Our new paper on Greater Antillean land mammal extinctions is published!

With great pleasure (and relief after nearly a decade of research) here I announce the publication of our paper Assessing the role of humans in Greater Antillean land vertebrate extinctions: New insights from Cuba on the journal Quaternary Science Review. Also, here is a link for its free download to all parties interested, in the next fifty days. 

I take this opportunity to thank all the coauthors of this paper, Ángelo Soto Centeno, Lázaro W. Viñola, Osvaldo Jiménez, Odlanyer Hernández de Lara, Logel Lorenzo, and Alexis Mychajliw – all respected specialists in their fields – for their significant contribution and participation in making this dream come true. Thank you. 

Here is a brief abstract: 

The Caribbean archipelago is a hotspot of biodiversity characterized by a high rate of extinction. Recent studies have examined these losses, but the causes of the Antillean Late Quaternary vertebrate extinctions, and especially the role of humans, are still unclear. Previous results provide support for climate-related and human-induced extinctions, but often downplaying other complex bio-ecological factors that are difficult to model or to detect from the fossil and archaeological record. Here, we discuss Caribbean vertebrate extinctions and the potential role of humans derived from new and existing fossil and archaeological data from Cuba. Our results indicate that losses of Cuba’s native fauna occurred in waves: one during the late Pleistocene and early Holocene, a second during the middle Holocene, and a third one during the last 2 thousand years, combining the arrival of agroceramists and later of Europeans. The coexistence of now-extinct species with multiple cultural groups in Cuba for over 4 thousand years implies that Cuban indigenous non-ceramic cultures exerted far fewer extinction pressures to native fauna than the later agroceramists and Europeans that followed. This suggests a determinant value to increased technological sophistication and demographics as plausible effective extinction drivers. Beyond looking at dates of first human arrival alone, future studies should also consider cultural diversity with attention to different bioecological factors that influence these biodiversity changes. 

Highlights

Cuban land mammal extinctions occurred in several waves after the middle Holocene, most intensively during the last 2000 thousand years

Cuba lost nearly half of its land mammal fauna during the late Amerindian subinterval (< 1500 thousand years)

Most important extinction episodes occurred after the arrival of agroceramist cultures, and later, Europeans

Cultural diversity, demographics, technological sophistication, and naturally occurring factors must be considered in human-induced extinction models

Future extinction models must consider the complex and concomitant combination of bioecological and climatological factors

[in Spanish]

Con gran gusto – y después de casi una década de investigación – anunciamos la publicación de nuestro artículo “Assessing the role of humans in Greater Antillean land vertebrate extinctions: New insights from Cuba” (Evaluación del papel de los seres humanos en la extinción de vertebrados terrestres de las Grandes Antillas: nuevas perspectivas desde Cuba) en la prestigiosa revista Quaternary Science Review. Aprovechen y compártanlo con los amigos y colegas que les interesen estos temas. Tomo este momento para agradecerle a los coautores Ángelo Soto Centeno, Lázaro W. Viñola, Osvaldo Jiménez, Odlanyer Hernández de Lara, Logel Lorenzo y Alexis Mychajliw por su arduo trabajo y participación en hacer este sueño realidad.

Resumen del trabajo:

El archipiélago caribeño es un “punto caliente” de biodiversidad caracterizado por una alta tasa de extinción. Los estudios recientes han examinado estas pérdidas, pero las causas de las extinciones de vertebrados del Cuaternario tardío de las Antillas, y especialmente el rol de los seres humanos, aún no están claros. Los resultados anteriores brindan apoyo a las extinciones inducidas por el hombre, pero a menudo se minimiza otros factores bioecológicos que son difíciles de modelar o detectar a partir del registro arqueológico o fósil. Aquí discutimos las extinciones de vertebrados en las Antillas Mayores y el papel que han jugado los humanos en las extinciones más recientes desde la perspectiva de datos arqueológicos y paleontológicos de Cuba. Nuestros resultados apoyan la hipótesis de que las pérdidas de la fauna nativa de Cuba ocurrieron en ondas: una durante el Pleistoceno tardío y el Holoceno temprano, una segunda durante el Holoceno medio, y una tercera durante los últimos 2 mil años. Estos dos milenios resultan ser los más importantes, combinando la llegada de los agroceramistas y luego de los europeos como importantes golpes a la fauna. La coexistencia de especies ahora extintas con múltiples grupos culturales en Cuba por más de 4 mil años implica que las culturas indígenas no-ceramistas ejercieron menos presiones de extinción sobre la fauna nativa que las agroceramistas y los europeos que siguieron. Esto sugiere un valor determinante para la sofisticación tecnológica y la demografía vías importantes de extinción. Más allá de mirar las fechas de la primera llegada humana, los estudios futuros también deben considerar la diversidad cultural y atención a diferentes factores bioecológicos que influyen en los cambios de biodiversidad.

