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Aegialomys Galapagoensis

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Mammalian Species 51(981):92–99

Aegialomys galapagoensis (Rodentia: Cricetidae)


Joyce R. Prado and Alexandre R. Percequillo www.mammalogy.org

Departamento de Ciências Biológicas, Escola Superior de Agricultura “Luiz de Queiroz”, Universidade de São Paulo, Av. Pádua
Dias, 11, Caixa Postal 9, 13418-900 Piracicaba, São Paulo, Brazil; joycepra@gmail.com (JRP); percequillo@usp.br (ARP)

Abstract: Aegialomys galapagoensis (Waterhouse, 1839) is a sigmodontine rodent commonly called the Galapagos rice rat. It is
a medium-sized rodent, characterized by dense, soft, yellow or copper, weakly grizzled pelage; the dorsum is grayish or brownish
and the ventral pelage is pale. It is endemic to the open habitats of the Galapagos Archipelago, occurring on three islands, Santa Fé

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Island (Barrington Island), San Cristóbal Island (Chatham Island), and Santiago Island (James Island). Restricted populations are
considered a major threat to A. galapagoensis conservation throughout most of its range, as well as competition with Rattus rattus. A.
galapagoensis is listed as “Vulnerable” by the International Union for Conservation of Nature and Natural Resources, and is listed as
“Critically Endangered” on the Ecuador National list.

Key words: Ecuador, Galapagos, Oryzomyini, rodent, South America

Synonymy completed 1 June 2019


DOI: 10.1093/mspecies/sez013
Version of Record, first published online October 14, 2019, with fixed content and layout in compliance with Art. 8.1.3.2 ICZN.
Nomenclatural statement.—A life science identifier (LSID) number was obtained for this publication: urn:lsid:zoobank.org:pub:
5F03FFB0-26FD-49BB-8C29-AFB1438044A2

Aegialomys Weksler, Percequillo, and Voss, 2006

Mus: Waterhouse, 1839:66. Part, not Mus Linnaeus, 1758; de-


scription of galapagoensis.
Hesperomys: Wagner, 1843:517. Part, not Hesperomys
Waterhouse, 1839; listing of galapagoensis.
Oryzomys: Allen, 1892:48. Part, not Oryzomys Baird, 1857; de-
scription of bauri.
Aegialomys Weksler, Percequillo, and Voss, 2006:5. Type spe-
cies Oryzomys xanthaeolus Thomas, 1894, by original
designation.

Context and Content. Order Rodentia, family


Cricetidae, subfamily Sigmodontinae, tribe Oryzomyini, Fig. 1.—An adult of Aegialomys galapagoensis from Santa Fé Island
genus Aegialomys. The subfamily Sigmodontinae is a mon- (Barrington Island), Galapagos, Ecuador. Used with permission of the
ophyletic lineage, comprising 11 tribes (sensu Salazar- photographer Heidi Snell/Visual Escapes.
Bravo et al. 2016) and 86 genera (sensu D’Elía and Pardiñas
2015). The tribe Oryzomyini is also a monophyletic group distributed throughout arid to semiarid lowland areas of
(sensu Weksler 2006), and currently comprises 29 genera coastal Peru and Ecuador (including some islands of the
and 140 species (Pardiñas et al. 2017), occurring from the Galapagos Archipelago) and also localities up to 2,800 m in
southeastern portion of the United States to southern Chile the Peruvian Andes (Percequillo 2015; Prado and Percequillo
and Argentina. The species of the genus Aegialomys are 2018).

