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110
n
Monitoring Biodiversity on a Trans-Andean Megaproject
*Correspondence: H. Zamora Meza, tommyzm@gmail.com.
The Peruvian Long- Snouted Bat, Platalina genovensium
Thomas, 1928 (Phyllostomidae, Lonchophyllinae),
in the Area of Influence of the PERU LNG Gas Pipeline:
Population Status and Recommendations for Conservation
Hugo T. Zamora Meza*, Catherine T. Sahley, César E. Medina Pacheco,
Yamileth E. Arteaga Miranda, Alain C. Escobar Montes, and Alexander Pari Chipana
ABSTRACT. As part of the biodiversity monitor-
ing and assessment program developed jointly by the
Center for Conservation Education and Sustainabil-
ity of the Smithsonian Institution and PERU LNG,
we carried out a study on the Peruvian long-snouted
bat, Platalina genovensium. We assessed potential
impacts of the construction of the gas pipeline on the
population of this species, which is found associated
with cacti in the Río Seco Basin in Ica, Peru. Data
were gathered in the dry and rainy seasons in three
sampling areas, which included plots near the pipe-
line (impact) and plots far from the pipeline (nonim-
pact) between 1,100 and 2,900 m. We used 10 mist
nets during four consecutive nights in each of the
sampling points and a harp trap installed at the en-
trance of a roost near sampling points. We recorded
data on the presence of owers and fruits of columnar
cacti in the area (Browningia candelaris, Armatocer-
eus procerus, Weberbauerocereus rauhii, and Neo-
raimondia arequipensis). We also took fecal samples
to analyze the diet of P. genovensium to obtain infor-
mation about potential factors that might be inuenc-
ing the presence and abundance of the species. After
four eld seasons of assessment and a total sampling
effort of 5,592 hours/net, we recorded two bat spe-
cies (P. genovensium and Tomopeas ravus) and cap-
tured and marked 39 individuals of P. genovensium.
The average abundance was 0.07 individual/net hour,
with a maximum capture in December 2009 (0.18 in-
dividual/net hour) and a minimum capture in March
2010 (0.03 individual/net hour). We obtained seven
recaptures, with one individual recorded at 3.49 km
from the original marking point and another individ-
ual found after 6 months. We conclude that P. geno-
vensium is strongly associated with columnar cacti,
and during our study, they fed on A. procerus and
W. rauhii. We found no statistical differences in bat
populations in impact and nonimpact areas along the
pipeline.
KEY WORDS: Platalina genovensium, columnar
cacti, desert ecosystem, bat pollination.
Introduction
Peru has 165 species of bats that represent 32% of the
total number of mammals recorded for the country
(Pacheco et al., 2009). Among these, 95 species are
members of the family Phyllostomidae, with seven
species belonging to the nectarivorous subfamily
Lonchophyllinae, which includes Platalina genoven-
sium (Thomas, 1928; Gardner, 2008; Pacheco et al.,
2009). The members of Lonchophyllinae, like the
members of Glossophaginae, are characterized by
having morphologic and physiologic characteristics
that are adaptations for the consumption of ower
nectar as one part of their diet (Freeman, 1995; Sim-
mons and Wetterer, 2002). The adaptations of both
of these subfamilies include an elongated rostrum,
reduced dentition, reduced size and number of teeth,
and a long tongue with papillae for the collection of
nectar from owers (Winkelman, 1971; Freeman,
1995), which allows them to play an important role in
the pollination of a signicant number of plants in the
Neotropics (Fleming et al., 2009). The benets of bat
pollination can include large amounts of pollen de-
posited per ower and long- distance pollen dispersal
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Zamora Meza et al. / Bats in the Area of the Pipeline n 111
disperser of columnar cacti reveal that it performs
these services for W. weberbaueri (Sahley, 1995;
Sahley and Baraybar, 1996; Baraybar, 2004). Ob-
servation of ower visits, diet studies via analysis
of fecal pellets, and the presence of 13C isotopes
in muscle tissue demonstrate that P. genovensium
has a diet based on crassulacean acid metabolism
(CAM) plants. These data indicate that this species
is a cactus- feeding specialist, at least in the southern
department of Arequipa (Sahley, 1995; Sahley and
Baraybar, 1996; Baraybar, 2004). Given that, many
columnar cacti of the western Andean slopes dem-
onstrate adaptation for bat pollination (Sahley, 1995;
Sahley et al., in press), and P. genovensium is likely
one of the most important pollinators and seed dis-
persers of several species of columnar cacti that are
considered keystone species in desert ecosystems.
In ecological landscape unit (ELU) 12 (Lang-
stroth et al., this volume) several species of colum-
nar cacti occur, and P. genovensium was reported to
occur in this area (Walsh Perú, 2005). Moreover, the
species of columnar cacti found in ELU 12, such as
A. procerus, N. arequipensis, W. rauhii (all of which
are endemic), and B. candelaris, present adaptations
that suggest chiropterophily. From these, W. rauhii
has been classied by the IUCN as vulnerable (Ara-
kaki et al., 2006).
