Submitted : July 9th , 2020 – Accepted : December 30 th , 2020 – Posted online : January 29 th , 202 1
To link and cite this article:
doi: 10.5710/AMGH.30.12.2020.3379
1 NEW RECORD OF THE VAMPIRE DESMODUS DRACULAE (CHIROPTERA) FROM
2 THE LATE PLEISTOCENE OF ARGENTINA
3 NUEVO REGISTRO DE DESMODUS DRACULAE (CHIROPTERA) DEL PLEISTOCENO
4 TARDÍO DE ARGENTINA
5
6 SANTIAGO BRIZUELA*,1 and DANIEL A. TASSARA2
7 1Universidad Nacional de Mar del Plata, CONICET, Departamento de Biología, Facultad de
8 Ciencias Exactas y Naturales, Mar del Plata, Argentina. Funes 3250, B7602AYJ Mar del Plata,
9 Argentina. [email protected]
10 2 Museo Municipal de Ciencias Naturales Pachamama, Niza 1065, B7609LNC Santa Clara del Mar,
11 Partido de Mar Chiquita, provincia de Buenos Aires, Argentina.
12
13 16 pag. (text + references); 3 figs.; 2 tables
14
15 Running Header: BRIZUELA AND TASSARA: LATE PLEISTOCENE VAMPIRE FROM
16 MIRAMAR
17 Short Description: The presence of Desmodus draculae, an extinct giant vampire bat, from the
18 Pleistocene of Buenos Aires is confirmed.
19 Corresponding author: Santiago Brizuela, [email protected]
20
21
22 Keywords. Vampire. Bat. Desmodontinae. Dentary. Miramar.
23 Palabras clave. Vampiro. Murciélago. Desmodontinae. Dentario. Miramar
1 24
25 BATS (Chiroptera) are one of the most diverse groups of extant mammals (Amador et al., 2018), but
26 this diversity is not represented in their fossil record (Teeling et al., 2005). In Argentina there are
27 presently four bat families represented, all Yangochiroptera, with more than 60 species (Díaz et al.,
28 2016). In contrast with this diversity, the fossil record of bats in Argentina is scarce and patchy
29 (Czaplewski, 2010), mostly accounting for very few, isolated, partial elements (Table 1).
30 One interesting record of fossil bats of Argentina is that of an isolated upper canine from the
31 late Holocene of southeast Buenos Aires referred to Desmodus cf. D. draculae (Pardiñas & Tonni,
32 2000), a large, extinct vampire. This species had a large late Pleistocene–late Holocene distribution
33 from Mexico to Argentina (see Ubilla et al., 2019). However there is some taxonomic uncertainty
34 (cf., aff.) regarding the specific assignation of the fossils of D. draculae at its latitudinal limits. Here
35 new fossil material from Late Pleistocene sediments outcropping near the city of Miramar
36 (Argentina) (Fig. 1) is described and discussed.
37 Institutional Abbreviations. CML, Colección Mastozoológica, Fundación Miguel Lillo,
38 Univerdidad Nacional de Tucumán, Tucumán, Argentina; LIEB-PV, Laboratorio de
39 Investigaciones en Evolución y Biodiversidad, Universidad Nacional de la Patagonia “San Juan
40 Bosco”, Esquel, Argentina; MLP, Museo de La Plata, Universidad Nacional de La Plata, La Plata,
41 Argentina. MPH-P, Museo Municipal Punta Hermengo, Collección Paleontológica, Miramar,
42 Argentina.
43 MATERIALS AND METHODS
44 The fossil here described is housed in the Colección Paleontológica of the Museo Municipal
45 Punta Hermengo (MPH-P). It was studied under stereoscope microscope, and compared to available
46 reference materials (Diaemus youngi CML 1343; Desmodus rotundus CML 8664), specimens
47 available in MorphoSource (https://www.morphosource.org), and bibliography. Images were
48 obtained by Scanning Electron Microscope (SEM) at the Laboratorio de Microscopía Electrónica of
2 49 the Universidad Nacional de Mar del Plata. Osteology follows Giannini et al. (2006) with
50 comments from Rädulet (2006). Systematics follows that of Cirranello et al. (2016). Dentary
51 measurements from Morgan et al. (1988), Barquez et al. (1999), and Davis et al. (2010) were used
52 for comparison.