© 2019 American Society of Mammalogists

92
51(981)—Aegialomys galapagoensis MAMMALIAN SPECIES 93

Aegialomys currently contains four extant species (sensu Context and Content. Context as for genus. No subspe-
Prado and Percequillo 2018): A. galapagoensis (Waterhouse, cies are currently recognized (Pardiñas et al. 2017; Prado and
1839; including Oryzomys bauri Allen, 1892), the yellowish rice Percequillo 2018).
rat A. xanthaeolus (Thomas, 1894), Baron's rice rat A. baroni
Nomenclatural Notes. The common name of Aegialomys
(Allen, 1897), and Ica rice rat A. ica (Osgood, 1944). The fol-
galapagoensis is Galapagos rice rat, or in Spanish, Rata Costera
lowing key was prepared with characteristics and measurements
de Galápagos. The generic name is composed of the Greek word
provided by Prado and Percequillo (2018).
aegialos, in reference to its coastal distribution (Weksler et al. 2006).
1. Palatal length long (range: 5.1–7.52 mm). Mesopterygoid fossa Waterhouse (1839) described galapagoensis under the genus Mus,
does not extend anteriorly between the maxillary bones …… a genus taxon name that at that historic time (early 19th century)
…… … ………… … … … .. .… ……………A. xanthaeolus assembled most of the rat-like muroid rodents. Later, galapagoensis
was transferred to the genus Hesperomys, a genus commonly em-

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Palatal length intermediate (range: 4.38–6.60). Mesopterygoid ployed for several species groups of South American cricetids (see
fossa extends anteriorly between the maxillary bones but not Cabrera 1961). Later, Thomas (1884) assigned galapagoensis to
between M3s������������������������������������������������������������������������2 Oryzomys, a subgenus of Hesperomys, and after that Allen (1892)
moved Oryzomys to the generic rank, during the description of a new
2. Length of the dorsal hairs around 11–13 mm (villiform hairs), species from the Galapagos Archipelago, O. bauri (here assigned
18–20 mm (setiform hairs), 27–29 mm (aristiform hairs) …… as a synonym of A. galapagoensis). This scenario remained un-
…… …… …… …… …… …… …… ...A. galapagoensis changed for more than a century, upon the allocation of the taxa
Length of the dorsal hairs around 7–12 mm (villiform hairs), of the species group name (galapagoensis and bauri) to a newly
10–18 mm (setiform hairs), 15–24 mm (aristiform hairs) … described genus Aegialomys by Weksler et al. (2006).
……… ……… ……… ……… …… . .. ….…… ……… .…3

3. Length of the setiform hairs between 10–15 mm��������� A. baroni DIAGNOSIS


Length of the setiform hairs between 16–18 mm ��������� A. ica
Aegialomys galapagoensis can be distinguished from others
species of Aegialomys by its very long, dense, and lax pelage. Its
viliform, setiform, and aristiform hairs are much longer compared
Aegialomys galapagoensis (Waterhouse, 1839) to the other three species of Aegialomys, with modal length about
13, 20, 29 mm, respectively. The other three species of Aegialomys
Galapagos Rice Rat present a tail length larger than the head–body length, while
A. galapagoensis exhibits a tail as long as the head–body length
Mus galapagoensis Waterhouse, 1839:66. Type locality in most specimens, although some specimens present a tail length
“Chatham Island [=San Cristóbal Island], Galapagos slightly shorter or a little longer than the head–body length, but not
Archipelago, Pacific Ocean,” Ecuador. as long as the other species; the tail is also bicolored dorsal-ventrally,
Hesperomys galapagoensis: Wagner, 1843:517. Name as in the other species of Aegialomys (Prado and Percequillo
combination. 2018; Fig. 1). The skull is large and robust (Fig. 2), with one of
Hesperomys (Oryzomys) galapagoensis: Thomas, 1884:453. the greatest overall size of skull in the genus (occipitonasal length
Name combination. Part. range: 31.34–35.85 mm), only comparable to A. ica (occipitonasal
Oryzomys bauri Allen, 1892:48. Type locality: “Barrington length range: 28.76–38.39 mm; A. xanthaeolus occipitonasal length
Island [= Santa Fé Island],” Galapagos Islands, Ecuador. range: 26.38–35.05 mm, and A. baroni occipitonasal length range:
O[ryzomys]. galapagoensis: Thomas, 1894:354. Name 26.92–36.36—Prado and Percequillo 2018). A. galapagoensis
combination. exhibits the more robust upper molar series (mean length of
[Oryzomys] Bauri: Trouessart, 1897:527. Name combination. molars, LM = 5.53 mm, range: 5.35–5.68; mean breadth of the first
[Oryzomys (Oryzomys)] galapagoensis: Trouessart, 1904:419. upper molar, BM1 = 1.74, range: 1.60–1.87) when compared to
Name combination. the other species of Aegialomys (A. xanthaeolus, LM = 4.81 mm,
[Oryzomys (Oryzomys)] bauri: Trouessart, 1904:419. Name range: 4.23–5.31, BM1 = 1.41 mm, range: 1.22–1.60; A. baroni,
combination. LM = 4.99 mm, range: 4.34–5.71, BM1 = 1.52 mm, range: 1.32–
Oryzomys galapagensis: Gyldenstolpe, 1932:23. Incorrect sub- 2.02; and A. ica, LM = 5.42 mm, range: 4.52–6.09, BM1 = 1.64 mm,
sequent spelling of Oryzomys galapagoensis (Waterhouse, range: 1.39–1.95). The mesolophid in the lower molars m1 and
1839). m2 is always present in A. galapagoensis, while in A. baroni, A.
[Aegialomys] galapagoensis: Weksler, Percequillo, and Voss, ica, and A. xanthaeolus, this character can be found only in less
2006:5. First use of current name combination. than 25% of the individuals.
94 MAMMALIAN SPECIES 51(981)—Aegialomys galapagoensis