In this area, Walsh Perú (2005) recommended
that the pipeline right of way be placed, where pos-
sible, along ridges, where cactus densities in general
are lower than in creek valleys. As part of the pipe-
line construction phase it was recommended that
hundreds of columnar cacti be removed from the
right of way and transplanted to nearby areas (Envi-
ronmental Resources Management Peru, 2008). The
removal and transplant process may have caused
stress to columnar cacti, affecting production of
owers and fruits or survival of some individuals.
This may, in turn, have affected the abundance of
species such as P. genovensium that feed on these
cacti. Therefore, our study had the following objec-
tives: (1) to determine the spatial and temporal distri-
bution of P. genovensium in the area of inuence of
the PERU LNG pipeline right of way (ROW), (2) to
determine the abundance of P. genovensium, (3) to
determine if the placement of the pipeline ROW had
an impact on P. genovensium populations within the
study area, and (4) to review the status of this spe-
cies and give recommendations for its monitoring
and conservation.
(Heithaus et al., 1974; Bertin, 1982; Sahley, 1995,
2001; Muchala and Thompson, 2010).
Unfortunately, the number of species and genera
of plants that depend on bat pollination have not been
quantied in Peru, especially in arid zones (Sahley
et al., in press). In desert ecosystems, numerous co-
lumnar cacti show oral adaptations to attract bats
(known as the chiropterophilous syndrome; Faegri
and Van der Pijl, 1966; Vogel, 1969) and depend on
them for pollination. In arid ecosystems new- world
nectar- feeding bats apparently have a stronger associ-
ation with plants than in humid ecosystems (Porsche,
1939; Vogel, 1969; Sahley, 1995; Fleming and Much-
hala, 2008). The specialized ecological associations
of nectar- feeding bats highlight the need to conserve
them, particularly in desert ecosystems where they
may play a critical role in the reproductive success of
columnar cacti.
The Peruvian long- snouted bat, Platalina geno-
vensium Thomas, 1928, Phyllostomidae, subfamily
Lonchophyllinae (Gardner, 2008), is a monotypic
species and one of two species of bats currently
classied as critically endangered by Peruvian leg-
islation (Supreme Decree 034- 2004- AG) and as near
threatened by the International Union for Conserva-
tion of Nature (IUCN, 2011). Its distribution range
encompasses the western slopes of the Andes of Peru
from the north in Piura to the south in the depart-
ment of Tacna (Ortiz de la Puente, 1951; Aellen,
1965; Jiménez and Pefaur, 1982; Sahley and Ba-
raybar, 1996; Aragón and Aguirre, 2008). Popula-
tions have been found in the inter- Andean valley of
Huánuco (Sanborn, 1936; Tuttle, 1970), which is the
only area where this species has been found on the
eastern slopes of the Andes. There are three speci-
mens reported for Arica, Chile (Galaz et al., 1999),
but Galaz et al. do not conrm the presence of vi-
able and resident populations in the north of Chile.
Distribution localities and observation indicate that P.
genovensium is strongly associated with desert eco-
systems that contain columnar cacti, such as Weber-
bauerocereus weberbaueri, Browningia candelaris,
Neoraimondia arequipensis, Coryocactus brevisty-
lus, Echinopsis chiloensis, Armatocereus procerus,
and Weberbauerocereus rauhii (Sahley and Baraybar,
1996; Ugarte and Salazar, 1998; Zeballos and López,
2002; Baraybar, 2004; Zamora, 2005; Aragón and
Aguirre, 2008; this study).
The few studies conducted that investigate
the role of P. genovensium as a pollinator and seed
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Monitoring Biodiversity on a Trans-Andean Megaproject
nearness to the pipeline right of way, each of the three
study areas (Ticcoc, Los Olivos, and San Juan) was
divided into two sites: one adjacent to the right of
way (impact site) and one farther from the right of
way (control site; Figure 1, Table 1). Mist nets were
used in each study area (for a total of six sites, includ-
ing impact and control sites) to determine the relative
abundance of P. genovensium. For each study area, 10
mist nets (12 m long by 2.6 m tall) were placed for
four consecutive nights during the new moon. Each
night was considered a replicate, as was each mist
net. Total capture effort was 5,592 hours/net. We also
utilized a harp trap to census P. genovensium bats in
an abandoned mine that was used as a roost.
To determine if there was a relation between the
number of individuals of P. genovensium and the
proximity of the pipeline ROW in each of the three
study areas (Ticcoc, Los Olivos, and San Juan), we
chose two sites within each area, one near the ROW
and one outside the presumed area of impact. These
Study Area
The study area comprises the dry creek valleys of Tic-
coc, San Juan, and Los Olivos in ELU 12 (called the
Coastal Bat Hills), between 1,094 and 2,566 m above
sea level (asl), in the district of Humay, province of
Pisco, department of Ica, Peru (Figure 1). The area of
study is characterized by a heterogeneous composition
of columnar cactus because of differences in altitude
among the three sites (Table 1). Columnar cacti pres-
ent in the study area include A. procerus, W. rauhii,
B. candelaris, and N. arequipensis in addition to other
ornithophilous species of cactus (adaptations for hum-
mingbird pollination), such as Cleistocactus hystrix
and C. peculiaris (Novoa Sheppard, this volume).