53 Stratigraphical provenance. The fossil site is located 700 m north of the mouth of the La
54 Ballenera stream (38° 19' 25.60" S; 57° 56' 6.74" W), 9 km south of Punta Hermengo (Miramar
55 city) (Fig. 1). The fossil was located in situ within a cave section (1.2 m diameter) which is
56 attributed to a Mylodontidae. The cave section is situated near the top of a 5 m sea cliff in which six
57 stratigraphic units can be recognized (Fig. 2). The lower deposits, units A–C, which are topped by
58 calcrete levels (C 1–3), correlated to units A–D of Tonni & Fidalgo (1982), units I–III of Alberdi et
59 al. (2006), and units A–E of Soibelzon et al. (2010). The faunistic content of these units
60 corresponds to Ensenadan Age/stage. The top units D–E, and the calcrete level atop (C 4) are
61 considered as Lujanian sensu lato, and could correlate to units E and F of Tonni & Fidalgo (1982).
62 Unit F is where the cave develops, this unit is transgressive to units D–E, and the surface form
63 where MPH-P 302 was recovered is at the level of unit E, close to C4. Unite F is assigned to the
64 Lujanian sensu stricto Stage/Age given that it correlates to unit G of Tonni & Fidalgo (1982) and
65 unit IV of Alberdi et al. (2006), which correspond to the Equus (Amerhippus) neogeus biozone
66 (Cione & Tonni, 2005). This is consistent with Glyptodon reticulatus shield fragments (MPH-P 034)
67 recovered in the superior part of unit F, above the cave.
68
69 SYSTEMATIC PALEONTOLOGY
70
71 Order CHIROPTERA Blumenbach, 1779
72 Family PHYLLOSTOMIDAE Gray, 1825
73 Subfamily DESMODONTINAE Wagner, 1840
3 74 Tribe DESMODONTINI Wagner, 1840
75 Genus Desmodus Wied-Neuwied, 1826
76 Type species. Desmodus rotundus (Geoffroy, 1810)
77 Desmodus draculae Morgan et al., 1988
78 Figure 3
79 Geographic occurrence. Mexico, Belize, Venezuela, Brazil, Uruguay, and Argentina. 20° N–38° S.
80 Stratigraphic occurrence. late Pleistocene–late Holocene.
81 Referred material. MPH-P 302, right dentary with no preserved teeth.
82 DESCRIPTION
83 A robust right dentary with the corpus mandibulae half the total length of the mandible (LM
84 = 18.79 mm). The margo alveolaris (length of mandible tooth row = 8.63 mm) presents six alveoli
85 (i2, c1, 3 postcanine), the three anterior (i1, i2, c) are circular–subcircular while the following are
86 oval, labiolingually compressed (pm1, pm2, m1). All alveoli have only one root. Posterior to the
87 last one there is a shallow depression.
88 In lateral view (Fig. 3.1) the dentary is excavated (not the coronoid process), with a strong
89 ventral ridge that extends posteriorly from the anterior premolar. In this same view, the most
90 posteromedial part of the symphysis is exposed, projecting posteroventrally a very short distance
91 from the ventral border of the dentary. Two foramina are observed, the anterior (below i2) and the
92 posterior (below pm1) mental foramina. The ramus mandibulae is close to quadrangular in outline.
93 The linea obliqua mandibulae of the coronoid process (= processus coronoideus) raises vertically
94 (close to 20° from the vertical), the coronoid process is of the same height of the condyloid process
95 (processus condylaris/condyloideus). Between these processes the incisura mandibulae is straight,
96 horizontal. The dorsal posterior border of the ramus mandibulae is vertical up to the small non
97 nominatus processus (see Rädulet, 2006 for synonyms) which is located at mid height (above the
4 98 margo alveolarias plane). Ventral to the non nominatus processus the posterior border of the ramus
99 mandibulae is inclined for a short distance.
100 Medially (Fig. 3.2) the symphysis is irregular, interdigitating with its counterpart. There is a
101 diastema from the symphysial articulation to the first incisive. Above the symphysis, lingual to both
102 incisors there is no evident pocket/depression for the upper incisors. In this view two foramina are
103 exposed. An anterior one, very small, on the symphysis below the canine. A second larger one, the
104 mandibular foramen, at the end of a groove in the center of the ramus mandibulae. In this view the
105 coronoid process is excavated. At mid height on the posterior part of the ramus mandibulae is a
106 short, horizontal crest that precedes the non nominatus processus. Ventral to the non nominatus
107 processus the posterior border of the dentary curves medially.
108 DISCUSSION
109 The general morphology of MPH-P 302 resembles that of vampire bats (Desmodontinae).
110 Among extant vampire bats the fossil shares with the Desmodontini (Desmodus and Diaemus) the
111 dentary dental formula (2121), the diastema between incisors, and an unfused symphysis, unlike
112 Diphylla (Allen, 1896; Burns, 1972; Davis et al., 2010; Scheffer et al., 2015). The fossil and
113 Desmodus present the coronoid process as tall as the condyloid process, with a straight, horizontal
114 incisura mandibulae, unlike Diaemus in which the coronoid process is taller than the condyloid
115 process. Furthermore, the diastema between left and right incisors is inferred large, more like that of
116 Desmodus (Scheffer et al., 2015). Desmodus is the only genus of Desmodontinae with known fossil
117 species (D. archaeodaptes, D. draculae, and D. stocki) (see Orihuela (2011) for discussion on D.
