 1980s, with an interesting biotic richness and endemism, characteristics that had attracted two of Cuba's foremost naturalists Carlos de la Torre and Johannes Gundlach a century before. They visited Palenque's hillside in search of unique mollusks, mammals, and birds.

Fig. 3: Karstic vegetation on the escarpment of Palenque hill, at about 180 meters above modern sea level.

A secondary but well preserved tropical semideciduous forest covers Palenque (fig. 3) with a flora that includes several kinds of trees including oaks and mahogany (Quercus and Swietenia spp.), gumbo limbo (Bursera simaruba), Royal Palms (Roystonea regia), plus other Thrinax-like palms in the upper levels. There is even coffee (Coffea arabica) brought into some caves by large fruit bats (Artibeus and Brachyphylla) and birds. Most of the lower level vegetation that extends to the agricultural savanna at the foothills include several kinds of grass, the poison ivy, locally known as Guao (Comocladia dodonea) and the toxic Chichicate (Urtica dioica).

Fig. 4. Main sinkhole of Nesfonte's cave, one of the largest cave of the Palenque, and probably the most extensive.
This is the main owl repository where most of the owl pellets have been accumulating for more than 2000 years. 

Fig. 5: The Cuban tarantula Phormictopus cubensis,  part of the penumbra cave fauna.

These fossil deposits originated from animal and plant remains mixed in with soil dragged into the caves by rainwater. Other were brought in by raptors, for example in vomitus, a form of hairball called pellet. Raptors, owls, and hawks do not fully digest bone and hair. Instead, they regurgitate them in the form of such pellets which accumulate in their cave roosts by the thousands (fig. 1, 6-7; note the round and brown pellet on the right of fig. 6). Because these raptors reuse the same roost areas from generation to generation their nests can include several thousand years of prey/pellet records. Even though their diet is selective, meaning they pick and choose from what prey is available in nature, theses pellets can provide a good record of the local fauna. The caves of Palenque provide an excellent record of that.

Fig. 6: A common vampire bat Desmodus rotundus skull in situ at the moment of discovery.
See the skulls of the brown bat Eptesicus fuscus (on far left) and large Cuban fruit bat Artibeus jamaicensis
skull (on upper right). A fresh owl pellet is next to the vampire bat skull,
 and some fresh material remains inside its braincase, testifying to its freshness.

The first serious discovery came from a newly discovered room in Cueva de la Caja or the Cave of the Box (fig. 3-4), also known as Cueva de Los Nesofontes or Nesofonte's cave, named so for the huge accumulation of Nesophontes fossil remains (more on this curious species on a future post). As archaeologist R. Viera and I explored a section of the cave, we accidentally stumbled upon a small mound on the floor of a newly discovered side room that had gone unnoticed on previous expeditions in 1994 and 2002 (Viera and Orihuela, 2006). In it, there were plant seeds scattered in a mound on the cave's floor and a vampire bat skull right on top like a cherry on the cream (fig. 6-8).

Figure 7: The vampire bat deposit as it looked before excavation on 22 December 2003.

What an interesting and unexpected discovery! At that time, vampire bats had been discovered only three times before, all exclusively from Cuba (Mayo and Woloszyn, 1974; Silva, 1979). As of today, vampire bats have not been discovered anywhere else in the greater Caribbean.

Desmodus rotundus is endemic to the New World neotropics, where it is well distributed from Mexico all the way down to northern Argentina, but in the past, that distribution included Florida, Cuba. The first fossil record came from Cueva Lamas in Havana. This fossil was discovered by Cuba's foremost paleontologist at the time Oscar Arredondo, who then sent the specimen to Dr. Karl Koopman, chiropterologist (=bat-ologist) who identified it as a common vampire bat Desmodus rotundus, identical to the continental species. In the following years, other researchers found two more specimens from which a new endemic subspecies, Desmodus rotundus puntajudensis, was erected (Woloszyn and Mayo, 1974; Jimenez et al., 2005). Later, the Cuban paleontologist William Suarez (Suarez, 2005) raised the species to the rank of full species, which made it a full Cuban endemic.

Fig. 8: Common vampire bat (Desmodus rotundus) after it's discovery at Nesophonte's cave.

However, the study of our well-preserved specimen (fig. 8-9) showed it was not referable to the new endemic form. Instead, our analysis of the skull characteristics of the continental form D. rotundus versus that of the endemic D. puntajudensis revealed that the Cuban fossils fell within D. rotundus, and did not represent a new form (Orihuela, 2011). This agreed with Koopman's original identification of the first specimen. As it seems, the Cuban fossil vamps are the same as the continental species, which may have arrived at the island during the last 10,000 years or less.