Methods
To determine if there was a relationship between
the abundance of P. genovensium individuals and
FIGURE 1. Map of the study area. Triangles are impact sites, and circles are control sites. The star indicates the location of the
conrmed roost site. (ICA is name of region; Qda. is quebrada [stream].)
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Zamora Meza et al. / Bats in the Area of the Pipeline n 113
roost was monitored with a harp trap installed at the
entrance of the mine.
Fruits of each cactus present in study areas were
collected in order to establish a reference collection for
later diet analysis using fecal samples. We analyzed
seven fecal samples from seven individuals captured
and recorded the presence of fruit pulp, seeds, pollen,
and insects using a stereomicroscope. We recorded the
presence or absence of owers and fruits in December
2009. In June and October 2010, we recorded phenol-
ogy data for 20 individuals of each species of colum-
nar cactus present at each of our census sites.
To determine if there were statistically signi-
cant differences in P. genovensium abundance be-
tween rainy and dry seasons and between control and
impact sites, we compared the number of individuals
captured between seasons and then sites using the
Mann- Whitney U statistic (StatPlus for Mac version
2009). We also compared mean weight and forearm
length between males and females using a t test after
testing data for normality.
Results
Morphology and Reproductive Condition
The mean weight of P. genovensium adults was 19.35
g (n = 23, standard deviation [SD] =1.86), and the
mean forearm length was 50.27 mm (n = 23, SD =
0.76). There was no signicant difference in weight
between adult males and females (t = 2.07, n = 23,
p = 0.78), but we did nd that females have a slightly
longer forearm length than males (for females, n = 12,
= 50.53 mm, SD = 0.77; for males, n = 11, = 49.93,
SD = 0.63; both sexes, n = 23, t = 2.07, p = 0.038).
We captured a similar proportion of males and
females during the entire study, although in the
sites were labeled impact and control sites (Figure
1, Table 1). We conducted two censuses during the
rainy season (December 2009 and March 2010) and
two during the dry season (June and October 2010)
in each study area and at each type of site (control
and impact).
We used mist nets in the study areas and a harp
trap near the conrmed refuge. Relative abundance
was calculated on the basis of the number of indi-
viduals captured at each site and the capture effort
expended (480 hours per net). Mist nets were placed
near columnar cacti that had owers and/or fruits or
were in potential movement corridors (narrow val-
leys and dry creek beds). Mist nets were opened from
6:00 to 12:00 and were checked every half
hour for the presence of bats. Because of security
policies implemented by PERU LNG, we were un-
able to census bats all night long. We recorded all
species of bats captured. Every individual captured
was individually marked by trimming fur on the cau-
dal part of the back and by making a small perfora-
tion in the wing membrane between the ngers. We
recorded length of forearm (in millimeters), weight
(grams), age (based on the ossication of third and
fourth digits), sex, and reproductive condition (based
on presence or absence of descended testicles in
males and vaginal opening and teat development in
females). After data were recorded, each individual
was photographed and set free.
During December 2009, 11 sites (abandoned
mines or naturally occurring tunnels) were identied
that could have served as potential roost sites for P.
genovensium. We conrmed the presence of P. geno-
vensium in one of these, an abandoned mine, located
in the San Juan site (Figure 1). The rest of the poten-
tial roost sites were not censused because of security
policies implemented by PERU LNG. The conrmed
TABLE 1. Characteristics of study areas by treatment.
Study area Treatment Altitude (m asl) Columnar cactus species
Ticocc Impact 1 2,819–2,940 Browningia candelaris
Control 1 2,460–2,710 B. candelaris, Weberbauerocereus rauhii
San Juan Impact 2 1,800–1,836 Neoraimondia arequipensis
Control 2 1,379–1,510 Armatocereus procerus, N. arequipensis
Roost
a 1,300 A. procerus, N. arequipensis
Los Olivos Impact 3 1,073–1,146 A. procerus, N. arequipensis
Control 3 1,122–1,236 A. procerus, N. arequipensis
a Only one roost was found in the study area.
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Monitoring Biodiversity on a Trans-Andean Megaproject
and 11 were captured with harp nets outside roosts.
With the exception of one individual of Tomopeas
ravus (Molossidae; critically endangered, H. Zamora
Meza, unpublished) no other species of bats were
captured. Therefore, P. genovensium is the only po-
tential bat pollinator and/or seed disperser in the area.
In December 2009, we captured 18 individuals
(using both mist nets and harp traps). This was the
largest number of individuals captured during the
entire study. In the months of March, June, and Oc-
tober 2010 we captured 4, 10, and 7 individuals, re-
spectively. The total number of captures in the rainy
season (December and March) was 22, whereas in
the dry season it was 17, but the difference was not
statistically signicant (Mann- Whitney U = 71.5, n
= 24, p = 0.35). The average relative abundance over
the entire study was 0.07 individuals/net hour, with a
maximum of 0.181 individuals/net hour in December
2009 and a minimum of 0.03 individuals/net hour in
March of 2010 (Table 2).