118 puntajudensis). Among all Desmodus spp. the fossil shares the large size (Table 2) and, more
119 significantly (autapomorphies), the straight ventral border and absence of pocket/depression behind
120 incisors (to accommodate the upper incisors), with Desmodus draculae (Morgan et al., 1988).
121 Therefore, from the aforementioned, MPH-P 302 is referred to D. draculae.
5 122 Unlike MPH-P 302, previous mid latitude records were assigned to D. draculae with caution
123 (cf. and aff.). This dentary removes the reasonable doubts and confirms the presence of this species
124 at mid latitudes (≈38 °S, Argentina) during the Late Pleistocene. At present the Desmontini have the
125 southernmost distribution of the Desmodontinae, where both Diaemus youngi and Desmodus
126 rotundus reach Argentina (Fig. 1). The southernmost present distribution of the Desmodontinae (D.
127 rotundus) in South America reaches north shore of the Río de la Plata (≈35° S) (Barquez et al.,
128 1999; Queirolo, 2016), more than 400 km north of La Ballenara site (≈38° S). Thus, the
129 environmental conditions of La Ballenera site would have been different from those of today to
130 allow the presence of Desmodus. The wide distributional limits of D. rotundus correlate, in the
131 North and South America, with the 10º C winter isotherm, beyond which cave temperature would
132 require much more energy to maintain normal body temperature (McNab, 1973). Therefore, mean
133 winter temperature during the Late Pleistocene at southeast Buenos Aires should have been at
134 minimum 10° C, a warmer condition than that of today. Associated rodent remains from the same
135 cave filing sediments (Cavia sp. MPH-P 224 and Reithrodon auritus MPH-P 210) suggest and open
136 environment, during a warm humid period, during the Late Pleistocene (Tonni et al., 1999, Pardiñas,
137 2004).
138 Desmodus rotundus is gregarious, know to form large colonies (up to 5000 individuals) with
139 smaller satellite ones, and small groups of up to 10 individuals (Barquez et al., 1999; Sampedro
140 Marín et al., 2008). It is also known to use a wide variate of temporal refuges (Barquez et al., 1999).
141 At least two main types of large refugees where available for bats during the Pleistocene in Buenos
142 Aires province: natural rock (orthoquartzite) shelters in the Tandilia and Ventaian mountain ranges
143 and large megafaunal excavated caves in loess (Dondas et al., 2009). In both of these types of
144 shelters bat fossils have been recovered (Quintana, 2016, this study) (Table 1). According to Frank
145 et al. (2011) in general extant bats are less frequent in sedimentary rock caves (i.e., megafaunal
146 caves) than in plutonic or volcanic ones. In particularly D. rotundus prefers caves with high relative
6 147 humidity and many clustered speleothems (Barros et al., 2020). It is not clear where D. draculae
148 congregated or roosted in the Late Pleistocene of southeast Buenos Aires, but this record suggests
149 that they could have exploited the large caves of giant extinct sloths (Mylodontidae), some of which
150 have been suggested as possible prey of these vampires (Trajano & De Vivo, 1991; Arroyo-
151 Cabrales & Polaco, 2008). An alternative hypothesis is that this dentary corresponds to prey
152 remains of a Strigiform (owls) which are known to prey on vampire bats (Barquez et al., 1999).
153 More data is needed to confirm or reject either hypothesis.
154 In summary MPH-P 302 presents autapomorphies and other characters that allow its referral
155 to D. draculae, and thus confirming the presence of this vampire at mid latitude (≈38° S) during the
156 Late Pleistocene of South America. The presence in southeastern Buenos Aires of D. draculae and
157 the associated rodents is indicative of warmer, a possibly more humid environmental conditions that
158 must have occurred during the last interglacial. This warmer period would have allowed the
159 migration of these taxa from populations further north, as evidenced by rodents as Clyomys sp.
160 (Vucetich et al., 1997), Plesiaguti sp. (Vucetich & Verzi, 2002), Bibimys sp, Scapteromys sp., and
161 Kunsia sp. (Pardiñas et al., 2004) from tropical-subtropical latitudes that also reached 38° S latitude
162 during the warm pulses of the Pleistocene. With the development of the Last Glaciation (85ka,
163 Rabassa et al., 2005) the paleoenvironmental condition would have been adverse (specially
164 regarding temperature) to D. draculae at these latitudes. Despite its wide distribution D. draculae
165 did not survive past the late Holocene, its extinction could be associated, among other things, with
166 the megafaunal extinction, upon which they would have preyed.