Recommended citation:

Orihuela, J., Viñola, L.W., Jiménez Vázquez, O., Mychajliw, A., Hernández de Lara, O., Lorenzo, L. and, Soto-Centeno, J. A. (2020a). Assessing the role of humans on Greater Antillean land vertebrate extinctions: new insights from Cuba. Quaternary Science Reviews, 249: https://doi.org/10.1016/j.quascirev.2020.106597

The Cuban Pallid bat Antrozous koopmani

Cuba has an endemic Pallid bat:  Antrozous koopmani, a species named in honor of the great bat zoologist, the late Karl Koopman. I posted a similar post on the Cubabat Facebook page, and the post was so successful, that I thought may be interesting in reposting an edited version here.

There are currently two species of Pallid bats, Antrozous pallidus and Antrozous koopmani, of the Vespertilionidae family. Antrozous pallidus has a range that extends from southern British Columbia, south to the coastal states east of Kansas, and down to Queretaro, Mexico with an insular population on Maria Magdalena Island, Nayarit. And A. koopmani an endemic species to the island of Cuba.

Drawing of one of Ramsden's Oriente specimens figured in Dr. G. Silva's
masterpiece Los Murcielagos de Cuba (1979). 

This insular and distinct population of Antrozous was not detected until 1920 when C. T. Ramsden acquires two specimens in Oriente province, eastern Cuba. One was caught at Caney, another from Guantanamo. This feat was not again repeated until the Drs. Gilberto Silva-Taboada and Karl Koopman captured another living specimen in 1956, on the foothill forests of Pan de Guajaibon in Pinar del Rio, western Cuba. The Ramsden specimens are the only known skin-preserved-specimens known to date.

Antrozous skull and dentary: Antrozous koopmani on the left, and
Antrozous pallidus (AMNH 2125) on the right.
The A. koopmani is from fossil site at Palenque Hill, Mayabeqye, Cuba.

In the upcoming years, however, several skeletal remains were recovered from fresh owl pellets in several caves in western Cuba. Robert T. Orr and Gilberto Silva used this material in the official description of the Cuban pallid bat Antrozous koopmani as an endemic species of the island of Cuba in 1960.

Today, the species has been reported from several fossil deposits in most provinces of western Cuba: Pinar del Rio, Mayabeque, Matanzas, and Sancti Spiritus. Which suggests that the species had a much wider distribution on the island during the Recent past, that could have lasted until several hundred years ago.

Antrozous koopmani is today Cuba’s rarest bat. It has not been captured alive again, at least with certainty, since 1956, and is now presumed extinct.

Please  visit Proyecto Cubabat's page to see the original post and many others there. 

Answer to Fossil Trivia III: The Fossil Bat in the Stone

Well, that was a tough one. Not many saw the fossil right away, but one learns in paleontology, as in any other field, with experience. It is with time and some getting use to, that one can beging to make out shapes and structures in rocks, that normally, would not be obvious to the observer.

In our last case, that was, I guess, a bit unfair. The fossil visible on the rock is that of a bat, a 30 million year old specimen from the Green River formation, and presently on the Vertebrate Paleontology collection of the American Museum of Natural History (AMNH, in NY).

The Green River is known for its superbly preserved fossils, among which figure bats. The low oxygen conditions, of what seemed to be a shallow lagoon, lake or estuary environment promoted preservation of any organism that fell in its waters, and settled at the muddy bottom. This bat represents one of those such events.

Archaeopterix specimen (cast) on exhibit at the AMNH.
Note the tail feather impressions. 

Other similar deposits exist at the Messel Pit fossil site, in Germany, where 40 million year old animals are remarcably preserved, along with embrios, stomach content, hair, and sometimes even color. The Solnhofen limestone deposits, also in Germany is another remarkable example. The first Archaeopteryx, the first feathered dinosaur discovered, A. lithographica, was found in the fine-grained limestones of Solnhofen's quarries in 1861. It was called lithographica, because, originally, the fine limestone extrated at those quarries were coveted for the printing industry. The fine-grained feature of Solnhofen's limestone allowed a high degree of detail of the engravings marked upon its printing blocks.