Abundance data in our study site (Figure 3)
present differing patterns of use in impact and con-
trol sites. At Ticcoc (2,460–2,940 m asl), where B.
candelaris and W. rauhii were present, we only cap-
tured individuals in control sites in December 2009
and October 2010, and we never captured individu-
als in the impact zone (Figure 3). In the San Juan
area (1,379–1,836 m asl), where N. arequipensis and
A. procerus were present, we obtained the greatest
number of captures, with 12 individuals. As in the
other study areas, after December 2009 the number
of individuals captured decreased (a maximum of 5
in December 2009 and a minimum of 1 in October
2010; Figure 3). Only one individual was captured
in the impact zone (in December 2009). In the Los
Olivos area (1,073–1,235 m asl), where N. arequipe-
nsis and A. procerus were present, we captured the
highest number of individuals (11) in our study in the
month of December 2009. In Los Olivos, however,
we observed a different pattern of captures in control
and impact sites. We captured more individuals in the
rst month (December 2009), we captured slightly
more males than females, whereas in the remaining
months, we captured a slightly higher number of fe-
males (Figure 2a). We captured both adults and juve-
niles during all months sampled (Figure 2b). Three
males with descended testicles were captured in De-
cember 2009, although the adult females captured did
not have signs of being reproductively active (open
vagina or developed teats). In March 2010, one male
had descended testicles, one female was in the early
stages of pregnancy, and one female was lactating.
In June 2010 (dry season), we recorded the pres-
ence of four females with open vaginas, one lactating
female, and three males with descended testicles. In
October 2010, the end of the dry season, we recorded
three males with descended testicles, but we did not
capture females that showed evidence of being repro-
ductively active.
Abundance
We captured 39 individuals of P. genovensium during
the present study; 28 were captured using mist nets,
FIGURE 2. Population composition of Platalina genoven-
sium in our study area based on (a) sex and (b) age.
TABLE 2. Relative abundance of Platalina genovensium in
our study, expressed in individuals per net hour.
Study area Dec 2009 Mar 2010 Jun 2010 Oct 2010
Ticcoc 0.02 0 0 0.03
San Juan 0.06 0.04 0.02 0.01
Los Olivos 0.101 0 0.01 0
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Zamora Meza et al. / Bats in the Area of the Pipeline n 115
Activity and Movement Patterns
Between 8:00 and 11:00 we recorded two activ-
ity peaks, with a total of 12 individuals captured at
the most active time (8:00 ), and no individuals
captured at 12:00 (Figure 4). We recaptured seven
individuals of P. genovensium, four in Los Olivos and
three in San Juan. The largest linear movement re-
corded was the recapture of an individual captured
in San Juan and recaptured four days later at 3.49 km
impact site (10) than in the control area (1; Figure 3).
Combining the data from the three areas and compar-
ing the number of P. genovensium individuals in im-
pact and control sites, we captured more individuals
in control sites (17) than in impact sites (11), which
suggests that the foraging patterns of P. genovensium
in our study area may have been inuenced by the
pipeline right of way. However, we did not nd a sta-
tistical difference between impact and control sites
(Mann- Whitney U test = 56, n = 24, p = 0.97).
FIGURE 3. Number of individuals of Platalina genovensium captured in impact
(black bars) and (gray bars) control sites in Ticcoc, San Juan, and Los Olivos.
FIGURE 4. Activity of Platalina genovensium based on mist net capture data.
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Monitoring Biodiversity on a Trans-Andean Megaproject
2009, 3 captures in March 2010, 3 captures in June
2010, and 4 captures in October 2010). These data
suggest that after December 2009 the amount of po-
tential food resources for P. genovensium may have
been below a threshold that could support the maxi-
mum population censused and that bats may have
increased their foraging areas or dispersed to areas
out of our study area.
Diet
Seven fecal samples from seven individuals captured
in mist nets were collected (Table 3). Of these, one
sample contained only pollen, one contained only
fruit pulp, two contained both pollen and fruit pulp,
and one contained seeds of A. procerus, pollen, and
fruit pulp. Another contained pollen and a seed husk,
and nally, the last sample contained the remains of a
deteriorated insect.
linear distance from the site of original capture, at the
abandoned mine where bats were roosting (Figure 5).
We also recaptured an individual after six months,
rst in December 2009 and then again in June 2010.
Cactus Phenology
In June and October 2010, we recorded the phenol-
ogy of columnar cacti in our study areas. In these
months we did not observe any owers or fruits of B.
candelaris (n = 20) and only one ower of N. arequi-
pensis (n = 20). The columnar cacti that produced
a greater number of owers and fruits during these
months were A. procerus and W. rauhii (Figure 6). In
June of 2010, there was greater production of fruits
and owers than in October 2010 for A. procerus and
W. weberbaueri (Figure 6). The number of captured
bats was low but consistent in these months com-
pared to December 2009 (18 captures in December
FIGURE 5. Recaptures and estimated longest (a, c) and shortest (b, d) linear distances traveled by Platalina genovensium at
(a, b) the San Juan site and (c, d) Los Olivos; D is linear distance from point of capture (dotted icon) to point of recapture (solid
icon). Codes indicate zone (Z) and treatments: control (C) or impact (I); numbers indicate the individual specimens, and the
letter indicates point of recapture.