extending over three ventral scales. Tail scales are generally dark
and about 20 per cm; tail bicolor (dark above and light below).
Pinnae are long and densely haired on internal and external
surfaces. Vibrissae moderately dense, not reaching beyond ears.
Mystacial vibrissae not extending posteriorly beyond the caudal
margins of the pinnae when laid back, and superciliary vibrissae
not extending posteriorly beyond pinnae. Although no sexual di-
morphism in Aegialomys was found by Prado and Percequillo
(2011, 2018), Clark (1980) found significant differences in body
size and weight with males heavier and with longer head–body
length than females.

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DISTRIBUTION

Known localities for Aegialomys galapagoensis occur on


three islands of the Galapagos Archipelago, Santa Fé Island
(Barrington Island), San Cristóbal Island (Chatham Island), and
Santiago Island (James Island). Prado and Percequillo (2018)
presented evidence for the presence of this species in Santiago
Island, but discussed the validity of this record, based on
specimens from the Natural History Museum (London, United
Kingdom). These authors suggest that one specimen may have
been mislabeled, being originally from San Cristóbal Island; the
other specimen was more likely obtained at Santiago Island. The
population of A. galapagoensis in San Cristóbal Island has been
considered extinct since the early 1900s (Heller 1904): recent
surveys performed by Patton and Hafner (1983) and Dowler
et al. (2000) recovered no specimens on this island (Fig. 3).

FOSSIL RECORD

Steadman et al. (1991:131, table 4) reported fossils of


Aegialomys galapagoensis on San Cristóbal Island, associated
with a Holocene vertebrate fauna with an estimated date range
Fig. 2.—Dorsal, ventral, and lateral views of skull and lateral view of
mandible of an adult female Aegialomys galapagoensis (MVZ [the
of 8,500–500 years before present.
Museum of Vertebrate Zoology at Berkeley] 145376) from Santa Fé
Island (Barrington Island), Galapagos, Ecuador. Occipitonasal length
is 35.23 mm. FORM AND FUNCTION

GENERAL CHARACTERS Form.—The external morphology of Aegialomys


galapagoenis follows the general pattern typical of Aegialomys.
Aegialomys galapagoensis is the largest species in the genus Digits II to V of the manus and pes have ungual tufts that are
(Percequillo 2015; Pardiñas et al. 2017; Prado and Percequillo longer than claws, and dI has ungual tufts as long as claws.
2018), similar in size to A. ica, a species from Southern Peru The length of claws is about 1 mm in the manus and about
(Prado and Percequillo 2018). It is a medium-sized rodent (total 2.5 mm in the pes. The pes is long and wide, dorsally covered
body length: 203–350 mm) characterized by very long, dense, with white hairs, with dI and dV smaller than the three cen-
and soft pelage. Dorsal coloration yellow or copper weakly griz- tral digits. The claw of dI extends slightly beyond one-half the
zled; ventral pelage grayish-yellow (Fig. 1). Pelage is a combina- length of the third phalanx of digit II, and digit V with claw
tion of short (11–13 mm) viliform hairs, long (18–20 mm) wavy extending to base of second phalanx of digit IV. The pes has six
setiform hairs, and very long (27–29 mm) and wide aristiform plantar pads, four interdigital at base of digits, one thenar, and
hairs. Tail length (range: 136–165 mm) as long as or slightly one hypothenar (Prado and Percequillo 2018).
shorter or slightly longer than head–body length, and slightly Skull is robust and strongly built. Rostrum is relatively
pilose with hairs on dorsal and ventral regions, apparently long and wide (nasal length range: 11.76–15.09 mm; rostrum
51(981)—Aegialomys galapagoensis MAMMALIAN SPECIES 95

