late holocene fauna from a cave deposit in western cuba ......taphonomy bone digestion evidence was...

Late Holocene Fauna from a Cave Deposit in Western Cuba:post-Columbian occurrence of the Vampire Bat Desmodus rotundus

(Phyllostomidae: Desmodontinae)

Johanset Orihuela*

*15921 SW 44th St Miami, FL 33185, Email: [email protected]

ABSTRACT.—Here I report a fossil microvertebrate fauna from a late Holocene cave deposit in northwestern

Cuba. This study provides new chronological data for the understanding of the post-Pleistocene survival

of some of Cuba’s rarest extinct bats, and a post-Columbian record of Desmodus rotundus. Remains were

excavated from a superficial, mound-like deposit with no dicernible stratigraphy. The bones of 13 bat

species, one owl, one rail, a small colubrid snake, one frog, a nesophontid shrew, a rat, a mouse, and two

capromyid rodents were extracted from the assemblage. The chiropterans were the most abundant and

included the extinct taxa Desmodus rotundus and Artibeus anthonyi, the rare Lasiurus insularis, and the

endangered Natalus primus. The mixed assemblage originated from accumulated owl pellets and carcasses

of roosting bats. Three Artibeus jamaicensis scapulae from the same level and bone coloration as the

Desmodus material yielded a modern radiocarbon content indicative of post-thermonuclear testing era

(114.9 % ± 0.6 pMC), indicating an essentially modern age for the specimens. Material from the deepest

level, however, is interpreted to be older due to factors such as a probable slow rate of accumulation,

difference in bone coloration, and mineralization. These results support the survival of some extinct bat

species well into the late Holocene of Cuba.

KEYWORDS.—Fossil Bats, Cave deposits, Desmodus, Cuba, Holocene extinctions, Taphonomy, Radiocarbon dates

Introduction

The need for additional fossil evidence andradiocarbon dates continue to limit ourunderstanding of West Indian vertebratepaleontology and historical biogeography.Despite the accumulating research, we donot have a clear chronologic resolution ofwhich species became extinct after thearrival of humans to the islands and theintroduction of exotic species, or whichbecame extinct due to past climatic fluctua-tions (Morgan and Wood, 1986; Diaz-Franco, 2004; Silva-Taboada et al., 2007).Only recently have reliable ‘late occur-rence’ dates been demonstrated for someWest Indian mammals, e.g., sloths andnesophontid insectivores (MacPhee et al.,1999; Steadman et al., 2005; MacPhee et al.,2007). However, causes and extinction datesof other Antillean mammals that are nolonger extant, such as Antillean monkeys,a number of capromyid rodents, and batsremain poorly understood (MacPhee and

Marx, 1997; MacPhee and Flemming, 1990;Davalos, 2004). The discovery and study ofadditional, fossil rich assemblages can pro-vide important data for improving our lastoccurrence records (Shipman, 1981; Andrews,1990; MacPhee et al., 1999; Steadman andDeleon, 1999; Jimenez et al., 2005).

Here I report an interesting cave depositassemblage discovered during a survey ofthe Serrania del Palenque, northwestern Cubain 2003. This research is important becausefew Cuban microvertebrate cave depositshave been radiocarbon dated, and becausethese new data provide a different inter-pretation to the existence of the vampirebat Desmodus rotundus in the Cuban fossilrecord. The common vampire bat Desmodusrotundus remains extant and widely distrib-uted in the continental Neotropics, but inthe Greater Antilles its ocurrence is exclu-sively indicated by fossil material fromCuba (Greenhall et al., 1983; Rey et al.,1988; Morgan, 2001; Suarez, 2005). Presently,

Caribbean Journal of Science, Vol. 46, No. 2-3, 297-312, 2012Copyright 2012 College of Arts and SciencesUniversity of Puerto Rico, Mayaguez

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it is recorded from only four late Quater-nary cave deposits in Cuba (Suarez, 2005).Although these specimens have never beendirectly dated, they come from assemblagesthat range in age from the supposed LatePleistocene of Cueva Lamas and Paredonesto the Middle Holocene of Cuevas Blancasand Centenario de Lenin (Woloszyn andMayo, 1974; Suarez, 2005; Jimenez et al.,2005). This report broadens our understand-ing of the occurrence of Desmodus rotundusin Cuba. Moreover, it provides new infor-mation on some of Cuba’s extinct bat taxa,which may help in determining when thesespecies disappeared.