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Zamora Meza et al. / Bats in the Area of the Pipeline n 117
(Sahley, 1996; Sahley and Baraybar, 1996; Baraybar,
2004; Zamora, 2005). Its altitudinal distribution coin-
cides with other published studies as well (Table 4).
In our study area, P. genovensium shares its habitat
with the insectivorous T. ravus and possibly other in-
sectivorous species not captured.
Morphology
Forearm and body weight measurements of captured
individuals in this study are similar to those of indi-
viduals measured in studies conducted in Arequipa
(Sahley, 1995; Sahley and Baraybar, 1996; Baraybar,
2004) and in the seasonally humid lowland habitats
of Atiquipa (Zamora, 2005) and Tacna (Aragón and
During the time that fecal samples were col-
lected, only the columnar cacti A. procerus and W.
rauhii were producing owers and fruits. These sam-
ples indicate that in our study area and in the months
sampled, P. genovensium feeds primarily from these
two species of columnar cacti.
Discussion
Our study provides data on new populations of P.
genovensium for south central Peru in the Humay Dis-
trict, Pisco Province. On the basis of 28 captures with
5,592 net hours, our data conrm that P. genovensium
has a relatively low abundance (0.07 individual/net
hour), which coincides with other published studies
TABLE 3. Diet of Platalina genovensium based on analysis of fecal pellets; Fl, cactus ower; Fr, fruit; a
dash (–) indicates not applicable.
Date Study area Material found Cactus species
Dec 2009 Ticcoc Deteriorated insect remains –
Los Olivos Pollen and cactus seed remains Armatocereus procerus (Fl+Fr)
Pollen A. procerus (Fl+Fr)
San Juan Seeds, pollen, fruit pulp A. procerus (Fl+Fr)
Oct 2010 Ticcoc Pollen and fruit pulp Weberbauerocereus rauhii (Fl+Fr)
Pollen and fruit pulp W. rauhii (Fl+Fr)
San Juan Fruit pulp A. procerus (Fr)
FIGURE 6. Production of owers and fruits by species of cactus in June 2010 and October
2010 at Ticcoc, San Juan, and Los Olivos sites. Codes indicate treatment (C, control; I,
impact) and phenological state (FL, owering; FR, fruiting).
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Monitoring Biodiversity on a Trans-Andean Megaproject
wing loading of species belonging to the Glossophag-
inae and Lonchophyllinae (Sahley, 1995). Sahley and
Baraybar (1996) suggest that this characteristic may
be an adaptation for ying in high- altitude habitats
(above 1,000 m asl), where this species is generally
found. The large body size found for P. genovensium
and other cactophilic bats may have several advan-
tages. Simmons and Wetterer (2002) suggest that
large body size could improve the insulation capacity
of P. genovensium, which may be important at high
altitudes where temperatures at night may reach close
to 0°C. Larger- bodied bats have a relatively lower
cost of transport than smaller- bodied bats (Sahley et
al., 1993). This could be advantageous when forag-
ing over large areas, which may occur if there are
species of cactus and food resources that are hetero-
geneous in space and time, as we have observed in
our study and others (Sahley and Baraybar, 1996;
Baraybar, 2004). For example, the nectarivorous bat
Leptonycteris yerbabuenae (previously curasoae;
Glossophaginae), a large cactophile that weighs 23
g, can y 30 km between its roosting site and forag-
ing areas (Sahley et al., 1993), and in one night of
foraging, L. yerbabuenae can y 100 km (Horner et
al., 1998). In our study, we recorded only relatively
short movements. However, our sample is small, and
we cannot make conclusions regarding nightly ight
distances for foraging activities in this study. Because
of seasonality in the owering and fruiting of colum-
nar cacti as well as an El Niño event that inuenced
Aguirre, 2008). Previous studies (Sahley and Baray-
bar, 1996; Baraybar, 2004) found that adult, nonpreg-
nant females had a slightly larger forearm and greater
weight than males, indicating possible sexual dimor-
phism in this species. The present study also found
that adult females have a slightly longer forearm than
males, but we did not nd a statistically signicant
difference with respect to weight. However, our data
are in accord with previous ndings that females
have slightly larger wings than males and that sexual
dimorphism is present in this species.