Lithographic limestone block used for printing. Taken from Pintrest. 

Fossil Matter Trivia II: Can you identify this fossil?

In our previous trivia, I posted the image of an ammonite. This was a group of cephalopod mollusks that went extinct in the late Cretaceous along with the dinosaurs. These were marine predators, some of them growing a few meters in diameter.

The specimen from the previous post is a Perisphinctes cubensis from the middle Jurassic black limestones and shales of Pinar del Rio, western Cuba. These fossils can be found on river rocks and rock walls.

But this post will be harder. Can you identify this fossil?

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!

The Bats of Cuba

Bats inhabit nearly every landmass on the planet, with the exception of the arctic and Antarctica, reaching their maximum diversity in the tropics. Current tallies rank bat diversity at 202 genera and over a thousand species! (Simmons, 2005).

Bats are highly specialized mammals. Not only can they truly fly (meaning powered flight), but they use echolocation to navigate and detect prey while in flight. Echolocation is a way of navigation by echoes, which bats and other mammals like dolphins emit to sense their environment and find food. Some birds, like the oilbird Steatornis caripensis, uses echolocation, but at a level we can hear. In addition to their characteristic webbed wings, their eyesight is better in the dark than ours, demystifying their bad reputation of poor eyesight.

Leach's single leaf-nosed bat Monophyllus redmani (subspecies clinedaphus) a
pollen-nectar feeder Microchiropteran common to the Greater Antilles

Chiropterans are divided into two groups: the small echolocating bats or Microchiroptera, and the non-echolocating and much larger Megachiroptera. The giant bats portrayed in movies often represent the large fruit-eating Megachiropterans. They are not all vampire bats!    

Ecologically, bats play keystone roles in the consumption of insects, distribution of plant seeds, and pollination of plants. Other bats are carnivores, also helping maintain the ecological balance of the ecosystems they inhabit.

Bats are the most diverse mammals of the Cuban archipelago representing over 75% of Cuba's mammalian fauna! Of the 34 species of mammals recorded in Cuba, 26 are bats. Currently, these 26 species are classified into six main groups including the nose-leaf bats (phyllostomids), the funnel-eared bats (natalids), the fisher bulldog bat (noctilionid), the free-tailed mastiffs (molossids), the insectivorous ghost-faced bats (mormoopids) and the vespertilionids. The majority are well distributed within the main island of Cuba, the isle of Pines, and a few of the thousand keys that make up the archipelago (Silva, 1983; Mancina, 2011, 2012).

Waterhouse's leaf-nosed bat Macrotus waterhousei
and the Cuban fig-eating bat Phyllops falcactus digitally
drawn by biologist and bat specialist Dr. Adrian Tejedor from
field sketches of specimens captured in Pinar del Rio.

The Cuban archipelago is the largest of the Antillean islands. It is located in the Antillean subzone of the Neotropics where it enjoys a warm weather and abundant rainfall most of the year. The geological formation of the Caribbean islands provided an intricate and complex mosaic of calcium-rich rocks, such as limestone, so important to cave formation and varied soils. Altogether, these variables give rise to lush vegetation, which supported by the warm climate, incites Cuba's biodiversity, especially of bats.

The Cuban archipelago, in the Caribbean basin, as seen in Google Earth.

Geologically, Cuba has been available for bat colonization since the latest Eocene (MacPhee and Iturralde, 1994). This means that there have been somewhat permanent group of islands where Cuba is located today, at least for the last 35 million years, giving ample time for bats to reach it and evolve there.
In the Pliocene - 5 million years ago - the islands had their largest subaerial extent, and thus their largest landmass increase to date (Iturralde, 2010). This was followed by the multiple landmass fluctuations experienced during the Quaternary glaciations. During the last 800,000 years, the Cuban archipelago increased and decreased in size at least 20 times, with the glacial periods decreasing sea levels, and the interglacials increasing it. This is within a range of ~20 meters from the modern standard. In effect, this had substantial effects on the formation of karstic features, such as caves, that serve some bats as roosts, affected plant distribution, and likely also the distribution and evolution of bats in the island. But most importantly, it likely played a role on the total number of species the archipelago could sustain.