Materials and Methods

Locality and Deposit

The deposit was discovered during a sur-vey of the Cueva de los Nesophontes inDecember 20, 2003. This cave is located atabout 180 m above sea level on the south-western slope of the Loma del Palenque(23, 00’ 91’’ N and 81, 20’ 91’’ W), a massiveEocene karstic formation 327m above sealevel in the Alturas Habana-Matanzas,Cuba (Nunez Jimenez, 1984). See Fig. 1; D.The deposit was mound-like in shape and

contained a large accumulation ofMeliccoca?(Sapindaceae) seeds, osteological material,insect exoskeletons, plus fine and loose red-dish cave sand (cinnamon rufous Color 40or ferruginous Color 41) see Figs. 2-3. Thecave sand was not damp. The deposit wasshallow and without a defined stratigraphy.The osteological remains were confinedto the first 8 cm of the deposit measuredfrom the highest part of the mound to thedeepest bone found below the floor level.Fresh guano was present toward the pos-terior region of the deposit and only onefresh owl pellet containing remains of Musmusculus was found at the surface (Fig. 3).The remains were in a small secondary

chamber on a passage leading to a larger gal-lery housing a colony of Artibeus jamaicensisparvipes. The Artibeus gallery had two largehorizontal openings or sinkholes from werefresh air rushed in occasionally (Fig. 1.2;see X). The area was inspected carefullybefore excavation and removal of the mate-rial. A small test pit measuring 30x30x30 cm

was dug to extract the matrix. The mate-rial was then dry sieved with a fine screenmesh (0.3 cm).Treatment of the material followed Silva-

Taboada (1974) except in color-coding theremains and his exclusion of juvenile speci-mens. The identifiable osteological elementsused for systematic taxonomic classifica-tion included the skull, mandibles, scapula,humerus, radius, pelvis, femur, tibia, and insome cases phalanges or digits. Only identi-fiable and diagnostic specimens were usedfor proper species assignment. These ele-ments were then sorted into long bones(humeri, radii, phalanges, and femora), cra-nial (skulls and mandibles), and miscellaneous

Fig. 1. (1. A-E) Map of the Cuban Archipelago show-ing sites where Desmodus rotundus material has beenreported. The localities are organized in chronologicalorder of publication. A is Cueva Lamas described byKoopman (1958). B is Cueva del Centenario de Leninin Punta Judas, material described by Woloszyn andMayo (1974) and Suarez (2005). C is the materialdescribed and dated by Jimenez et al., (2005) fromCuevas Blancas, Habana. D is the material reportedby Suarez (2005) for Cueva de los Paredones, in westHabana. E is the material reported here from Cuevade los Nesophontes, Palenque Hill, Matanzas-Habana.Shaded areas indicate land 300 m above sea level.Fig. 1.2. This plan shows a sketch of the deposit’s

locality within Cueva de los Nesophontes. I. showsa passageway leading to the rest the cave. II.-III.show the small dolines or sinkholes where air andred soil have been entering the cave. X marks thedeposit’s location.

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categories to calculate element frequencyand breakage (Table 2). The specimens wereclassified with the aid of published literatureand comparison with museum collections.Chiropteran systematic classification fol-lowed Silva-Taboada (1979), Wetterer et al.,(2000), and Simmons (2005). Subspecies wereassigned following their geographical rangein Cuba. Other mammalian classification fol-lows McKenna and Bell (1997). Avian osteol-ogy nomenclature and diagnosis followedWetmore (1922 and 1939) and Olsen (1979),and for the Buthidae scorpion Moreno (1942)and de Armas (2006).

The studied collections mentioned hereare housed in the American Museum ofNatural History (AMNH), New York, theFloridaMuseumofNatural History (FLMNH)in Gainesville, FL, the speleological col-lections of Jorge Ramon Cuevas (JRC) andNorbert Casterett (NC). The material isdeposited in the Cuban National Museumof Natural History (MNHNCu) in Havana.

Specimens were measured with digitalcalipers calibrated to the nearest 0.01 mm,and the measurements are expressed in mil-limeters (mm). Measuring methods followedSilva-Taboada (1979) and Suarez (2005).Descriptive statistical analyses were per-formed with STATISTICA-software version 5(1995). These analyses compared variationbetween the specimens from this deposit,and museum collection material.

The material destined for AMS radiomet-ric dating was collected in situ from thearea nearest the Desmodus skull. Since theskull itself could not be dated due to itsrarity, three cream-colored Artibeus jamaicensisscapulae from the most superior part of themound, in direct association to the Desmodusskull, were chosen to be dated. In addition,four Brachyphylla nana radii encrusted withmatrix, colored dark cinnamon and takenfrom below the floor level, were not dat-able due to their lack of collagen (RonaldHatfield person. comm.). Even though dating

Fig. 2. The deposit in profile. A general aspect of the deposit. B shows the depth of the bones and colorrelationship. C shows the largest seeds on the deposit. Including the large accumulations of Meliccoca sp. seeds,still undecomposed, plus Spondias sp. and other non identified taxon.D is a close up photograph of theDesmodusspecimens in situ. Note the dark, organic residue inside the skull. The dated material came from this area.