Fig. 9. Three of the four known Desmodus rotundus skulls from the Cuban Quaternary fossil record.
 Our specimen is on the far right. Line illustration from Orihuela, 2011 (copyrighted). 

An endemic form would suggest long isolation for such evolution to occur. And the presence of both, an endemic species (or subspecies) and the continental form, seems unlikely out of so poor of a record. So far, the fossil Desmodus specimens are associated with ages less than 8000 years before the present, and thus within our own current inter-glacial period, but none date to the late glacial period as was originally considered. In fact, we do not know if they were true relics of the past glacial fauna.

Osvaldo Jimenez Vazquez and his colleagues discovered vampire specimens from a cave deposit in southern Mayabeque province (formerly Havana province), providing the first radiocarbon date associated with vampire bat fossils in Cuba. Their remains dated, although indirectly, about 7000 years before the present (Jimenez et al., 2005). Our specimen posed a more complex problem and brought about new questions. They were dated, indirectly as well, between the 1950s to the 1990s (Viera and Orihuela, 2006; Orihuela, 2012; Orihuela et al. in prep.). Unfortunately, neither Cuban vampire fossil has been dated directly due to the rarity of the fossils.

Fig. 10: Pteronotus parnelli parnelli (Mormoopidae) from a transient local population at Nesophonte's cave.

What is sure is that there were vampire bats in Cuba until very recently, even if just accidentally. Our research using maximum entropy software such as Maxent, DIVA and R, employed to predict and model which bioclimate variables could have limited bat distribution, and if any of the variables could have lead to their extinction, suggests that the climate of the last 7000 years, up to our modern climate, is still appropriate to sustain many currently extinct Cuban bats including the vampire bats (work in prep.). Their extinction is considered concomitant to the extinction of the large sloths (like those I mentioned in my previous post here), which were likely part of their diet. Now we know that these sloths did not disappear during the late glacial epoch either, but instead during the last 5000 years, which is within our current inter-glacial.

So what does this mean? Where vampire bats accidentally present in the Cuban fossil record? If so, why are their fossils so rare considering how easily adaptable and reproducible the species is on the mainland? Are Desmodus fossils present in other Antillean fossil records? These are questions our current research is hoping to resolve. Stay tuned!

I want to take this opportunity to thank those that have helped on this research. First, I am eternally thankful to all those that participated in this expedition and those colleagues that helped with the analysis, their encouragement and help. These are archeologist Ricardo Viera, geographer Leonel Perez-Orozco, biologist Adrian Tejedor, Candido Santana and Joel Monzon, and many other more friends and colleagues that have made this research possible. These expeditions would have been impossible without their enthusiastic support.

This post is dedicated to the memory of Johannes Gundlach, whose birthday we celebrate today.

References

Atlas Nacional de Cuba 1969-1985.

Arredondo, O. 1958. El Vampiro Cubano. Scout 10:6-7.

Jiménez, O., M. M. Condis, and E. García. 2005. Vertebrados post-glaciales en un residuario fósil de Tyto alba scopoli (Aves: Tytonidae) en el occidente de Cuba. Revista Mexicana de Mastozoología, 9:84-111.

Koopman, K.F. 1958. A fossil vampire bat from Cuba. Breviora 90:1-4.

Silva-Taboada, G. 1979. Los Murciélagos de Cuba. Editorial Academia, La Habana. 424pp.

Orihuela, J. 2011. Skull variation of the vampire bat Desmodus rotundus (Chiroptera: Phyllostomidae): Taxonomic implications for the Cuban fossil vampire bat Desmodus puntajudensis. Chiroptera Neotropical 17(1): 963-976.

Orihuela, J. 2012. Late Holocene fauna from a cave deposit in Western Cuba: post-Columbian occurrence of the vampire bat Desmodus rotundus (Phyllostomidae: Desmodontinae). Caribbean Journal of Science, 46 (2): 297-313.

Orihuela, J., and A. Tejedor. 2012. Peter's ghost-faced bat Mormoops megalophylla (Chiroptera: Mormoopidae) from a pre-Columbian archaeological deposit in Cuba. Acta Chiropterologica 14(1): 63-72.

Orihuela, J., R. Viera, and L. Vinola. 2017. New bat records based on modern and fossil remains from the province of Matanzas, Cuba.

Suárez, W. 2005. Taxonomic Status of the Cuban Vampire Bat (Chiroptera: Phyllostomidae: Desmodontinae: Desmodus). Caribbean Journal of Science 41 (4):761-767.

Woloszyn, B.W., and N.A. Mayo. 1974. Postglacial remains of a vampire bat (Chiroptera: Desmodus) from Cuba. Acta Zool.Cracoviensia 19:253-265.