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Fig. 3.—Shading indicates presumed distribution area of Aegialomys galapagoensis based on collecting localities and corresponds (left to right) to
Santiago Island, Santa Fé Island, and San Cristobál Island, Galapagos, Ecuador.

breadth range: 5.6–6.92 mm); zygomatic notch deep and wide; bridge flat (palatal bridge length range: 5.41–6.55 mm, palatal
lacrimal small and in contact with frontal and maxillary. The width range: 6.37–7.60 mm). The mesopterygoid fossa has an
zygomatic arches are robust, divergent posteriorly, wider near anterior margin that is variable in shape and extending above the
the zygomatic root (zygomatic width range: 16.25–19.78 mm). maxillary bones. Auditory bullae inflated, with short and wide
The interorbital region is strongly divergent posteriorly, with Eustachian tube. Mandible is robust, the ramus deep, with coro-
supraorbital margins sharp and acute, forming developed crests noid and condyloid processes well developed. (Fig. 2; Prado and
(interorbital width range: 4.92–5.91 mm). The braincase is elon- Percequillo 2018).
gated with a marked temporal bead. The parietals expand over Upper molar series robust, long, and wide. First upper molar
the lateral surface of temporal region; lambdoidal and occipital with anterocone divided by anteromedian flexus in about 27%
crests are sharp. Rostrum with nasal projected anteriorly, greatly of specimens analyzed by Prado and Percequillo (2018), first
surpassing premaxillary and incisors. Jugal is present. The stape- lower molar with anteroconid undivided; mesoloph present in
dial foramen is very small or absent (in some specimens, it is not the first and second molars; mesolophid present in the first molar
discernible in the suture of the ectotympanic and petrosal; this in all specimens, and in the second molar in 78% of specimens
latter condition was not mentioned by Voss 1988, but reported analyzed by Prado and Percequillo (2018; Fig. 4).
also for other genera of the tribe Oryzomyini, Cerradomys, Fifth lumbar vertebra with well-developed anapophysis;
Drymoreomys, and Sooretamys by Percequillo et al. 2008, hemal arch between second and third caudal vertebrae with spiny
Percequillo et al. 2011, and Chiquito et al. 2014, respectively). posterior process; hemiglandular and unilocular stomach without
Carotid circulatory pattern 3 is present (Voss 1988; Carleton extension of glandular epithelium in the corpus. The phallus
and Musser 1989; Weksler 2003). The basicranial flexion is presents glans penis complex, with small distal and trifurcated
weakly pronounced with foramen magnum oriented caudally. bacular cartilage (with a central, short and thin digit), a pair of pre-
Rostrum with incisive foramen long (occupying most of the di- putial glands present, smooth tissues (not spiny) on margin of ter-
astema), with lateral margins wider medially and anteroposte- minal crater rim do not hide bacular projections; spineless dorsal
rior margins rounded, configuring a long and convex foramen. papilla; urethral processes without subapical lobes; the phallus is
The posterior margins of incisive foramina penetrate between elongated, with a length/diameter ratio of 1.8. The male accessory
first molars in most specimens (ca. 36% in A. galapagoensis, reproductive glands consist of a pair of preputials, bulbourethral,
4% in A. xanthaeolus, 13% in A. baroni, and 48% in A. ica— ampullary, vesiculars, and four pairs of prostate, very similar to
Prado and Percequillo 2018), or leveled with the alveoli of first other oryzomyines (Patton and Hafner 1983).
molars, or almost reaching the alveoli (length of incisive fo- Function.—The shape of the hindfeet, assessed by length
ramen range: 6.42–7.59 mm, width of incisive foramina range: and width, and the length of the claws along with the very large
2.23–2.71 mm); posterior margin of zygomatic plate anterior and fleshy plantar pads (thenar and hypothenar pads large and
to M1; palate intermediate (sensu Weksler 2003) with palatal distinct), with interdigital pads set close together, suggest that
96 MAMMALIAN SPECIES 51(981)—Aegialomys galapagoensis

