Tent Roosts of Macconnell's Bat (Vampyressa macconnelli)

Author(s): Mercedes S. Foster

Source: Biotropica, Vol. 24, No. 3 (Sep., 1992), pp. 447-454
Published by: Association for Tropical Biology and Conservation
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 BIOTROPICA 24(3): 447-454 1992

Tent Roosts of Macconnell's Bat (Vampyressa macconnell)l1

Mercedes S. Foster

National Ecology Research Center, Fish and Wildlife Service, National Museum of Natural History,

Washington, D.C. 20560, U.S.A.

ABSTRACT

Between 11 September and 2 November 1990, 26 tent roosts, four of which were occupied by Macconnell's bat

(Vampyressa macconnelli), were discovered in an area of mature flood plain forest in Manu National Park, Madre de

Dios, Peru. All were constructed in the terminal blade of unseparated pinnae in fronds of Astrocaryum macrocalyx

Burret (Palmae). Tents generally were not found in fronds of A. macrocalyx shorter than 2 m or longer than 4 m,

or in other species of palms or broad-leaved plants that have been reported to harbor tent roosts of bats. The proximal

four to nine unseparated pinnae of the terminal blade were severed to form the tent. Tents averaged 50.7 cm long,

17.1 cm maximum width, and 16.7 cm maximum depth, and were an average of 2.33 m above the ground. From

one to seven bats were present in the four occupied tents but were not present in a tent every day. The density of

active tents was ca 0.74/ha.

RESUMEN

Entre el 11 de septiembre y el 2 de noviembre 1990, se encontraron en el area de bosque maduro inundable del

Parque Nacional del Manu, Madre de Dios, Peru, 26 toldos diurnos de descanso del murcielago, cuatro de que

estaban ocupados por el murcielago de Macconnell (Vampyressa macconnelli). Casi todas se construyeron en las frondas

de Astrocaryum macrocalyx Burret (Palmae), en la parte terminal de la hoja donde las hojitas no se separan. Generalmen-

te, no se encontraron los toldos en frondas de A. macrocalyx con un largo inferior de 2.0 m o mayor de 4.0 m, ni

en palmeras de otras especies, ni en plantas con hojas anchas. Las proximas cuatro a nueve hojitas no separadas se

cortaron para formar el toldo. El promedio del largo del toldo era 50.7 cm, el ancho mrnximo, 17.1 cm, y la profundidad

mrnxima, 16.7 cm. Su altura promedio sobre el suelo era 2.33 m. Se encontraron desde uno hasta siete murcielagos

en los toldos ocupados. Los murcielagos no se encontraron todos los dias en los toldos. Toldos en uso se encontraron

en una densidad de 0.74/ha.

Key words: Astrocaryum macrocalyx; bat tent; Macconnell's bat; Peru; Vampyressa macconnelli.

YELLOW-EARED BATS of the genus Vampyressa are aceae. Timm (1984) found a single male V. pusilla

found from southern Mexico to southern Peru, in a tent made from a Philodendron leaf. Koepcke

(1984) found two tent roosts of Macconnell's bat

southern Brazil, and Paraguay, where they occupy

gardens, plantations, and mature and disturbed rain (V. macconnelli) in fronds of Geonoma sp. and an-

other in a leaf of Anthurium sp., although the species

forest (Emmons 1990). Despite their wide distri-

bution, these bats are considered uncommon, and has also been reported to roost elsewhere (Sanborn

little is known of their natural history (Nowak & 1951, Goodwin & Greenhall 1962, Handley 1976).

Between 11 September and 2 November 1990,

Paradiso 1983, Lewis & Wilson 1987). Roosting

I discovered 26 tents of V. macconnelli. The tents

habits of three species, Vampyressa brocki, V. bidens,

showed little variation in structure but differed in

and V. melissa, are unknown (Emmons 1990), and

roost records are extremely limited for two of the several respects from those reported by Koepcke

(1984). Herein I describe the tents, characterize

other three species. Nevertheless, it appears that

plants selected for their construction, and provide

Vampyressa spp., like other Stenoderminae, roost in

notes on use of tents by these bats. I also report on

tents.