167 ACKNOWLEDGMENTS
168 Authors would like to thank G. S. Morgan, N. J. Czaplewski, an anonymous reviewer as well as the
169 editorial team for their comments and suggestion which allowed us to improve our manuscript. We
170 are also a grateful to R. Barquez and P. Gaudioso for facilitating images of reference materials, also
171 to N. J. Czaplewski, J. Arroyo-Cabrales, and C. A. Iudica for facilitating bibliography. We also
7 172 thank D. Boh and C. A. Quintana for their predisposition and assistance. M. Oppedisano assisted us
173 with the SEM. Finllay, SB would like to thank A. Elbakyan. The fossil was recovered under
174 Collection permit 2018-3-P-213-1 of the Dirección Provincial de Museos y Preservación
175 Patrimonial de la Provincia de Buenos Aires.
8 176
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14 308
15 309
310 Figure Captions
311
312 Figure 1. Distribution of Desmodontini in Argentina and fossil bats of Buenos Aires province. 1)
313 Argentina (in grey) with distribution of Desmodus rotundus (opened circles), Diaemus youngi
314 (close circles) and bat fossils of Buenos Aires province (black squares). 2) Closeup of southeastern
315 Buenos Aires fossil bat sites: 1, Desmodus draculae, La Ballenera site (late Pleistocene); 2,
316 Desmodus cf. D. draculae, Centinela del Mar (late Holocene); 3, Noctilio, Necochea (middle
317 Pleistocene); 4, Chiroptera indet., Cueva Tixi (late Holocene). Scale equals 500 km.
318
319 Figure 2. Schematic profile of La Ballenera fossil site.
320
321 Figure 3. Desmodus draculae from the Late Pleistocene of Buenos Aires. Dentary (MPH-P 302)
322 in lateral (1) and medial (2) views. Abbreviations: AMF, anterior mental foramen; CON.P,
323 condyloid process; COR.P, coronoid process; DI, diastema; IN.M, incisura mandibulae; NNP, non
324 nominatus processus; PMF, posterior mental foramen; SYM, symphysis. Scale equals 2 mm.
16
TABLE 1. Fossil bats of Argentina. Taxa Age Locality/Province elements reference Chiroptera indet. Late Holocene Cueva Tixi, Buenos humerus Quintana (2016) Aires (pers. com SB) Desmodus cf. D. draculae Late Holocene Centinela del Mar, isolated C Pardiñas and Tonni Buenos Aires (2000) Histiotus macrotus Late Holocene Several localities in several, not specified Sauthier et al. (2013) Chubut and Santa Cruz Myotis chiloensis Late Holocene Several localities in several, not specified Sauthier et al ., (2013) Chubut
Tadarida brasiliensis Late Holocene Several localities in several, not specified Sauthier et al . (2013) Chubut
? Histiotus Late Pleistocene Epullán Grande Cave, partial left dentary Iudica et al . (2003) Neuquén (MLP-96-V23-1) Desmodus draculae Late Pleistocene La Ballenera, right dentary this study Buenos Aires (MPH-P 302) Noctilio Middle Necochea, Buenos right C Merino et al . (2007) Pleistocene Aires (MLP 07-V-1-1)
Mormopterus barrancae Early Miocene Gran Barranca, partial left dentary Czaplewski (2010) Chubut (MLP-93-IX-18-15);
left m1 (MLP-93-IX- 18-18) Mormopterus sp. Early Miocene Gran Barranca, partial right dentary Czaplewski (2010) Chubut (MLP-93-IX-18-19)
Indet. Phyllostomidae Early Miocene Gran Barranca, right m3 (MLP-93- Czaplewski (2010) Chubut IX-18-16) Indet. family Eocene Laguna Fría, Chubut left m2 (LIEB-PV Tejedor et al . (2005) 999); right talonid (LIEB-PV 1000)
TABLE 2. Comparative measurements of Desmodontinae dentary characters. sp ML LMdT CoH Diaemus youngi 15.1 3.9 - Diphylla ecaudata 13.2±0.2 - - Desmodus rotundus 15.8 ± 0.4 4.8 ± 0.09 6.3 ± 0.26 MPH-P 302 18.79 8.63 8.28 Desmodus draculae 21.9 8,5 8,3 9.4 Desmodus stocki 17.4±0.26 5.3±0.1 6.7±0.21 Data in mm from Morgan et al. , 1988, Barquez et al. , 1999; Davis et al. , 2010. Abbreviations: CoH, Coronoid Height; LM, Length of Mandible; LmdT, Length of mandible tooth row.