Taxonomic status of Andersen’s fruit-eating bat (Artibeus jamaicensis aequatorialis) and revised classification of Artibeus (Chiroptera: Phyllostomidae)
Author
Larsen, Peter A.
Author
Marchán-Rivadeneira, María R.
Author
Baker, Robert J.
text
Zootaxa
2010
2648
45
60
journal article
10.5281/zenodo.198735
08d3d60e-f0e9-40ab-9eec-d5105e6b56f7
1175-5326
198735
Artibeus aequatorialis
Andersen, 1906
Andersen’s Fruit-eating Bat
Artibeus jamaicensis aequatorialis
Andersen, 1906
Artibeus jamaicensis aequatorialis
Andersen, 1908
A
[
rtibeus
].
l
[
ituratus
].
aequatorialis
:
Hershkovitz, 1949
: 447
; name combination.
Holotype
.
BMNH
0.2.9.13: adult male, skin and skull; collected by Perry
O
. Simons, no. 395,
17 June 1899
.
Type
locality.
Ecuador
: Department of El Oro, Zaruma;
1000m
.
Distribution.
West of the Andes Mountains from northern
Peru
, northward throughout western
Ecuador
to
Colombia
(
Fig. 3
). The northernmost and southernmost distribution of
A. aequatorialis
in
Colombia
and
Peru
, respectively, is unknown. Elevation ranges from
22m
(
Ecuador
: Guayas) to
1,106m
(
Ecuador
: El Oro). This species seems to exploit a variety of habitats, ranging from the dry forests of northern
Peru
and southern
Ecuador
to the tropical forests of the Chocó in
Colombia
. Occurs sympatrically with
Artibeus fraterculus
and/ or
A. lituratus
throughout most of its distribution.
TABLE 4.
Revised classification of
Artibeus
. Asterisk indicates
A. schwartzi
may be of hybrid origin (Larsen
et al.
2010).
Artibeus jamaicensis
,
A. lituratus
,
and
A. obscurus
are in bold, indicating potential unrecognized species diversity within Mexican, Central American, and Brazilian populations, respectively (see Guerrero
et al.
2004, Davis 1984, and Redondo
et al.
2008). Note: Simmons (2005) incorrectly cited Allen (1897) for the description of
palmarum
. The original description of
palmarum
was by Allen and Chapman (1897).
Genus
Artibeus
Leach, 1821
Artibeus aequatorialis
Andersen, 1906
A. amplus
Handley, 1987
A. concolor
Peters, 1865
A. fimbriatus
Gray, 1838
A. fraterculus
Anthony, 1924
A. hirsutus
Andersen, 1906
A. inopinatus
Davis and Carter, 1964
A. jamaicensis
Leach, 1821
A. j.
jamaicensis
Leach, 1821
A. j. parvipes
Rehn, 1902
A. j. paulus
Davis, 1970
A. j. richardsoni
Allen, 1908
A. j. triomylus
Handley, 1966
A. j. yucatanicus
Andersen, 1908
A. lituratus
(
Olfers, 1818
)
A. l. intermedius
Allen, 1897
A. l. koopmani
Wilson, 1991
A. l.
lituratus
Olfers, 1818
A. l. palmarum
Allen and Chapman, 1897
A. obscurus
(
Schinz, 1821
)
A. planirostris
Spix, 1823
A. p.
fallax
Peters, 1865
A. p. grenadensis
Andersen, 1906
A. p. hercules
Rehn, 1902
A. p. trinitatus
Andersen, 1906
A. schwartzi
*
Jones, 1978
Specimens examined.
(n = 79) (* indicates specimen used in molecular analyses).