Brooke (1987) located 115 tents (11 occupied) experiments to assess the suitability of fronds of

different palm species for tents.

of V. nymphaea in the broad leaves of Pentagonia

Vampyressa macconnelli (Thomas 190 1) is more

donnell-smithii, an understory shrub in the Rubi-

commonly referred to in the literature as either Me-

sophylla macconnelli or Ectophylla macconnelli. My

use of the name combination V. macconnelli for this

' Received 12 August 1991, revision accepted 20 De-

bat follows Owen (1987).

cember 1991.

447

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 448 Foster

TABLE 1. Properties of roost tents of Vampyressa macconnelli in fronds of Astrocaryum macrocalyx.

Cut pinnae

Height above Complete Partial Rachis dist. Tent dimensions (cm)

ground (m) no.a no.b (cm)' Len. base Width max. Depth max.

N 24 50 50 24 19 15 14

Min 1.65 4 0 4.7 40.0 12.0 10.2

Max 3.60 9 4 25.2 69.8 29.0 21.7

Mean 2.33 6.6 2.1 11.7 50.7 17.1 16.7

SD 2.44 1.3 1.2 5.1 8.0 4.9 3.2

a Pinnae cut through completely; combined value for right plus left side of rachis.

b Pinnae cut through the blade but not the central vein; combined value for right plus left side of rachis.

c Distance along the rachis occupied by the bases of the pinnae cut through completely.

For each tent I identified the plant species, noted

METHODS

frond characteristics, and measured the fronds and

tents. I measured angles with a protractor. I was

Observations were made in mature flood plain forest

in the vicinity of Cocha Cashu Biological Station, unable to take all measurements from damaged

Manu National Park, Madre de Dios, Peru (1 155'S, tents.

77?18'W; elev. ca 380 m). The area is described To assess the suitability of fronds of different

in Terborgh (1983). The first tent was located when species for tents and to determine the rate at which

I brushed against it and flushed the bats. Its struc- cut pinnae dry and break, I constructed tents in

three species of palms by cutting the fronds in a

ture was distinctive enough to enable recognition of

other tents, even in the absence of bats. Some ad- pattern mimicking the bat cuts. In addition, I count-

ed the number of unseparated terminal pinnae on

ditional tents were located at random. Most, how-

ever, were located during systematic examination of both sides of the rachis, noted the rachial distance

covered by these pinnae, and measured the length

all trees and shrubs in 51 40 x 40 m plots in

connection with vegetation studies. of the leading edge of the most anterior pinna in

Tents that did not contain bats were subjectively Astrocaryum fronds of different lengths to determine

classed as probably active, probably inactive, or their suitability for tents.

disintegrating, based on the integrity of the structure Occasionally, when scanning Astrocaryum fronds

for tents, for an instant I mistook suspended dead

and the dryness of the cut areas. Tents that looked

active were checked for 1-10 days to see whether leaves for bats. To determine if such debris was

bats occupied them. The ground under each tent present frequently enough to provide negative re-

was searched carefully for seeds or bat droppings. inforcement to sight-hunting predators, I counted

I netted a single 8.5-g nonlactating female bat unmodified fronds with dried leaves suspended be-

from the first tent located. It measured: total length, neath the terminal blade.

46 mm; tail, 0; forearm, 31.8; foot, 6.2; ear, 13.0; Mean values were compared using the Wilcox-

and tragus, 6.0. It had pale brown fur and dark on-Mann-Whitney test for large samples corrected

wing membranes. The proximal half of each ear for ties (Siegel & Castellan 1988). Tests were one-

and tragus, the base of the nose leaf, and the lower tailed and tested whether character values for 2-

lip were bright yellow. I examined the teeth with 4-m fronds exceeded values for fronds of other

a 14 x hand lens. The second lower molar was lengths.