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of specimens by association in West Indiancave deposits can sometimes be erroneous,in this case taphonomic analyses suggestthat the dated specimens were depositedwithin the same span of time as indicatedby their color, depth, and level of minerali-zation in comparison to their position inthe deposit.The radiometric dates (AMS= Accelerator

Mass Spectrometry) reported in this workwere performed by Beta Analytic Inc. inMiami, FL and followed their standards.The bone collagen extraction was performedsuccessfully and the lab reported no prepa-ration difficulties (Ronald Hatfield person.comm.). In literature, dates are usuallyreported in radiocarbon years before pres-ent (Present=1950 A.D., rcybp=B.P.) forsamples that are older than 1950 A.D. How-ever, results for samples containing moreC14 than the modern reference standards,

and younger in age are reported as “Per-cent Modern Carbon” (pMC). These resultsindicate that the dated specimens con-tained excess radiocarbon, and can be con-fidently judged to be less than 50 years olddue to thermonuclear weapon testing (Huaand Barbettii, 2004; Jull et al., 2004).The color numbers were assigned with

Pantone Color Chart (PMS) and follow thecolor nomenclature of Smithe (1975).

Results

Age of the Bones

Three Artibeus scapulae yielded an AMSC14 percent age of 115.9±0.6 pMC (Beta-210380) which resolves into standard nota-tion as 1989-1992 C.E. (1s) or 1957-1993 C.E.(2s). The dark chestnut and cinnamoncolored (Color 33) Brachyphylla materialextracted from the deepest level did notyield enough collagen for dating due totheir mineralized state.The deposit’s age prior to the C14 dating

was hypothesized to be very young (i.e. lateHolocene) due to the coloration of the bones,the presence of Mus and Rattus, the well-preserved state of the bones, seeds, andinsect exoskeletons. The bones in the moundshowed mild signs of exposure to humid-ity and regular air currents, all which aredestructive agents of bones (Andrews, 1990;Tarhule-Lips and Ford, 2006).

Taphonomy

Bone digestion evidence was scarce, onlythe fresh pellet material and the Desmodushumerus (JRC2389) showed distinct diges-tion marks (Fig. 8F). Post-depositionalmodifications such as mineralization, soilcorrosion, flaking, or exfoliation due tohumidity, and bone coloration were verycommon. The bones buried under the floorline had a darker coloration, higher degreesof mineralization and calcium carbonate(CaCO3) calcifications than the ones aboveon the mound. The bones in and above themound, however, showed mild flaking andbone discoloration, but no mineralization.Rounded bones or other water-transportrelated modifications were not found inthis assemblage.

Fig. 3. Deposit mound and assemblage showing itskey components. Photograph taken in situ before removalof material. Notice the abundance of seeds, bones, andinsect exoskeletons. Scale on lower left hand = 30 mm.

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The coloration and the degree of bonemineralization varied, generally, in rela-tion to their vertical position within thedeposit; however, this was not the casewith their degree of CaCO3 incrustation.The bones found above the floor line didnot make metallic sounds when strokedagainst each other, but the ones foundbelow the floor line did make the dis-tinct metallic sound characteristic of highdegrees of mineralization (see Linares,1968; Silva-Taboada, 1974). The bonesfound in the most superior level of themound (3 cm above floor the line) had colorsranging from light cream to light yellow-orange hues (Colors 54 and 18), whereasthose below the floor level (4 - 8 cm) variedfrom dark flesh-ocher and cinnamonbrown to chestnut and near black (Colors132D and 33).

Bats comprised the most common groupin the deposit (83%), of which species thatexclusively use caves for roosting were the

most common. Many of the species foundare presently very rare, extirpated, or extinctin Cuban territory and rare in museum col-lections; these are Natalus primus, Desmodusrotundus, Chilonatalus macer,Artibeus anthonyi,and Lasiurus insularis (Silva-Taboada, 1979;Morgan, 2001; Tejedor et al., 2004, 2005).See Tables 1 and 2.

Systematic Paleontology

Order Chiroptera Blumenbach, 1779Suborder Microchiroptera Dobson, 1875Family Mormoopidae de Saussure, 1860

Mormoops blainvillei Leach, 1821

This taxon is represented by three incom-plete right humeri (JRC 2087, 2169, and2206), one complete left humerus (JRC2161),and a complete pelvis (unnumbered). Thehumeri are dark brown and cinnamonbrown in color (PMS176 or Color 33) andwere found at floor line, below the mound.

Table 1. Summary of present and absent taxa in relationship between bone color and their level in thedeposit. N represents the total number of specimens for that taxon.

Mound Level Subsoil Level

N SpeciesCream

PMS157/Color 54Yellow-Cream

PMS164/Color18Dark Clay

PMS173/Color 132DChestnutPMS1545

BATS96 Brachyphylla nana + + + +66 Artibeus jamaicensis + + – –40 Monophyllus redmani – + + –20 Eptesicus fuscus + + – –11 Pteronotus parnellii – + + –4 Mormoops blainvilleii – + + –2 Desmodus rotundus + – + –2 Artibeus anthonyi – + + –2 Tadarida brasiliensis – + – –2 Macrotus waterhousii – – – +1 Lasiurus insularis – – + –1 Natalus primus – – + –1 Chilonatalus macer – + – –

AMPHIBIANS5 Osteopilus sp. – – + –

RODENTS1 Mus musculus + – – –1 Rattus sp. – + – –

AVES1 Nesotrochis sp. – – – +1 Tyto alba – + – –

Number of specimens arranged in order of abundance and assorted by locality and bone color plus stratigraphichorizon. Shaded (+) marks the presence of a taxon at the stratigraphic horizon indicated; (–) indicates absence.