Within the nectarivorous subfamily Lonchophyl-
linae, P. genovensium is the largest species. Simmons
and Wetterer (2002) examined sizes of cactophilic
nectarivorous bats and concluded that they have lon-
ger forearm lengths, larger skulls, and larger body
sizes than other species of nectarivorous bats. Sahley
(1995, 1996) hypothesized that the morphologies of
P. genovensium and the giant hummingbird Patag-
ona gigas, with which it shares its habitat, habit of
cactophily, and which has a similar body size, have
responded to similar coevolutionary pressures with
regard to adaptation to large and tubular columnar
cactus owers. Simmons and Wetterer (2002) also
hypothesize that the cranial characters and forearm
length in P. genovensium are a coevolutionary re-
sponse to feeding from cactus owers.
Platalina genovensium has the largest wing area
in proportion to body size of both new- world necta-
rivorous subfamilies, and therefore it has the lowest
TABLE 4. Published distribution localities for Platalina genovensium. A dash (–) indicates data not available.
Number of Duration
Publication Location Altitude (m asl) individuals of study
Thomas (1928) Lima (Lima) – 1 –
Sanborn (1936) Huánuco (Huánuco) – 1 –
Ortiz de la Puente (1951) Cerro el Agustino (Lima) – 1 –
Aellen (1965) Angola, Sullana (Piura) – 1 –
Jiménez and Pefaur (1982) Yarabamba and Yura (Arequipa) 2,100–2,300 1 –
Sahley and Baraybar (1996) Yarbamba, Batolito, Charcani (Arequipa) ~2,500 87 1990–1994
Galaz et al. (1999) Azapa Valley, Tarapacá (Chile) ~300 3 –
Baraybar (2004) Yarbamba, Batolito, Charcani (Arequipa) 2,200–2,600 165 1991–1993
Zamora (2005) Lomas of Atiquipa, Caravelí (Arequipa) 600–1,000 49 1993–1994
Walsh Perú (2005) Pisco (Ica) ~1,000 6 2 months
Zamora Meza (personal observation, 2007) Lomas of Atiquipa (Arequipa) 600–1,000 50 1 month
Aragón and Aguirre (2008) Burros creek bed (Tacna) 300–800 20 12 months
Zamora Meza (pers. obs., 2007–2008) Lomas of Atiquipa (Arequipa) 2,200–2,600 18 12 months
Zamora Meza (pers. obs., 2009) Lomas of Atiquipa (Arequipa) 2,200–2,600 15 6 months
This study (2009–2010) Pisco (Ica) 1,000–2,600 33 10 months
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Zamora Meza et al. / Bats in the Area of the Pipeline n 119
utilized by P. genovensium while in their roosts, and
this information could be useful for identifying the
presence of this species in potential roost sites as well
as obtaining additional information on its distribution
and abundance.
In our study area, we found spatial and tempo-
ral variability with regard to columnar cactus species
composition and owering and fruiting schedules.
When our roost and mist net censuses are combined,
the data indicate that the bat population remained
stable year- round, with a maximum of 22 individuals
recorded in the rainy season and a minimum of 17 in-
dividuals recorded in the dry season, indicating resi-
dency in our area during the entire year. Sahley and
Baraybar (1996) found individuals of P. genovensium
year- round in the rst year of their study, but during
a prolonged drought caused by an El Niño–South-
ern Oscillation event, the abundance of this species
dropped signicantly, indicating likely migration out
of the study area. In the present study, there were spa-
tiotemporal differences in the use of foraging areas
across time and space, as demonstrated by the num-
ber of bats captured in mist nets, which did show a
marked decrease after December 2009. It is possible
that P. genovensium individuals had temporally dis-
persed out of our study area to forage, which sug-
gests that this species has larger foraging areas that
we were not able to measure.
When comparing the abundance of bats between
control and impact areas, more individuals were cap-
tured at control versus impact sites, and although this
difference was not statistically signicant, it could
indicate that the pipeline ROW could be inuencing
patterns of foraging and nightly dispersal.
Reproductive Condition
We caught potentially reproductive adult males (with
descended testicles) during all four census periods.
We found adult, reproductively active females in
March 2010 (rainy season; 1 pregnant and 1 lactat-
ing) and June 2010 (dry season; 1 lactating). The
low number of captures does not permit us to make
conclusions regarding the yearly reproductive cycle
for this species. Sahley and Baraybar (1996) found
pregnant and lactating females in the month of March
in Arequipa and found that the presence and repro-
ductive condition of reproductive females was linked
to the presence of owers and fruits of the columnar
cactus Weberbauerocereus weberbaueri. More data
cactus phenology, Sahley (1996) and Sahley and Ba-
raybar (1996) suggest that P. genovensium may be a
migratory species, following altitudinal and longitu-
dinal nectar corridors located on the western slopes
and river valleys of the Andes. In order to determine
the level of dispersal and foraging areas of this spe-
cies, however, studies utilizing radiotelemetry, as has
been done with other nectarivorous species such as
L. yerbabuenae (Sahley et al., 1993), are necessary,
along with a genetic study to determine if differen-
tiation occurs among populations of P. genovensium
(as per Wilkinson and Fleming, 1996) in its area of
distribution.