considerably wider than the first and had a relatively

shallow cleft between the lingual cusps. The third

RESULTS

lower molar was a tiny spike. The second upper

molar was also about twice the size of the first and TENT STRUCTURE.-All of the 2 6 tents I located

square rather than triangular. I subsequently ex- were in fronds of Astrocaryum macrocalyx Burret

amined specimens of Artibeus spp., V. pusilla, and (Palmae), which has pinnately compound leaves. In

most fronds, terminal pinnae remain unseparated,

V. macconnelli in the collections of the National

Museum of Natural History in Washington, D.C. forming a broad blade along the distal 10 percent

On the basis of the described characteristics, I iden- of the rachis. Bats constructed their tents in this

tified the tent bat as Vampyressa macconnelli. blade by severing the central vein and lateral shafts

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 Tent Roosts of Bats 449

FIGURE 1. Terminal blade of Mstroaty&m macrocalyx showing pinnae CUt tO form a tent roost used by Vampyressa

macconnell:.

of the first (proximal) four to nine unseparated pinnae were cut decreased distally (e.g., most prox-

pinnae, which occupy about 40 percent of the ra- imal pinnae, #1: x = 8.8 cm, range = 4.5-18.0

chial length of the blade (Table 1). In nearly all cm; N = 43; pinnae #5: x = 6.8 cm, r = 3.3-

tents, the shafts of an additional one to four pinnae 12.1, N= 44; pinnae #9: x = 3.8 cm, r= 1.8-

were prtially cut, but not the central vein. The 6.5, N = 16; Fig. 1). The angle at which the pinnae

distances from the frond rachis at which successive diverged from the rachis decreased distally as well

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450 Foster

It A~~~

FIGURE 2. Tent roosts of Vampyressa macconnelli and palm fronds at Cocha Cashu Biological Station, Manu

National Park, Madre de Dios, Peru. (a) V. macconnelli hanging in a tent roost in an Astrocaryum macrocalyx frond.

(b) Lateral view of a tent in a frond of A. macrocalyx. (c) Dead leaf resembling a small bat hanging from the underside

of the terminal, unseparated pinnae of a frond of A. macrocalyx. (d) Terminal, unmodified blades of Scheelea cephalotes

(left), Hyospathe elegans (center), and A. macrocalyx (right) fronds.

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 Tent Roosts of Bats 451

(e.g., angle of pinna #1: x = 420, r = 26-49', N 13.4, range = 9-23, N = 64) than fronds of 1.0-

- 20; #5: x = 19?, r = 13?-23?, N = 20; #9: x 2.0 m (x = 10.3, range = 9-13, N = 40; Z =

- 110, r = 7?-160, N = 11; Fig. 1). The path -6.719, P < 0.001) or >4.0 m (x = 5.8, range

described by the cuts was straight or somewhat "J" = 3-9, N = 40; Z = -8.582, P < 0.001). The

shaped. The cuts generally produced a hole 0.2- extent of the unseparated pinnae along the central

4.0 cm long and 0.2-1.0 cm wide. The severed rachis was also significantly greater on 2.0-4.0 m

edges were irregular and obviously chewed over 1 fronds (x = 29.7 cm, range = 13.5-58.0, N =

to 2 mm. 32) than on those >4.0 m (x = 8.2 cm, range =

The parts of the blade formed by the distal 3.0-17.8, N = 20; Z = -5.954, P < 0.001) or

portions of the severed pinnae folded ventrally to from 1.0-2.0 m (x = 15.9 cm, range = 9.7-22.8,

form the sides of a large chamber. The cut ends of N = 20; Z = 4.901, P < 0.001). The length of

the walls projected backward, curving to meet in the leading edge of the blade was approximately

the midline and close the chamber posteriorly. The the same in all groups (2.0-4.0 m: x = 32.6 cm,

anterior portions of the blade curved forward over- N= 64 vs 1.0-2.0 m: x = 29.2 cm, N = 20; Z

lapping in the midline. In some tents, overlap was = -1.589, P > 0.05; 2.0-4.0 m vs >4.0 m: x

complete, and the chamber was closed anteriorly = 30.3 cm, N = 20; Z = -1.063, P > 0.05).

and dorsally. In other tents, a narrow slit remained Even within the 2.0-4.0 m range, the bats

along the anterodorsal midline (Fig. 2a). seemed to have selected among fronds. The number