Ecuador
: Azuay, Manta Real Manglar (
2°33'60"S
,
79°21'20"W
), 1 Ψ (MEPN 913072); Bolivar, Barraganate (
1°27'20"S
,
79°19'60"W
), 1 ɗ, 1 Ψ (MEPN 80412, 80460); Carchi, El Pailon (
1°0'7"N
,
78°14'11"W
), 1 ɗ, 2 Ψ (MEPN 871842, 871752, 871850); El Oro, Bosque Petrificado Puyango (
3°53'3"S
,
80°4'41"W
), 2 Ψ (TTU
102596
, 103794 [TK 135290*]), El Progreso (
3°19'60"S
,
79°49'0"W
), 2 ɗ, 3 Ψ (QCAZ 2111, 2113, 2109,
2116–17
),
9 mi
S Zaruma (
3°40'60"S
,
79°37'0"W
), 1 ɗ (TCWC 12283), El Faique (
3°42'7"S
,
79°37'18"W
), 1 Ψ (TTU
102771
[TK 135391*]); Esmeraldas, Borbon comuna Corriente Grande (
0°30'0"N
,
79°58'0"W
), 2 ɗ (QCAZ 2150, 2154), Borbon comuna Viruela (
1°6'51"N
,
78°59'37"W
), 1 ɗ, 3 Ψ (QCAZ 2168, 2173, 2178, 2181), Casa Comunal Mataje (
1°12'4"N
,
78°33'42"W
), 1 Ψ (MEPN 882048), Chontaduro (
0°55'60"N
,
79°23'16"W
), 1 ɗ (MEPN 851433), Comunidad Valle del
Sade
(
0°31'0"N
,
79°22'0"W
), 1 ɗ, 1 Ψ (MEPN 85932, 85971), E San Lorenzo, Banana Plantation (
1°9'11"N
,
78°27'54"W
), 2 Ψ (TTU 85369–70), Esmeraldas (
0°58'60"N
,
79°39'20"W
), 1 ɗ, 2 Ψ (QCAZ 2479, 2466, 2484), Estacion Experimental La Chiquita (
1°13'55"N
,
78°45'57"W
), 5 Ψ (TTU 85284; QCAZ 2344,
2376–77
, 2380), Finca in the road from
Lita
to San Lorenzo (
1°5'16"N
,
78°41'29"W
), 1 ɗ (TTU
103068
), Mataje (
1°21'21"N
,
78°43'27"W
), 2 ɗ, 2 Ψ (TTU
103109
–10 [TK 135905*–TK 135906*],
103107
,
103196
), near to Quingue (
0°43'0"N
,
80°4'60"W
), 1 ɗ, 1 Ψ (QCAZ 6070, 6072), Quininde, Rio
Sade
(
0°19'60"N
,
79°28'0"W
), 1 Ψ (MEPN 85996), Rio Verde (
1°4'0"N
,
79°25'0"W
), 1 ɗ, 1 Ψ (MEPN 851431, 851436), San Francisco de Bogota (
1°4'21"N
,
78°42'41"W
), 2 ɗ (TTU
103180
–81 [TK 135701*–TK 135702*]), San Lorenzo, la Guarapera banana farm pasture (
1°9'41"N
,
78°28'52"W
), 2 ɗ, 3 Ψ (TTU 85409, 85417–19, 85370), Zapallo Grande (
0°46'60"N
,
78°58'60"W
), 1 Ψ (QCAZ 67); Guayas, El Triunfo (
2°19'0"S
,
80°24'0"W
), 1 ɗ, 1 Ψ (QCAZ 2609, 2607), Manglares Churute (
2°27'17"S
,
79°37'39"W
), 3 ɗ, 2 Ψ (TTU 103689, 103692 [TK 134602*], 103712, 103696, 103697), Naranjal,
7 mi
N of Bucay (
1°30'0"S
,
79°54'0"W
), 1 ɗ (AMNH 62934), La Union (
1°58'0"S
,
80°1'0"W
), 4 ɗ, 2 Ψ (QCAZ
2050–51
,
2055–56
, 2049, 2054); Loja,
15 mi
N Catacocha (
4°4'0"S
,
79°37'60"W
), 1 ɗ (TCWC 12282); Los Rios, Buena Fe (
0°54'0"S
,
79°28'60"W
), 1 ɗ (MEPN 81312); Manabi, Cauque River, El Destino (
0°0'0"N
,
80°5'60"W
), 1 ɗ (AMNH 64547), Cerro Pata de Pajaro (
0°1'60"N
,
79°58'0"W
), 1 ɗ (MEPN 913112), Parque Nacional Machalilla (
1°31'0"S
,
80°42'0"W
), 1 Ψ, 1 ɗ (MEPN 964794, 902898), Pedernales (
0°4'60"N
,
80°2'60"W
), 1 Ψ, (MEPN 913113); Pichincha, Bosque Protector La Perla (
0°0'0"N
,
79°22'60"W
), 2 ɗ, 1 Ψ (QCAZ 61, 427, 399), Nanegal (
0°10'10"N
,
78°40'5"W
), 1 Ψ (QCAZ 1998), Santa Rosa, Rio Toachi (
0°19'0"S
,
78°57'0"W
), 1 Ψ (QCAZ 4236), Union del Rio Toachi, Otongachi (
0°25'0"S
,
79°0'0"W
), 1 ɗ (QCAZ 4711).
Emended diagnosis.