A lone Cuba fruit bat Artibeus jamaicensis parvipes in roost in Cueva Ambrosio,
Varadero, alongside Amerindian cave pictographs.

During the Quaternary, Cuba, and the Bahamas acted like a single archipelago. Today that archipelago is mostly drowned by higher sea levels. Increased sea levels likely flooded potential cave roosts affecting strict cave dwellers. There are bats that have adapted not only to live in caves but also preferring hotter environments within cave systems. Caves with hotter than normal chambers are called "hot caves" because the temperature in some of its rooms increases to above 40 degrees C with a relative humidity above 90 percent. These hot environment form in chambers with restricted access, in which large colonies of the bats roost. Their body heat, perspiration, urine and droppings, all within a very restricted and poorly ventilated cave room results in the abnormal increase in room temperature and humidity. Hot cave bats include the pollen-eater Phyllonycteris poeyi and Pteronotus quadridens.

The changes in world climate during the last 2 million years, or that Quaternary epoch that we've been referring to, brought changes in rainfall, temperatures, and potential land size, therefore potentially affecting different species. However, bats were not significantly culled by the Quaternary climatic fluctuations, as far as we can tell today from the fossil record, in comparison to other mammals groups, like monkeys and sloths, which disappeared completely. Cuba lost only three species during the last glacial maximum, ~18,000 years ago, as indicated by the fossil record of Cueva El Abron, in Pinar del Rio (see Suarez and Diaz-Franco, 2003; Balseiro, 2011). Others survived until a thousand years ago or less (Orihuela, 2012; Orihuela and Tejedor, 2012, Orihuela et al, in prep).

The greater bulldog bat Noctilio leporinus. This is Cuba's largest bat. It feeds mostly on fish,
but it has been observed eating insects near street lamps.

Cuban bats, like most bats, inhabit most ecosystems where they have evolved adaptations to many forms of feeding. There are bats that eat insects, seeds, fruits, nectar, pollen, and some that feed only on blood, such as the infamous vampire bats, or fish - as the Noctilio leporinus above. In the past, there were vampire bats in Cuba. Vampire bats are locally extinct in Cuba today, but their fossil remains suggest their presence on the main island up to several hundred years ago! (Suarez, 2005; Orihuela, 2011, 2012) (see my previous post on vampire bats here).

Thomas Horsfield on the right and John Edward Grey on the left.
Both men dedicated time to collecting and describing the first Cuban bats during the XIX century.

We owe the first published account on Cuban bats to Thomas Horsfield, who sent a letter to the prestigious Zoological Journal in 1828 while he resided in Cuba. Horsfield and the British naturalist William Sharp MacLeay sent specimens to the British museum. These specimens allowed John Edward Grey to properly describe the first species in 1840 in his article "Description of some Chiroptera discovered in Cuba", published in the Annals of the Natural History, volume IV (Silva, 1983).

The Antillean fruit-eating bat Brachyphylla cavernarum and the big free-tailed bat Nyctinomops macrotis
lithographs from Grey's first description of Cuban bats. These are also the first graphic depiction of any Cuban bat.
Lithographs made by the french J. Basire in 1839. B.cavernarum here is likely B. nana but nana was not properly
described until the early XX century.

Since then, and no doubt thanks to the efforts of many naturalists such as Johannes Gundlach in the XIX century and Gilberto Silva-Taboada of the XX, among others, the knowledge on the natural history of Cuban bats grew steadily. Their research quickly demonstrated the diversity of the Cuban bat fauna. There are more species in Cuba, with species representing most of the known New World groups, than in all the North American continent!

  Parnell's mustached bat Pteronotus parnelli from Nesofontes Cave near Matanza, Cuba.

The Cuban bat fauna is surrounded in interesting stories of accidental discoveries and rediscoveries. Such as it happened to the two bat specialists, the Cuban mastozoologist G. Silva-Taboada and Karl Koopman of the American Musem of Natural History (NY) while mistnetting bats in the Pan de Guajaibon, Pinar del Rio, in the 1950s. There they caught a living Cuban pallid bat Antrozous koopmani, the only one ever caught alive for decades. This is a species similar to the pallid bat Antrozous pallidus of the arid western U.S. The Cuban pallid was previously known from a handful of isolated skulls and two specimens preserved in spirits collected in the early decades of the XX century. The feat his yet to be repeated. Antrozous koopmani is today the rarest of Cuban bats, and is presumed nearly extinct (Mancina, 2012).