of the neurocranium, which complement the growth in the early


postnatal life (Voss 1991), are relatively less variable.
The sex ratio of A. galapagoenis was biased toward males,
as males were trapped significantly more than females (Clark
1980). Dowler et al. (2000) recovered similar results, with a sex
ratio favoring males (1.3:1). Harris and MacDonald (2007) also
recovered a bias toward males in the Santiago Galapagos mouse
Nesoryzomys swarthi, another Galapagos endemic oryzomyine
rodent. They hypothesized that transient males caused this asym-
metry, as males and females exhibit similar survival rates, al-
though males present a lower interannual capture rate. Harris and
MacDonald (2007) concluded that transient males could also ex-

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plain the sex ratio biased towards males in A. galapagoensis, and
added, that home range sizes in males are larger in both species.
According to Clark (1980), reproduction occurs in the hot
season, with young individuals collected in May and March,
while Brosset (1963) collected young specimens in March, July,
and December. Sampling in August 1995, Dowler et al. (2000)
observed no lactating females, but reported males with scrotal
testis (11 individuals of 27). The number of embryos per female
was 2–7 (Brosset 1963; Clark 1980), and it was suggested that
Fig. 4.—Occlusal view of the maxillary and mandibular molar rows of
Aegialomys galapagoensis (MVZ [the Museum of Vertebrate Zoology embryo production is limited by the amount of rainfall which
at Berkeley] 125470) from Santa Fé Island (Barrington Island), is associated with food availability (Clark 1980; Harris and
Galapagos, Ecuador. MacDonald 2007). Harris and MacDonald (2007:216) stated
“females adjust their reproductive output according to resource
availability, perhaps through failed implantation or embryo re-
Aegialomys galapagoensis is a terrestrial species with some absorption.” Brosset (1963) revealed that the young are born
scansorial abilities. completely naked and blind; they develop fur and open their
The current concept of the tribe Oryzomyini includes eyes from day 4 to day 9; adult pelage starts appearing at day
pentalophodont, typical forest-specialist genera and also 13. The dynamics of body size revealed that there were two re-
tetralophodont, open-dweller genera (sensu Weksler et al. 2006). cruitment periods throughout the year: young individuals ap-
Some genera in the tribe are transitional habitat specialists, such pear during the warmer season, coupled with the presence of
as Microakodontomys, Oligoryzomys (Hershkovitz 1993), and many large year-old rats. In the nonbreeding season, young rats
Cerradomys (Percequillo et al. 2008), that exhibit polymor- grow and old rats die, compressing the size distributions toward
phism regarding the mesoloph, which is reduced or absent in normality (Clark 1980).
some individuals or species. Similarly, A. galapagoensis is also
a specialist in open, transitional habitats (halophytic vegetation
mixed with cacti forest) and as such exhibits reduced or ab- ECOLOGY
sent mesolophid in some specimens. Therefore, there is a well-
known correlation between the presence of the mesoloph and According to Clark (1980) the population of Aegialomys
mesolophid and habitat, although there is not a clear relation on galapagoensis on Santa Fé Island was remarkably stable, when
their form and function. compared to other Oryzomyini (see also Harris and MacDonald
2007). The number of individuals is highly correlated with the
density of vegetation, and this relation between vegetation den-
ONTOGENY AND REPRODUCTION sity and population density varies seasonally, suggesting that this
species is “resource-limited.” This fact may be related to avail-
There is no sexual dimorphism during age development, and able food and protection from aerial predators, with food being
the ontogenetic variation described for Aegialomys galapagoenis the finite resource, instead of cover (Clark 1980; see also Harris
follows the pattern described for A. xanthaeolus (Prado and and MacDonald 2007). As reported by Prado and Percequillo
Percequillo 2011) and other rodents of the tribe Oryzomyini (see (2018), James Patton (pers. comm.) indicated that he trapped
Carleton and Musser 1989; Voss 1991): the variation is larger in 39 specimens in 100 traps (39%), a large trapping success, at
measurements related to craniofacial and incisor components, as Barrington Cove on Santa Fé Island in only one night. Dowler
these have more conspicuous growth related to ontogenetic devel- et al. (2000), trapping also on Santa Fé Island, obtained a cap-
opment. But the dimensions of the molars and the measurements ture success of 82%, with 69 specimens caught in 84 traps. More
51(981)—Aegialomys galapagoensis MAMMALIAN SPECIES 97