Compared with
A. jamaicensis
,
A. aequatorialis
has a larger, broader, and more robust skull and dentition. In skull size proportion
A
.
j
.
richardsoni
is most similar to
A
.
aequatorialis
, while the subspecies
A
.
j
.
paulus
is the smallest form within
A
.
jamaicensis
followed by increased size in
A
.
j
.
parvipes
,
A
.
j
.
yucatanicus
, and
A
.
j
.
jamaicensis
. Nasals are moderately tubular in both
A. jamaicensis
and
A. aequatorialis
, with the orbitonasal shield being concave (
Patten 1971
). The pre- and post-orbital ridges and processes are poorly developed in
A. aequatorialis
, with a less arched and broader appearance of the rostrum with respect to
A
.
jamaicensis
. The zygomatic arch is more slender in
A
.
jamaicensis
. A higher coronoid process and longer distance from the condyle to the coronoid process is present in
A
.
aequatorialis
with respect with
A
.
jamaicensis
, and the angular process is relatively small in both taxa. Dentition is proportional to the skull size, being more robust in
A
.
aequatorialis
. Protocone and hypocone of the first upper molar (M1) are moderately well developed, with a wider talon in
A
.
aequatorialis
. In
A
.
aequatorialis
and
A
.
jamaicensis
, the second upper molar (M2) is distinctly more reduced in size and shape with paracone expanded and a developed labial cingulum (
Patten 1971
). Metacone and metaconule of M2 form a distinct lobe in both species. M1 and M2 are separated by a bigger gap in
A
.
aequatorialis
. The third lower molar (m3) is reduced in size (when present, see below) and does not show well developed cusps in either
A
.
aequatorialis
or
A
.
jamaicensis
. The dental formula in both
A
.
aequatorialis
and
A
.
jamaicensis
is
I 2
/2, C 1/1, P 2/2, M 2/2–3, total 30–32. Of the
134 specimens
examined,
A
.
aequatorialis
and
A
.
jamaicensis
presented two upper molars and few specimens were missing the m
3 in
one or both sides of the mandible. Externally,
A
.
aequatorialis
and
A
.
jamaicensis
are very similar with major differences in body size proportions. For both species, the color of fur varies from pale to dark brown, with lighter color of the ventrum. Both species are characterized by a black patagium and uropatagium (
Patten 1971
).
Artibeus aequatorialis
has been characterized by white wing tips (
Patten 1971
), however, this characteristic is variable and is also present in
A
.
jamaicensis
. Facial stripes are not well defined in
A. aequatorialis
and are variable in
A. jamaicensis
.
Two congeneric species,
A
.
fraterculus
and
A
.
lituratus
, occur in sympatry with
A
.
aequatorialis
. In body and skull size,
A
.
aequatorialis
is intermediate between
A
.
lituratus
and
A
.
fraterculus
.
Artibeus fraterculus
is paler than
A
.
aequatorialis
and is known only from the Pacific slope of central-southern
Ecuador
and northern
Peru
. Nasals are not tubular, M2 is close to M1, and metacone and metaconule are not set apart from the teeth in
A
.
fraterculus
as in
A
.
aequatorialis
(
Patten 1971
)
.
Artibeus lituratus
differs mostly from
A
.
aequatorialis
in having a more robust, larger, and wider skull and dentition, with distinct supraorbital stripes and brownish fur (
Jones 1978
). The pre- and post-orbital constriction is narrower in
A
.
lituratus
, with the optic spicules extended.
FIGURE 3.
Map of the distribution of
Artibeus aequatorialis
. Star indicates the type locality of
A. aequatorialis
(Zaruma [3°40'60"S, 79°37'0"W], Ecuador). The distributional range is based on literature references (hollow squares; Andersen 1906 and Marques-Aguiar 2008) and specimens examined in the Species Account (star and solid squares). The northernmost (western Colombia; Valle del Cauca, Cali [3°26'14"N, 76°31'21"W], and Rio Raposo [3°43'0"N, 77°7'60"W]) and southernmost (western Peru; Tumbes, Faical [3°44'57"S, 80°45'2"W]) extent of
A. aequatorialis
remains to be defined. Shaded areas represent areas of the Andes Mountains higher than 1800 meters.
There has been a historical debate about the recognition of
A
.
planirostris
as a morphologically and genetically defined species distributed east of the Andes in South
America
(see
Larsen
et al.
2007
). With respect to body and skull size proportion,
Artibeus aequatorialis
is smaller than
A
.
planirostris
, and the former differs from
A
.
planirostris
in skull and dentition mainly by: shape of the zygomatic arch (slender =
A. aequatorialis
; robust =
A. planirostris
); development of paracone when compared to metacone of M1 (metacone bigger than paracone =
A. aequatorialis
; metacone smaller than paracone =
A. planirostris
); location of M1 with respect to M2 (M1 and M2 separated by a large gap =
A. aequatorialis
; M1 and M2 without substantial gap =
A. planirostris
); and metacone and metaconule of the M2 (metacone and metaconule separated as a distinct lobe from rest of M2 by a fissure =
A. aequatorialis
; metacone and metaconule not separated as a distinct lobe from the rest of M2 =
A. planirostris
).
Measurements.
Artibeus aequatorialis
is larger than
A
.
jamaicensis
in all fourteen cranial and mandibular measurements, and is generally intermediate between
A
.
fraterculus
and
A
.
lituratus
(Appendix II).