The pelvis is colored in a light cream color(PMS157 or Color 54), and was found atthe top of the mound in association withEptesicus and Desmodus material. No sig-nificant quantitative or qualitative variationwas found between these specimens andmuseum specimens.

Pteronotus parnellii parnellii Gray, 1843

This subspecies is represented by 11 ele-ments: four dentary bones, two right(JRC2219-2220) and two left (2218-2221);two incomplete humeri (JRC2082-2083); threeunnumbered auditory bullae, one completepelvis, and one unnumbered complete scap-ula. The mandibular and humeral materialwas found at the level of and below the floorline, and is colored in light reddish brown(PMS164). The remaining elements werefound on the mound and have a colora-tion of light orange-yellow to cream huesas the mound material to which they wereassociated. All mormoopids occurred atthe same level. There was no significantvariation between these specimens andmuseum specimens.

Family Phyllostomidae Gray, 1825Desmodus rotundus E. Geoffroy, 1810

This species is represented by a distalhumerus portion (JRC2389) and a nearlycomplete skull (JRC600). The skull wasdirectly associated with Eptesicus fuscusand the dated Artibeus jamaicensis scapulae(See Fig. 2 and 3). The skull is missing theright zygomatic arch and has a small per-

foration on the supraorbital area (Fig. 4). Itpreserves all dentition except the canines,making it the most complete specimenfound in Cuba. The skull shows evidenceof weathering such as flaking and exfo-liation on the lateral side of the braincasethat suggests mild exposure to cave ero-sion (Andrews, 1990). The skull is creamcolor (PMS5875/1365 or Color 54), butthe humerus is colored in a dark cinna-mon brown (PMS176 or Color 223B) andshows digestive modifications as shownby Andrews (op. cit.) see Fig. 8F. Skullmeasurements are total length of the skull25.37, palatal length 9.87, and height of theskull 12.06, maximum zygomatic width11.83, canine width 6.14, and postorbitalwidth 5.51.The Cuban vampire bat fossils have

been previously classified as a subspecies,D. r. puntajudensis, (Woloszyn and Mayo,1974), or as endemic species D. puntajudensis(Suarez, 2005). However, I believe that theCuban vampire bat fossils are conspecificwith the mainland extant D. rotundus, andnot a new form. These specimens werecompared to a wide geographical range ofneontological material, and were found tobe inseparable from D. rotundus (Fig. 5).However, the taxonomic status of the Cubanvampire bats are fully discussed elsewhere(Orihuela, 2011).

Brachyphylla nana nana Miller, 1907

This is the most common taxon in thedeposit. It is represented by 96 elements:30 humeri, 21 mandibular rami, 10 radii,7 incomplete skulls, three femora, and onescapula. This species was found on alllevels of the deposit. Their bone colora-tions cover a large range of hues from verydark brown (cinnamon brown: PMS4625 orColor 33) to light cinnamon (Color 123A).Some of the deeply stained material hadthin films of CaCo3 and matrix adheredto the bone. There was no significant varia-tion between these specimens and museumspecimens. However, the skulls andhumeri appear more robust than the recentmaterial to which it was compared (seeFig. 5 A-C).

Table 2. Table of osteological elements and theirfrequency in the assemblage.

Osteological Elements Number Percentage (%)

Long bones total 180 60%Humeri 93 31%Radii 67 22%Cranial 64 21%Mandibles 43 14%Complete elements 177 60%Incomplete elements 121 41%Total identifiable 297Total 324 100%

Summary of osteological elements and their frequencyin the assemblage.

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Monophyllus redmani clinedaphusMiller, 1900

This subspecies is represented by 40 ele-ments: 14 humeri, five incomplete skulls,five femora, 13 phalanges, and three radii.This species was encountered in the moundand below the floor level. The bones showa wide range of colors from light cream(PMS157 or Color 54) to light and dark red-dish brown (PMS718/4625 or Color 123B).Some of the specimens found below thefloor line are slightly mineralized, with cal-cifications within the brain cavity. Two ofthe skulls, JRC53 and 481, found below thefloor level and the darkest in color (Darkclay Color 123B-C), are significantly differ-

ent (p<0.050) in five skull measurementsthan the compared neontological sample, orthe set given by Silva-Taboada (1974-1979).These specimens are smaller in total skulllength, maximum palatal length, and max-imum dental length and larger in post-orbital width and skull height.

In 1974, Silva-Taboada reported fossilsslightly larger than a neontological sample.In this case, however, the fossils are signif-icantly smaller than the neontological spec-imens I compared or the ones reported bySilva-Taboada. Morphological and size com-parisons revealed that it does not belongto M. r. portoricencis from Puerto Rico.

Macrotus waterhouseii minorGundlach in Peters, 1865

A complete right ( JRC2080) and a rightdistal humerus (JRC2079) with slight incrus-tations of calcium carbonate represent thissubspecies. These were the deepest speci-mens found below the floor level, and theones with darkest brown coloration (chestnutandmaroon (PMS4625/1545 or Color 31-32).They were in close association with theNesotrochis femur of the same coloration.There was no significant variation betweenthese specimens and museum specimens.