The list is followed by the greater funnel-eared bat Natalus primus, a critically endangered species known alive only from the single location of La Barca cave in Guanahacabibes, extreme western Cuba. There it was rediscovered by biologist Adrian Tejedor in 1991. Tejedor has written several interesting articles and a monograph on the rare and interesting funnel-eared bats (Natalids) of Cuba and the Caribbean (Tejedor, 2011 and citations therein). Cuba has other two funnel-eared bats. One of them, Nyctiellus lepidus, is one of the smallest bats in the world, known in Cuba as the "butterfly bat". The other, Chilonatalus macer, is similar to the Natalus major on the right of the image below but smaller.  Cuban natalids are all are endemic.

Left: Pteronotus quadridens from Hispaniola. Right: Hispaniolan funnel-eared bat
Natalus major from Cueva de Los Patos, also in Hispaniola.
These are small insectivores that live only in caves.

Other interesting records include Myotis sodalis, likely an errant from Florida found mummified by G. Silva in the city of Havana during the winter of 1966 (Silva, 1983). Eumops perotis, likely a vagrant or erroneous record dating back to 1839, a tree-dwelling Lasiurus insularis found by Ricardo Viera on the banks of the Yumuri River, Matanzas, in 2004, and our rediscovery of Desmodus rotundus in 2003; the fifth vampire bat fossil record from Cuba, among other informative, but isolated discoveries (Silva, 1983; Viera, 2004; Orihuela, 2010; Orihuela et al, in prep.). To this adds an array of new species and new deposits rich in bat fossils (Silva, 1974; Suarez, 2005; Suarez and Diaz-Franco, 2003; Mancina and Garcia, 2005; Jimenez et al., 2005; Balseriro, 2011; Orihuela, 2012).

Most of these latter species, however, are either accidental records, critically endangered, extirpated or extinct today. In addition to the extant 26 species, there are 8 disappeared species for a total of 34 known to have existed in Cuba at least during the last 20,000 years. The complex account of Cuban bat extinctions is reserved for an upcoming post; a topic most interesting to me, and the focus of most of my research.

Stay tuned!

The greater Cuban funnel-eared bat Natalus primus, severely endangered
and extant only in Cueva la Barca, Guanacahabibes, western Cuba.
Digital painting by, and copyright of Adrian Tejedor.

References

There are more citations, especially on area restriction, bat habitat, feeding, and climatic changes of the Quaternary that, if included, would have made this post a bit more tedious. I think, however, that the references below, in addition to the work of Angel Soto-Centeno, David Steadman, Danny Rojas and Liliana Davalos, will provide a good background for those interested in keeping up with this ever-growing body of literature.

Balseriro, F. 2011. Los murcielagos extinctos. pp: 171-177 en Borroto-Paez, R. y C. A. Mancina (eds) Mamiferos en Cuba. UPC print, Vaasa

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.

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

Gonzalez, Alonso, et al. 2012. Libro Rojo de los Vertebrados de Cuba. Editorial Academia, La Habana. See "Mamiferos" pp:269-291 by mastozoologist Carlos Mancina.

Mancina, C. A., and L. Garcia. 2005. New genus and specis of fossil bat (Chiroptera:Phyllostomidae) from Cuba. Caribbean Journal of Science, 41: 22-27.

Mancina, C. A. 2011. Introduccion a los murcielagos pp: 123-133 en Borroto-Paez, R. y C. A. Mancina (eds) Mamiferos en Cuba. UPC print, Vaasa

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.  

Silva Taboada, G. 1974. Fossil Chiroptera from cave deposits in Central Cuba, with a description of two new species, and the first record of Mormoops megalophylla. Acta Zoologica Cracoviensia, 19: 33-83.

Silva Taboada, G. 1983. Los Murcielagos de Cuba. Editorial Academia, La Habana.

Suarez, W. and S. Diaz-Franco. 2003. A new fossil bat (Chiroptera:Phyllostomidae) from a Quaternary cave deposit in Cuba. Caribbean Journal of Science, 39:371-377.

Tejedor, A. 2011. Systematics of the funnel-eared bats (Chiroptera: Natalidae). Bull. Amer. Mus. Nat. Hist. 353.

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.