In shape, tents most closely resembled inverted of unseparated terminal pinnae (x = 14.5, range

canoes (Fig. 2b). Because fronds curve downward = 12-19, N = 48; Z = 3.762, P < 0.001) in

distally, the portion of the rachis to which the uncut, fronds with tents, their rachial distance (x = 30.5,

unseparated pinnae are attached, i.e., the top of a range = 20.3-41.7, N = 24; Z = 2.600, P <

tent, was usually horizontal. The leading and trailing 0.005), and the length of the leading edge of the

edges of tents sloped downward. Tents bulged in blade (x = 37.3, range = 25.6-50.0, N = 24; Z

the center because of changes in pinnae widths along = 2.153, P < 0.025) all exceeded these values for

their lengths. An unseparated pinna is tightly pleat- the 2.0-4.0 m fronds sampled at random (see

ed where it attaches to the rachis. A few centimeters above).

distally, it begins to flatten and is ca 17-22 mm Nine of the 10 active or probably active tents

wide. At 50-67 percent of its length, it reaches a were in the next to newest fronds in the tree; usually

maximum width of 20-26 mm, decreasing again the newest frond was still growing. The other tent

to ca 18 cm a few centimeters from the tip. Tent was in a tree with two newer fronds. Unused tents

dimensions are summarized in Table 1. were in trees with 3-7 newer fronds.

Edges of the cut pinnae in the tents I constructed I examined 205 2.0-4.0 m long fronds in 51

turned brown after ca 1 wk. The brown area was Astrocaryum palms for suspended leaves. Forty-two

2 to 3 mm wide after ca 1 mo. The ventral edges fronds (20.5%) had one or more dried leaves hang-

of tents (i.e., trailing edge of the most proximal cut ing near the rachis of the terminal unseparated pin-

pinna) sometimes began to curl after 3 to 4 wk. nae (Fig. 2c). These leaves were attached with spi-

The most common damage to tents was the loss of der's web or insect material, or their petioles were

one or both sides. wedged in holes or separations between pinnae.

Another common species of palm at Cocha Ca-

PLANT CHARACTERISTICS.-The Astrocaryum fronds shu that might be suitable for tents is Scheelea

used by V. macconnelli for tents ranged from 2.2 cephalotes (Poepp. ex Mart.) Karsten, in which the

to 4.7 m long (x = 3.23 m, SD = 0.58, N= 26), terminal pinnae of the shorter (usually <2.0 m)

but only one frond exceeded 4 m. All were in fronds remain unseparated to form a terminal blade

relatively small palms that lacked a trunk. Fronds (Fig. 2d). The unseparated pinnae are weaker and

of mature trees usually exceed 4 m and often reach narrower than those of Astrocaryum. Because they

8 m. All fronds used for tents lacked spines along form an extremely acute angle with the central ra-

the ventral rachis in the area of the tent, although chis, the leading edges of the anterior pinnae are

palms of this species are generally extremely spiny. separated by only a few centimeters, and the blade

There was no evidence that spines had been re- is narrow. In addition, the pinnae, though attached

moved. over most of their lengths, are usually separated

Several frond characteristics varied significantly along the basal few centimeters (Fig. 2d). Thus,

among fronds of different length. Fronds of 2.0- when I cut Sheelea fronds (N = 7) in the fashion

4.0 m had more unseparated terminal pinnae (x = of the tents in Astrocaryum, the pinnae did not hold

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 452 Foster

together to make a roof posteriorly nor did they tent roof adjacent to the midrib or to a pinneal vein

overlap anteriorly to close the tent in front. The within 2 cm of the midrib, beginning 1.5-13.0 cm

resulting tube-shaped structures were 5.0-7.0 cm from the proximal end of the tent and covering

wide and 5.0-8.0 cm deep, considerably smaller distances of 8.8-20.6 cm.

than the Astrocaryum tents (Table 1). Twenty tents were located in the vegetation

The unseparated terminal pinnae in fronds of plots, which had a combined area of 81,600 M2.