Artibeus anthonyiWoloszyn and Silva-Taboada, 1977

This extinct fossil species is representedby two complete left humeri (JRC2394 and2122), plus a possible right distal humerus(JRC2407). One of the specimens, JRC2122,and the darkest in color (PMS173 or Color123A-B) was found below the mound. Theremaining two, and lightest in color (PMS164or Color 18) were found above the floor lineand in association with Natalus and Lasiurusspecimens. This species was identified dueto its robust appearance and larger sizein total length of the humerus comparedto Artibeus jamaicensis (Woloszyn and Silva-Taboada, 1977). No other discrete morphol-ogy was found to be diagnostic. Maximumlengths for two of these specimens measured34.14 and 37.67 mm, in contrast to the max-imum of 34.2 mm given by Silva-Taboada(1979) for the Cuban subspecies A. j. parvipes,or the maximum measurement, 33.68 mm,

Fig. 4. The skull of Desmodus rotundus, showing thesuperior, left lateral and ventral aspects of the almostcomplete skull. Notice the erosion-caused flaking ofthe periosteum on the parietal bones. Adhered limecan be noted on the maxilla. Scale = 10 mm.


obtained from specimens in this deposit.However, the dimension of the smallestspecimen overlaps with the maximum givenby Silva-Taboada (op.cit). Nonetheless, thisspecimen was assigned to this taxon duethe much larger epitrochlea-capitulum diam-eter of 6.24 mm, as compared to the maxi-mum of A. j. parvipes 6.19 mm from thissample and neontological material (Fig. 7;D-D1).There is evidence indicating that Artibeus

anthonyi has survived into the middle tolate Holocene (Jimenez et al., 2005; personalobservation). Therefore, in this work thisspecies is not considered a Pleistocene marker(Suarez and Diaz-Franco, 2003; Mancinaand Garcia-Rivera, 2005).

Artibeus jamaicensis parvipes Rhen, 1902

This subspecies is represented by 66 spec-imens and is the second most common

taxon in the assemblage: 17 adult humeri,plus ten juveniles; eight mandibular ramii,plus 12 juveniles; 12 radii, six scapulae (threewhich were dated), 14 teeth, and three skulls(two adults and one juvenile). This specieswas present only on the mound were itwas associated with the Eptesicus andDesmodus material. The specimens werecolored light to yellow/orange (Color 18 orColor 223B-C).

Family Natalidae Gray, 1866Natalus primus Anthony, 1919

This species is represented by a singleleft complete humerus (JRC2085) foundbelow the floor line. Like most of the mate-rial recovered from that area it has a lightorange-yellow coloration (PMS 164 orColor 18). The specimen measured a totallength of 27.02 mm, maximum proximalwidth of 3.44 mm, and maximum distal

Fig. 5. Scatter plot showing the relationship of the Desmodus rotundus material from this deposit (filled circle)versus neontological specimens of D. rotundus from museum collections at FLMNH and AMNH (open circles),in addition to the fossil taxa D. stocki (open cross) and D. draculae (filled triangle).

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width of 3.88 mm. No variation was foundbetween this specimen and other museummaterial. Fig. 8D.

Chilonatalus macer Miller, 1914

This species is represented by a single,left proximal humerus (JRC2208) foundbelow the floor level, and it is colored inlight orange-yellow (PMS164 or Color 18).The specimen’s maximum proximal widthis 3.13mm and least medial width is 1.16 mm.No variation was found between this spec-imen and museum material. Fig. 8F.

Family Vespertililionidae Gray, 1821Eptesicus fuscus dutertreus Gervais, 1837

This subspecies is represented by threeskulls (JRC594, 595, and 596), four mandi-bles, two scapulae, four humeri, and seventeeth. This species was found only abovethe floor line, inside the mound, and indirect association to the Desmodus mate-rial (fig. 2 D and 3). These specimens arelight cream colored (PMS 157 or Color 54)and the darkest are light orange-yellow(Color 18). No variation was found betweenthese specimens and neontological material.

Fig. 6. Brachyphylla nana skulls. A-C, left lateral (A), superior (B), and inferior (C). The first-row specimenresembles a juvenile. The second-row specimen is mineralized and was found in the deepest part of the depositassociated with the Macrotus waterhouseii material. Some photographs have been reversed. Monophyllus redmaniskulls. D-G, Left lateral (D), superior (F), and inferior (G) views. Note the degree of mineralization and Calcitegrowth on the first and second skulls. The last row skull was found to be significantly smaller in three linearmeasurements than previously reported in Cuba, Silva-Taboada (1974, 1979).