Bat Abundance in the Study Area
Similar to what has been found in other studies (e.g.,
Aragón and Aguirre, 2008; Sahley and Baraybar,
1996; Zamora, 2005) in arid habitats where this
species lives, P. genovensium was not found to be
common in our study area. The mean relative abun-
dance in our study area (0.07 individual/net hour)
was higher than that found in the Lomas del Morro
Sama in the southernmost department of Tacna (0.01
individual /net hour; Aragón and Aguirre, 2008). Sah-
ley and Baraybar (1996) found a maximum density
of 0.68 individual/km2 from censuses of roost sites,
but they did not obtain data on relative abundance.
In the lomas of Atiquipa, near the coast of the de-
partment of Arequipa, Zamora (2005) captured only
four individuals of P. genovensium in nine months
of sampling. In two roost sites near his study area,
however, he found 20 and 60 individuals, respec-
tively. This nding indicates that P. genovensium
individuals have a high capacity for dispersal when
foraging and/or are able to avoid mist nets. Thus, we
emphasize the need to locate and census roost sites
of this species in order to obtain accurate estimates
of abundance.
The studies of Zamora (2005) and Aragón and
Aguirre (2008) were conducted in the seasonal,
humid, and lowland desert habitats known as lomas,
which are not considered the most commonly used
habitat of P. genovensium, which has generally been
found at higher altitudes on western Andean slopes
(Koopman, 1978; Baraybar, 2004). However, it is
apparent that more information is needed on habi-
tat requirements, potential seasonal movements, and
roost requirements over time. A recent study by Malo
et al. (2011) has identied the echolocation signals
Alonso_5th-rev-proof.indb 119 8/5/13 12:53 PM
120
n
Monitoring Biodiversity on a Trans-Andean Megaproject
of known chiropterophilic genera, serve as a food
source for P. genovensium.
Conclusions
Platalina genovensium was recorded year- round
in the three areas sampled and in both wet and dry
seasons at altitudes ranging from 1,094 to 2,566 m
asl. However, P. genovensium occurred in low num-
bers at our study sites. Individuals feed from pollen,
nectar, and fruit from the columnar cacti A. procerus
and W. rauhii and possibly from the columnar cacti
N. arequipensis and B. candelaris. Infrequently, it
fed from insects, but these were unidentied, and we
could not verify if they were associated with cactus
owers or were caught independently.
The number of individuals of P. genovensium
varied among dry river valleys, seasons, and control
and impact sites (far from and close to the pipeline
right of way, respectively). There were more cap-
tures in the rainy season than in the dry seasons and
more captures in control than impact sites, although
none of these differences were statistically signi-
cant. These spatiotemporal variations indicate that
P. genovensium is highly sensitive to the presence
of owers and fruits of columnar cacti in its habitat.
Although we cannot conclude that the population of
P. genovensium was negatively impacted by the pipe-
line ROW, its foraging patterns and foraging areas
may have been inuenced by seasonality as well as
by changes in the availability of owers and fruits
near the ROW.
Platalina genovensium is the only bat pollinator
and seed disperser for several species and genera of
columnar cacti that are endemic to Peru. Its role in the
reproductive biology of columnar cacti in our study
area, as well as in other areas of the western Andes
in central and southern Peru where no other bat pol-
linators or seed dispersers occur, is likely signicant.
Recommendations for Monitoring and
Conservation
Ecological landscape unit 12, in Humay, Ica, con-
tains a community of mostly endemic columnar cacti,
of which some are classied as vulnerable, some as
information decient, and others, as is the case for
Cleistocactus spp. (Novoa Sheppard, this volume),
as critically endangered (although Cleistocactus has
ornithophilic adaptations). In addition, this ELU is
are needed to elucidate the reproductive patterns and
population dynamics of P. genovensium.
Diet and Association with Columnar Cacti
Data from the present study indicate that in our study
area, P. genovensium is dependent on columnar cacti
for its food. During the time in which we conducted
our study fecal analysis showed that this species con-
sumes pollen and fruit from W. rauhii and A. procerus,
which is not surprising as these species demonstrate
adaptations for chiropterophily. In Arequipa, Peru,
where detailed diet studies have been conducted,
P. genovensium demonstrates a strong association
with W. weberbaueri. A δ13C isotope study of muscle
tissue samples of P. genovensium demonstrates that it
feeds heavily on CAM plants or insects that may feed
on CAM plants, such as cacti (Sahley and Baraybar
1996). Baraybar (2004) analyzed 371 fecal samples
of P. genovensium from Arequipa, and of these, 64%
contained pollen, 29% contained both pollen and
fruit, 5% contained fruit, and only 2% contained in-
sect remains. On the basis of his study and previous
data, Baraybar concluded that P. genovensium is a
cactus specialist and not a generalist feeder.
Zamora (2005) collected 28 fecal samples in the
Atiquipa lomas; 57% contained pollen, 18% con-
tained both pollen and seeds of the columnar cactus
Echinopsis gaymardi, 11% contained pollen and in-
sects, and 14% contained only unidentied remains
of insects. The higher number of insect remains sug-
gests one of two possibilities: either P. genovensium
is actively seeking insects for food or it is also con-
suming insects found in cactus owers, as Sahley
(1995) suggested.