Hyospathe elegans Mart., another common under- Six of these were probably active, for a density of

story palm, resemble the terminal blade of Astro- ca 0.74 active tents/ha. With 30 percent of active

caryum (Fig. 2d). However, the pinnae and their tents occupied by an average of 3.3 bats, a crude

central veins seem weaker in Hyospathe and diverge estimate of bat population density is ca 1.1 bat/

from the rachis at a greater angle. When I severed ha. No bat droppings were apparent under any of

pinnae in the manner of the bat roosts in Astro- the roosts.

caryum, the leaf blade (N = 5) sometimes folded

anteroventrally, closing in front but not laterally, so

that no tent was formed. In other fronds (N = 3),

DISCUSSION

the cut blade folded ventrolaterally, but did not

close at either end or above, so that hanging bats Koepcke (1984) found two tents of V. macconnelli

would be exposed from almost any angle. about 1 m above the ground in fronds of Geonoma

Several of more than a dozen other palm species sp. The manner of construction was the same as

found in the flood plain forest at Cocha Cashu (R. that of tents I found in Astrocaryum. In both in-

B. Foster, pers. comm.) might be suitable for tents stances, the cuts in the fronds followed either the

(e.g., Phytelephas macrocarpa R. & P., Geonoma "V" or "J" shapes described for tents of Artibeus

spp.) as might the many species of Araceae, Cyclan- spp. and Uroderma (Barbour 1932, Chapman 1932,

thaceae, Marantaceae, and Musaceae from which Foster & Timm 1976, Timm 1987).

tents of various bat species have been reported pre- The Geonoma tent pictured by Koepcke (1984)

viously (Timm & Mortimer 1976; Koepcke 1984; was essentially pendant, with the bats in the apex,

Timm 1984, 1987). Many broad-leafed species of while the Astrocaryum tents were more horizontal,

Zingiberaceae and Rubiaceae are also present. I with the bats located proximally. The tents in As-

examined individuals of these families whenever trocaryum were also considerably higher above the

they were encountered but did not locate any bat ground (Table 1). These differences undoubtedly

tents. reflect differences in the strengths of the rachis and

veins of the leaves of these species and the sizes of

OCCUPATION OF TENTS BY BATS-.Three of the 26 the trees.

tents were active (contained bats when located), Koepcke (1984) found tents of V. macconnelli

seven were probably active, one was probably in- in both Geonoma sp. and Anthurium sp. Goodwin

active, and 15 in various stages of disintegration and Greenhall (1962) also reported finding these

were assumed to be inactive. Tents contained 1-7 bats under a leaf of Antharium (sic) huegelli (=

bats (x = 3.3, N = 3). The seven individuals Anthurium hookeri Kunth), but did not mention

returned after being flushed twice within an hour, that the leaf was modified. I did not find tents in

and then disappeared and did not return during the species of either genus. At Cocha Cashu these bats

next seven days. Another tent had two roosting bats showed a clear preference for young Astrocaryum

that were never flushed. They occupied the tent one fronds 2.0-4.0 m long. Bats may be selecting young

day, were absent the next, and (presumably the leaves that are easier than older ones to chew, as

same ones) were present again the day after, but suggested by Choe and Timm (1985) for Artibeus

not during the next eight days. A single bat did watsoni and by Timm and Lewis (1991) for Uroder-

not return in four days after being disturbed. Finally, ma bilobatum. Also, terminal blades of shorter and

one bat occupied a tent 27 days after I constructed longer Astrocaryum fronds may be too small and

it in an Astrocaryum frond. It also did not return have too few pinnae to form tents with adequate

in 6 days after being flushed. shelter. Leaves of other plant species, such as Schee-

Roosting bats occupied the proximal half of a lea cephalotes, also may not be as strong as those

tent (Fig. 2a), hanging face to face and slightly of Astrocaryum macrocalyx, or may not form as

offset on either side of the central rachis. Opposite effective a shelter. Others (Timm & Mortimer 1976;

and adjacent individuals seemed to touch. Tiny holes, Choe & Timm 1985; Brooke 1987, 1990; Timm

presumably made by bat claws, were present in the & Lewis 1991) have reached similar conclusions

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 Tent Roosts of Bats 453

about the suitability of leaves of certain size or surely flush roosting bats by moving the fronds when

species for roost tents of other bat species.