Lasiurus insularis Hall et Jones, 1961

This species is represented by a restoredright humerus (JRC2078 and 2142) with atotal length of 35.89 mm, maximum distalwidth of 3.81 mm, and maximum proximalwidth of 4.70 mm. This specimen was recov-ered below the floor level, and directlyunder the mound. It is colored in a lightbrown or flesh ocher (PMS173 or Color 132D)Fig. 7A and 8G. No significant variationwas detected between this specimen andmaterial reported by Silva-Taboada (1979)or modern material at AMNH.The occurrence of Lasiurus in cave depos-

its is not rare as previously assumed. Thistaxon has been previously reported fromowl pellet deposits in Cuba by De Beaufort(1934), Silva-Taboada (1979), and Mancinaand Garcia-Rivera (2005). Additionally, thisgenus is reported from other cave depositsin the U.S. (Quay and Miller, 1955; Myers,1960) and the island of Bermuda (Gradyand Olson, 2006).

Family Molossidae Gervaisin de Castelnau, 1855

Tadarida brasiliensis muscula Gundlachin Peters, 1862

This subspecies is represented by aleft m3 and an incomplete right dentary(JRC2002). These were found with theDesmodus material and have the same levelof coloration. No variation was foundbetween these specimens and neontologi-cal material.

Aves

Gruiformes Bonaparte, 1854Family Rallidae Vigors, 1825

Genus Nesotrochis Wetmore 1938

An incomplete left proximal femur(JRC2012b) represents the rail family. Thisspecimen was found in the deepest bone-bearing level of the deposit, associated withthe non-datable Brachyphylla material. It iscolored in a dark chestnut hue as the other

Fig. 7. Lasirus insularis and Artibeus sp. humeri. A shows a complete L. insularis humerus. B-D, variation rangeof Artibeus humeri shown in anteroposterior and oblique profiles. B is an Artibeus jamaicensis parvipes specimenencountered on the top of the deposit mound. C is an intermediate Artibeus cf. anthonyi specimen found abovethe floor line. D is an Artibeus anthonyi specimen found below the floor line.

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Fig. 8. Diversity of encountered taxa. A shows frontal and lateral views of a proximal Nesotrochis sp. (cf.)N. picapicensis found in the deepest bone-bearing horizon of the deposit. B shows superior and lateral views of aTyto alba phalanx or digit. This specimen was found in close association with the Desmodus material. C showssuperior, inferior and transverse views of two small colubrid snake fused vertebrae. D, Natalus primus distaland proximal humerus in frontal views. E, Chilonatalus macer proximal humerus lateral and frontal views.F, Desmodus rotundus humerus comparing the specimen from this deposit (on the right) and a neontologicalspecimen (on the left). G, proximal and distal aspects of Lasiurus insularis humerus. The full view of thisspecimen is shown in figure 6. H shows an almost complete exoskeleton of a Buthidae scorpion, possibly of thegenus Centuroides sp. Note the small auxiliary telson, and the slightly hairy vesicule (in magnification). Scaleequals 7 mm inside magnification. I, frog osteological elements, possibly Osteopilus sp. J, complete Lipotyphlan(= Insectivoran) radius, probably Nesophontes.


elements found at this depth, with smallencrustations of matrix. The specimen’s mea-surements are as follow: greatest length takenfrom the front of the femoral head to theposterior of greater trochanter is 12.97 mm;greatest proximal width: 10.75 mm; leastwidth of shaft: 4.73 mm and least width ofthe femoral head: 4.83 mm (fig. 8A). Thisspecimen probably belongs to the Cubanform N. picapicensis Fischer et Stephen, 1971.This taxa is supposed to have gone extinctaround the 18-19th Century, and is there-fore, not considered a Pleistocene marker(Jimenez, 1997).

Strigiformes Wagler, 1830Family Tytonidae Ridgway, 1914

Tyto alba. The Common barn owl isrepresented by a single phalanx or digit(JRC2012a). See fig. 8B.

Discussion

The assemblage reported here is the resultof an accumulation of owl pellets and batroosting activity. The presence of guanonear the deposit, and the high concentrationof large seeds of Meliccoca sp. and Spondiassp. suggest that this area has been recentlyused by bats. This is additionally supportedby the presence of juvenile Artibeus andBrachyphylla specimens found at the moundlevel, plus the presence of an active colonyof Artibeus jamaicensis parvipes in the adja-cent gallery.There were slightly more post-cranial

than cranial elements in the assemblage(Tables 2 and 3). This ratio correspondsto those usually found in Tyto alba pelletassemblages. Andrews (1990), however, hasshown that short-eared owls Asio flammeusand little owls Athene sp. also have high per-centages (³ 90%) of long bone occurrence,with small degrees of bone fragmentation.These raptors are present in the Cubanfauna (Garrido and Kirkconnell, 2000),and therefore could have contributed tothe deposit.Low levels of bone breakage has been

associated with nocturnal predators, espe-cially owls because of the small amount ofprey loss through digestion, and becauseowl pellets enhance the preservation of