recently, Tirira (2007) also reported this rodent as a common autosomes include two pairs of small meta-submetacentrics, one
species on Santa Fé Island. pair of distinctive and large acrocentrics, and 24 additional pairs
The population is remarkably dense around beaches because from small to large. Concerning the sex chromosomes, the X is a
A. galapagoensis is highly attracted to fish waste left on the large acrocentric and the Y, a small acrocentric.
beaches by fishermen (Brosset 1963). In agreement with Brosset There is little information regarding population genetics of
(1963), Prado and Percequillo (2018:108) reported that Patton’s A. galapagoensis. In a study about population genetics of the na-
specimens were trapped in “rough ground of large boulders cov- tive rodents of the Galápagos Islands, microsatellite and mtDNA
ered by a narrow strip of very dense but low halophytic vegeta- markers were employed to examine genetic differentiation be-
tion about 5–10 m wide before it opens to Opuntia echios forest tween two sampling periods (1997 and 2006—Johnson 2009).
with a sparse understory.” In these areas, A. galapogoensis was Genetic differentiation between the two sampling periods was
found to only occupy shelters under lava blocks and under the high and it is possible that this represents a cyclic population
shrubs of Scutea spicata (Brosset 1963). bottleneck related to El Niño Southern Oscillation events. Also

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Aegialomys galapagoensis exhibits great longevity: the high haplotype diversity and low nucleotide diversity suggest
maximum survival in the wild was 599 days for one male and closely related haplotypes in the populations.
one female. The median survival time for recaptured animals Prado and Percequillo (2018) recovered A. galapagoensis
was 165 days for males and 167 days for females, and 37% of as sister to all other species of the genus employing Cytb and
these individuals were alive 365 days after first capture, and 13% Adh1-I2 genes and as sister to the two southern forms of the
were alive after 500 days (Clark 1980). genus, A. baroni and A. ica, with only the Adh1-I2 marker.
Clark (1980) hypothesized that A. galapagoensis, exhibiting Despite this incongruence, Prado and Percequillo (2018) es-
high survival and low reproduction, could be categorized as a tablished that A. galapagoensis represents a unique lineage,
pulse averager, a demographic trait exhibited by desert rodents of falsifying the hypothesis of Patton and Hafner (1983) that
the family Heteromyidae. These pulse averagers “are temporally sailors brought Aegialomys from Galapagos to the island from
fixed, once-a-year breeders that rely on general predictability of the Peruvian coast.
environmental seasonality” (Harris and MacDonald 2007:208).
This hypothesis was confirmed by studies conducted by Harris and
MacDonald (2007) with the Santiago Galapagos mouse, which CONSERVATION
exhibits a demographic history very similar to A. galapagoensis.
Aegialomys galapagoensis has been described as predomi- Charles Darwin was the first collector of Aegialomys
nantly nocturnal, although during reproductive season it becomes galapagoensis, and about it he commented “This mouse or rat is
more active during the day (Brosset 1963); however, Dowler abundant in Chatham Island, one of the Galapagos Archipelago.
et al. (2000) reported that individuals were active, running I could not find it on any other island of the group. It frequents
through lava fields before dark, with some captures occurring the bushes, which sparingly cover the rugged streams of basaltic
during the day. A. galapagoensis is omnivorous, but has a pref- lava, near the coast, where there is no fresh water, and where the
erence for insects and fish remains (Brosset 1963). According land is extremely sterile” (cited in Waterhouse, 1839:66). It now
to Tirira (2007), researchers have to maintain their belongings has been pointed out in the literature (Heller 1904; Patton and
(including food) in metal containers to keep them protected from Hafner 1983; Tirira 2001, 2011) that the population from San
A. galapagoensis, as otherwise it will eat or gnaw at the items. Cristobál Island is extinct, and except for data associated with
Data on captivity show that in a period with less food they may the specimen that Darwin secured all available data is from the
practice cannibalism (Brosset 1963). population of Santa Fé Island.
The predation of a young individual of A. galapagoensis On Santa Fé Island, A. galapagoensis is considered
by a centipede (Scolopendra galapagoensis) on Santa Fé Island “Vulnerable” by the International Union for Conservation of
has been reported (Clark 1979). The specimen inside the nest Nature and Natural Resources (Weksler 2018). This listing
was only slightly furred, with closed eyes, and about 40 mm follows D2 criterion which describes populations that occupy a
of body length.. The centipede measured about 200 mm and it limited number of locations (typically five or less) or the pop-
held the rat in its anterior appendages, carrying the individual ulation occupies restricted areas (typically less than 20 km2).
out of the nest. A. galapagoensis has a host-specific ectopar- In addition, the restricted population “is prone to the effects
asite, the laelapine mite (Acari: Mesostigmata: Laelapidae) of human activities or stochastic events within a very short
Gigantolaelaps galapagoensis (Gettinger et al. 2011). time period in an uncertain future, and is thus capable of be-
coming Critically Endangered or even Extinct in a very short
time period” (International Union for Conservation of Nature
GENETICS and Natural Resources Standards and Petitions Subcommittee
2017:69). In the Libro Rojo de los Mamiferos del Ecuador
The karyotype of Aegialomys galapagoensis has a diploid (Tirira 2011), this species is currently considered “Critically
number (2n) of 56 chromosomes and a fundamental number Endangered,” under the criteria B1ac(i), that is related to ex-
(FN) for autosomes of 58 (Gardner and Patton 1976). The treme fluctuations in extent of occurrence. Jiménez-Uzcátegui
98 MAMMALIAN SPECIES 51(981)—Aegialomys galapagoensis