At our study site in Ica, the evidence indicates
that P. genovensium is a cactus specialist and a likely
pollinator and seed disperser of A. procerus and
W. rauhii. It also probably visits and feeds from the
other columnar cacti in the area, such as B. candelaris
and N. arequipensis, but because of the low ower-
ing and fruiting of these species during our study, we
were unable to conrm this possibility.
The genus Weberbauerocereus has been shown
to be pollinated by P. genovensium (Sahley, 1996),
and pollen from Armatocereus cartwrightianus has
been identied as a food source for the glossophagine
bat Anoura caudifer (Arias et al., 2009). Therefore, it
is not surprising that the columnar cacti in our study
area, W. rauhii and A. procerus, which are members
Alonso_5th-rev-proof.indb 120 8/5/13 12:53 PM
Zamora Meza et al. / Bats in the Area of the Pipeline n 121
Arias, E., R. Cadenillas, and V. Pacheco. 2009. Dieta de mur-
ciélagos nectarívoros del Parque Nacional Cerros de
Amotape [Diet of nectarivorous bats from Cerros Amo-
tape National Park]. Revista Peruana de Biología, 16(2):
187–190.
Baraybar, L. 2004. Parámetros biométricos y ecológicos del
“Murciélago Longirostro Peruano” Platalina genoven-
sium Thomas, 1928 (Phyllostomidae), en la Provincia de
Arequipa (1994) [Biometric and ecological parameters of
the “Peruvian long- snouted bat” Platalina genovensium
Thomas, 1928 (Phyllostomidae), in Arequipa Province].
Bachelor’s thesis. Universidad Nacional de San Agustín,
Arequipa, Peru.
Bertin, R. I. 1982. Floral Biology, Hummingbird Pollina-
tion and Fruit Production of Trumpet Creeper (Campsis
radicans, Bignoniaceae). American Journal of Botany,
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Environmental Resources Management Peru. 2008. Final Re-
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Faegri, K., and L. Van der Pijl. 1966. The Principles of Pol-
lination Ecology. Oxford, U.K.: Pergamon Press.
Fleming, T. H., C. Geiselman, and W. J. Kress. 2009. The
Evolution of Bat Pollination: A Phylogenetic Perspec-
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Fleming, T. H., and N. Muchhala. 2008. Nectar- Feeding Bird
and Bat Niches in Two Worlds: Pantropical Comparisons
of Vertebrate Pollination Systems. Journal of Biogeogra-
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Freeman, P. W. 1995. Nectarivorous Feeding Mecha-
nisms in Bats. Biological Journal of the Linnean So-
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- 8312.1995.tb01104.x.
Galaz, J. L., J. Torres, and J. Yáñez. 1999. Platalina genoven-
sium (Thomas, 1928), un quiróptero nuevo para la fauna
de Chile (Phyllostomatidae: Glossophaginae) [Platalina
genovensium (Thomas, 1928), a new quiropteran for the
Chilean fauna (Phyllostomatidae: Glossophaginae)]. No-
ticiario Mensual del Museo Nacional de Historia Natu-
ral (Chile), 337:6–12.
Gardner, A. L. 2008. Mammals of South America, Volume 1.
Marsupials, Xenarthrans, Shrews, and Bats. Chicago:
University of Chicago Press.
Heithaus, E. R., P. A. Opler, and H. G. Baker. 1974. Bat Activ-
ity and Pollination of Bauhinia pauletia: Plant- Pollinator
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aging Behaviour and Energetic of a Nectar- Feeding Bat,
Leptonycteris curasoae (Chiroptera: Phyllostomidae).
also home to P. genovensium, which is found in low
numbers. Platalina genovensium faces several threats,
such as being collected for medicinal use and the de-
struction of its habitat (Sahley and Baraybar, 1996).
It is possible that it is an endemic species from Peru
since no resident population has been reported for
Chile (Galaz et al., 1999). At present, it is listed as
critically endangered within Peru and as near threat-
ened by the IUCN red list. We recommend that the
IUCN red list classication be modied to reect its
distribution, population data, and legal status in Peru.
As for the construction of the pipeline right of way,
there was no statistically signicant effect on popula-
tions, but given the rarity and ecological importance of
this bat, we recommend the following to obtain more
conclusive data regarding the potential longer- term
impacts of cactus removal on the ROW and effects on
P. genovensium: (1) additional monitoring of bats that
is conducted throughout the entire night and includes
a census of all roost sites; (2) longer- term monitoring
of cactus phenology as well their demography and
reproductive biology since the status of most species
is unknown; (3) monitoring bats using radiotelemetry
and camera traps at owers to better determine activ-
ity, movement patterns, foraging areas, and possible
migratory corridors; (4) measuring the importance of
P. genovensium as a pollinator or seed disperser of co-
lumnar cacti; (5) continued monitoring of transplanted
cacti to determine if there are impacts on survival, phe-
nology, and reproduction; and (6) in future projects,
following the strategy utilized during the construction
phase of the pipeline to avoid disturbing columnar
cacti as much as possible.
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