their osteological content (Andrews, 1990;Kusmer, 1990). However, due to the pre-dominance of long bones such as humeriand radii, other nocturnal avian raptorscannot be entirely ruled out. Low tram-pling is indicated by the low degrees ofbone fragmentation, which is less than50% in this assemblage. The breakagelevels in osteological material from owlpellets are not high compared to otherpredators such carnivores or diurnal raptors(Andrews, 1990; Kusmer, 1990; Jimenezet al., 2005). Some of the bat skulls encoun-tered in this deposit had cerebral cavity per-forations that are usually associated withowl predation (Dodson, and Wexlar, 1979;Andrews, 1990). The Desmodus rotundusskull had these perforations. Its cerebralcavity contained grass seeds and pelletmaterial, suggesting that this specimencame from a pellet.The absence of some taxa are worthy of

attention. For example, snakes are rarely ornever encountered in owl pellet deposits(Jimenez et al., 2005), as are the scorpions,or the Lasiurus sp. tree bats. In addition,otherwise common species such as the batsPhyllonycteris poeyi and Phyllops falcatus, theintroduced House Sparrow Passer domesticus,and the Boromys spiny rats are absent. Themice, rats, and the nesophontid lipotyphlanwere never occurred in the high numbersseen in other deposits (pers. observations).In other deposits within the same cave,“classic” owl pellet assemblages have beenfound with very high accumulations of suchgenera (i.e., Rattus,Mus, Boromys, Nesophontes,and passerine birds).

Conclusions

The presence of the common vampire batDesmodus rotundus is of great importancebecause it indicates the presence of this taxonin post-Columbian Cuba, a more recentpart of the Holocene than ever expected forthis species. This prompts a series of ques-tions on the occurrence and extinction ofthe vampire bats in Cuba. Why are fossilremains of this species so scanty? CouldD. rotundus still be present on the Cubanfauna? If so, why are they not commoneven with the abundance of suitable prey?

J. ORIHUELA308

Is D. rotundus a systematic visitor, Recentor accidental? Could disease transmissiondue to the blood feeding habits be involvedin its demise, and not the extinction ofthe edentates?

The deposit mound seems to have formedvery recently, as much as a couple hundredyears, as taphonomic evidence, the radio-metric date, and the presence of Rattussuggest. The deposit appears to have beenundisturbed until discovery. There was noevidence of water transportation, i.e., scat-tering of bones along the walls as observedby H. E. Anthony in Daiquiri, Cuba (1919),and Linares in Venezuela (1968). Otherappreciable forms of deposit disturbance,such as mixing of different colored bones orbone-rounding modification was also absent.This evidence suggest that the deposit formedslowly, with the mound material above thefloor level (i.e., zone with decomposed seedsand light colored non-mineralized bones)being the most modern in age. This is fur-ther supported by the state of the seeds (somestill with dried pulp, see figs. 2-3) and thefine state of the bones under cave corrosiveconditions such as air currents, humidity,or temperature fluctuations (Andrews, 1990;Tarhule-Lips and Ford, 2006).

Consequently, for the deepest material(i.e., below the floor level, mineralized, anddark-colored) greater age is inferred. Thedate, however, is unknown since datingwas unsuccessful at that level. Since thechemical processes that lead to changes inbone coloration are so far chronologicallyunpredictable, sometimes taking thousandsof years or less than five years due to theever-changing microenvironments insidecaves, the coloration of the bones cannot

Table 3. Systematic species account. The numbersof remains for each taxon are in parentheses.

Arachnida

Amblypygi? sp.

ScorpinidaButhidae?Centuroides or Tityus sp. (1)

Crustacea

Geocarcinus sp. (1)Amphibia

AnuraHylidae? Osteopilus sp. (5)

Reptilia

OphidiaColubridaesmall colubrid (2)

DinosauriaAvesGruiformes

Rallidae cf. Nesotrochis sp. (1)Strigiformes

TytonidaeTyto alba (1)

Mammalia

RodentiaMuridaeMus musculus (1)Rattus sp. (1)

Capromyidaemesocapromyid (3)

ChiropteraYangochiropteraMormoopidae

Mormoops blainvilleii (4)Pteronotus parnellii parnelli (11)

PhyllostomidaeDesmodontinae

Desmodus rotundus (2)Brachyphyllinae

Brachyphylla nana nana (96)Glossophaginae

Monophyllus redmani clinedaphus (40)Phyllostominae

Macrotus waterhouseii minor (2)Stenodermatinae

Artibeus anthonyi (3)Artibeus jamaicensis parvipes (66)

NatalidaeNatalus primus (1)Chilonatalus macer (1)

VespertilionidaeEptesicus fuscus dutertreus (20)Lasiurus insularis (1)

Table 3. Continued.

Mammalia

MolossidaeTadarida brasiliensis muscula (2)

Lipotyphla (=Insectivora )

SoricomorphaNesophontidae

? Nesophontes sp. (1)

Summary of taxa encountered in the deposit.Material followed for taxonomic and systematicclassification is indicated in Materials and Methods.