et al. (2007) stated that the major threat to this species is Baird, S. F. 1857 [1858]. Mammals. In Reports of explorations and surveys
introduced rats, although on this island the presence of such to ascertain the most practicable and economical route for a railroad
from the Mississippi River to the Pacific Ocean. Part 1. General re-
rats was not recorded. The Charles Darwin Foundation and port upon the zoology of the several Pacific railroad routes. Beverly
Galapagos National Park Service monitors the presence of the Tucker, Washington, D.C., 8, 1–757, 60 plates.
black rat (Rattus rattus) on islands with endemic native species Brosset, A. 1963. Statut actuel des mammifères des îles Galapagos.
(Jiménez-Uzcátegui et al. 2007). Mammalia 27:323–341.
Cabrera, A. 1961. Catalogo de los mamiferos de America del Sur.
Brosset (1963) stated that the two elements that promote Revista Museo Argentino Ciencias Naturales (Ciencias Zoologicas)
extinction of this species were the considerable population of 4:309–732.
feral cats and the population of R. rattus. But according to Clark Carleton, M. D., and G. G. Musser. 1989. Systematic studies of oryzomyine
rodents (Muridae, Sigmodontinae): a synopsis of Microryzomys. Bulletin
(1980) this species is not a threatened species: he estimated that of the American Museum of Natural History 191:1–83.
the population numbers vary between 104 and 105 individuals Chiquito, E. A., G. D’Elía, and A. R. Percequillo. 2014. Taxonomic
and he believes that the only danger to A. galapagoensis would