(Continued)


be effectively used for the dating of thedeposit. The dark red bone coloration ofthe buried material is most likely due tothe incorporation of mineral impurities inassociation to taphonomic, physical, andchemical processes that occur within caves(Andrews, 1990; Robles et al., 2002). Nev-ertheless, the presence of specimens withdifferent coloration in the same depositcould indicate different times of expo-sure to the cave environment and to thematrix. This could mean that the red col-ored bones were exposed longer to thered sand matrix than the grayish bonesabove the mound not in contact (NunezJimenez, 1984; Pregill et al., 1994; McFarlaneet al., 2002).There is room for improvement in our

understanding of West Indian paleon-tology and historical biogeography. Thedocumentation of its past diversity as pre-served in the fossil record will continueto expand and deepen our understating ofCuba’s vertebrate extinctions, and therefore,the Caribbean.

Sinopsis.—En este trabajo se describe undeposito fosil cavernario procedente deloccidente de Cuba. Allı se descubrieronrestos de microvertebrados en un depositomonticular en el suelo del salon. De estese excavaron restos de trece taxones demurcielagos, dos aves, una culebra pequena,ranas, una musarana nesophontida, roedoresmuridos y capromidos entre semillas yarena ferralıtica roja. Entre las especiesde murcielagos se encuentran el extinctoArtibeus anthonyi, el raro Lasiurus insularis,Natalus primus, y el desaparecido y rarısimovampiro comun Desmodus rotundus. Estaacumulacion de especies y ausencia de otras,resultan inusual en depositos cubanos. Eldeposito parece haber sido formado por lasacumulaciones de perdigones regurgitadosde lechuzas u otras rapaces, mas laactividad de murcielagos. Los altos nivelesde preservacion, aun bajo condicioneserosivas, la presencia de taxones introducidos,y el fechado radiocarbonico (114.9±0.06 pMC,calibrado a 1957-1993 A.D.) indican unaedadHoloceno tardıa (i.e., post-Columbina),esencialmente moderna para el deposito.Esto datos indican procesos de extincion

trans-Holocenicos, en vez de abrupto orepentinos al final del Pleistoceno paramurcielagos cubanos.

Acknowledgments.—I thank my friend andcolleague Dr. Adrian Tejedor for alwaysfinding the time and patience to guide me,without his constructive criticism none ofthis research could have been possible. Forallowing access to specimens in museumcollections I am greatly indebted to ChrisNorris, Susan Bell, Judy Galkin, and A.Tejedor at the AMNH in New York.David Steadman, Richard Hulbert, andTom Webber (FLMNH) at the Universityof Florida; Bill Amaral and Judy Chupaskoat MCZ at Cambridge for providing valu-able information on their specimens, andGilberto Silva-Taboada, Osvaldo Jimenez,and William Suarez at MNHNCu in Cubafor always answering my questions., andproviding access to specimens (OJ). Forfieldwork, curation of specimens, andexcavation logistics I thank Leonel P.Orozco, Ricardo Viera Munoz, and CandidoSantana (members of the SpeleologicalSociety of Cuba), Alberto Sanchez, and JoelMonzon. For making the dating processa pleasure I thank Ronald Hatfield andthe staff at Beta Analytic Inc. I thank A.Tejedor, Ross D. E. MacPhee, David W.Steadman, Gary S. Morgan, TamaraCastano, and Pere Bover for providingcritical comments and revisions thatimproved this article. Finally, for encour-aging my love for West Indian biogeogra-phy and paleontology I thank Leonel P.Orozco, A. Tejedor, Gilberto Silva-Taboada,H. E. Anthony, David Steadman, R.D.E.MacPhee, Karl Koopman, Gary S. Morgan,and Oscar Arredondo.

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Appendix 2: Additional Informationon flora, and fauna

Additional Flora and Fauna

Plants were represented by seeds of atleast two species of Euphorbia (Euphorbiacea),plus seeds of Meliccoca (Sapindaceae) thatoccurred in large concentrations on top ofthe mound, two Spondias (Anacardiaceae)or ? Giras (Lecythidaceae) seeds and onethat remains unidentified (see fig. 3). Theseaccumulations are most likely the result ofthe large fruit bats Artibeus and Brachyphylla(Silva-Taboada, 1979; Appendix 1). Otherorganisms found in the deposit includedland mollusks of the genus Zacrhysia andChondropoma, exoskeletons of amblypygids,spinobolidid centipedes and other uniden-tified insects, plus a well-preserved smallbuthidae scorpion exoskeleton. Also, fivesmall frog specimens, probably pertainingto the Hylidae family, two unidentified smallcolubrid snake vertebrae, and one land crabmandible cf. Geocarcinus sp. (see figs. 3 and7; See Table 3 for complete faunal list).Other mammals included two small to

medium sized capromyid rodent ribs (prob-ably a mesocapromyine). These were foundin association to the undated Brachyphyllaspecimens and the Nesotrochis specimen.A Rattus sp. first upper molar was foundwithin the mound and below the Desmodusmaterial. A fresh owl pellet was found closeto the Desmodus skull containing remainsofMus musculus. A small eulipotyphlan radiusprobably pertaining to the Nesophontidaeshrew family was found at the floor level,below the bone mound, and in associa-tion with L. insularis and A. anthonyi speci-mens (fig. 7 J).

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late holocene fauna from a cave deposit in western cuba ......taphonomy bone digestion evidence was...

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