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review of genus Sooretamys Weksler, Percequillo & Voss (Rodentia:
be the presence of R. rattus. Further, the great temporal differen- Cricetidae: Sigmodontinae): an integrative approach. Zoological
Journal of the Linnean Society 171:842–877.
tiation found in Johnson (2009) following an El Niño Southern Clark, D. B. 1979. A Centipede preying on nestling rice rat (Oryzomys
Oscillation event may be a signal of the potential impacts of bauri). Journal of Mammalogy 60:654.
global climate change. Clark, D. B. 1980. Population ecology of an endemic Neotropical island
As all modern records are from Santa Fé Island (one rodent: Oryzomys bauri of Santa Fe Island, Galapagos, Ecuador.
Journal of Animal Ecology 49:185–198.
of the few islands without rodent invasive species of genus D’Elía, G., and U. F. J. Pardiñas. 2015. Subfamily Sigmodontinae
Rattus), there are some concerns about the conservation of Wagner, 1843. Pp. 63–73 in Mammals of South America, vol. 2,
A. galapagoensis. Dowler et al. (2000) stressed that there are few rodents (J. L. Patton, U. F. J. Pardiñas, and G. D’Elía, eds.). University
data for the ex situ management of this species (and other en- of Chicago Press, Chicago, Illinois.
Dowler, R. C., D. S. Carroll, and C. W. Edwards. 2000. Rediscovery
demic species of Galapagos, as the small Fernandina Galapagos of rodents (genus Nesoryzomys) considered extinct in the Galapagos
mouse Nesoryzomys fernandinae), and that such programs Islands. Oryx 34:109–118.
should be started as soon as possible to protect A. galapagoensis Gardner, A. L., and J. L. Patton. 1976. Karyotypic variation in
from extinction. oryzomyine rodents (Cricetidae) with comments on chromosomal
evolution in the Neotropical cricetinae complex. Occasional Papers
Museum of Zoology. Louisiana State University 49:1–48.
Gettinger D., F. Martins-Hatano, and S. L. Gardner. 2011. Some
ACKNOWLEDGMENTS Laelapine Mites (Acari: Laelapidae) ectoparasitic on small mammals
in the Galapagos Islands, including a new species of Gigantolaelaps
from Aegialomys galapagoensis. Journal of Parasitology 97:574–576.
We are thankful to the editor of the Mammalian Species, Gyldenstolpe, N. 1932. A manual of Neotropical sigmodont rodents.
namely Meredith J. Hamilton, and to the reviewers for their Kungliga Svenska Vetenskapsakademiens Handlingar 11:1–164.
valuable comments on the manuscript. We are also grateful Harris, D. B., and D. W. MacDonald. 2007. Population ecology of the
endemic rodent Nesoryzomys swarthi in the tropical desert of the
for all the curators that provided access for their specimens we Galápagos Islands. Journal of Mammalogy 88:208–219.
examined in our review of the genus Aegialomys; and for Heidi Heller, E. 1904. Mammals of the Galapagos Archipelago, exclusive of
Snell, for kindly providing the photo of the living specimen of the Cetacea. Proceedings of the California Academy of Sciences
3:233–251.
A. galapagoensis. We are also indebted for the financial sup- Hershkovitz, P. 1993. A new central Brazilian genus and species of
port provided by the São Paulo state agency of research sup- sigmodontine rodent (Sigmodontinae) transitional between akodonts
port Fundação de Amparo à Pesquisa do Estado de São Paulo and oryzomyines, with a discussion of muroid molar morphology and
(acronym: FAPESP) and the Brazilian agency of research evolution. Fieldiana Zoology 75:1–18.
International Union for Conservation of Nature and Natural
support Conselho Nacional de Desenvolvimento Científico e Resources Standards and Petitions Subcommittee. 2017.
Tecnológico (acronym: CNPq); collection study grants from Guidelines for using the IUCN Red List Categories and Criteria.
the American Museum of Natural History, the United States Version 13. Prepared by the Standards and Petitions Subcommittee.
Natural History Museum, the Field Museum and Museum of http://www.iucnredlist.org/documents/RedListGuidelines.pdf.
Accessed 1 July 2019.
Comparative Zoology, allowed the senior author to examine Jiménez-Uzcátegui, G., V. Carrión, J. Zabala, P. Buitrón, and
specimens and type specimens of this genus back in 2001, B. Milstead. 2007. Status of introduced vertebrates in Galapagos.
which would became the beginning of this contribution. Pp. 136–141 in Galápagos report 2006–2007. Charles Darwin
Foundation. Puerto Ayora, Galápagos, Ecuador.
Johnson, S. 2009. Population genetics of the native rodents of the
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