climbing. Brooke (1990) suggested that close prox-

Vampyressa macconnelli is considered uncom- imity and continued use of tents, which has been

mon in high ground forest around the Cocha Cashu reported for several species (e.g., Koepcke 1984,

station (Terborgh et al. 1984), which is confirmed Timm 1987), implies a low level of predation on

by my estimate of bat density. Nevertheless, the bats roosting in such structures. On the other hand,

availability of suitable fronds for roost tents prob- Central American Squirrel Monkeys (Saimiri oer-

ably does not influence the abundance of this bat. stedi) may represent a significant mortality factor

Astrocaryum of the size used are common in the for tent-occupying bats in western Costa Rica. These

forest understory. The tents that I constructed seemed monkeys recognize and actively search tents. They

perfectly functional after a month, although the cuts attempt to catch bats by jumping on the tents and

I made with scissors probably did not dry as quickly knocking them and their contents to the ground

as those chewed by bats. Koepcke (1984) suggested (Boinski & Timm 1985). Bats flushed from the

that tents might be used for up to six months, and roost are also exposed to predation by double-toothed

Brooke (1987) reported a tent of V. nymphaea that kites (Harpagus bidentatus) or other raptors that

was used for nine months. The presence of two tents forage with monkeys (Boinski & Timm 1985).

each in four palms, and the successive, rather than Common Squirrel Monkeys (Saimiri sciureus) and

simultaneous construction of the tents in the palm capuchins (Cebus albifrons and C. apella), as well

in which it was possible to determine the sequence as double-toothed kites, are found at Cocha Cashu

suggest that the bats have other, as yet unidentified, (Terborgh et al. 1984).

microhabitat preferences. Close proximity of active tents and their con-

The use by a V. macconnelli of a tent that I tinued use could increase the vulnerability of the

made shows that these bats will occupy tents that bats to predators that initially locate only one tent

they do not construct. Brooke (1987) also reported or that revisit sites of previous prey contact. This

this for V. nymphaea, and both she and Timm may explain the intermittent use of some tents,

(1987) observed that some bats will occupy tents particularly after disturbance. Additional negative

of other species. Because, like others who have re- reinforcement for diurnal, sight-hunting predators

ported tent roosts, I did not observe any tent being may derive from contact with dead leaves suspended

made, it is possible that the roosts occupied by the from the underside of the terminal blade in Astro-

V. macconnelli at Cocha Cashu were constructed by caryum. These often resemble small hanging bats

another species of bat. Eighteen species of stenoder- and at casual glance can deceive an experienced

mine bats have been recorded at the site in addition observer (Fig. 2c). Many diurnal forest raptors occur

to V. macconnelli (Terborgh et al. 1984, Ascorra in the study area (Terborgh et al. 1984), but none

et al. 1991). At least nine of these roost in tents: has been reported to forage at bat tents.

Uroderma bilobatum and U. magnirostrum (Timm

1987); Vampyressa pusilla (Timm 1984) and V.

ACKNOWLEDGMENTS

nymphaea (Brooke 1987); Artibeus andersoni, A.

I am grateful to the Direcci6n General de Flora y Fauna

gaucus, A. gnomus (Timm 1987), A. cinereus

of the Peruvian Ministry of Agriculture and to the ad-

(Goodwin & Greenhall 1961), and A. jamaicensis

ministration of Manu National Park for permission to

(Foster & Timm 1976). However, the only bats in

carry out research in the park. M. Silman drew my at-

the tents I found were V. macconnelli. In addition,

tention to two tents and lent equipment. R. B. Foster

tent construction differed significantly from that re- assisted with the identification of the palms. J. Terborgh

provided logistical support. R. Altig, A. L. Gardner, R.

ported for the other nine species yet was the same

Hole, Jr., J. W. Kress, R. W. McDiarmid, S. F. Smith,

as that described by Koepcke (1984) for V. mac-

R. M. Timm, and D. E. Wilson provided literature ref-

connelli, all of which strongly suggest that the tents

erences and other information. L. Marquez helped with

were made by this species.

the preparation of the Spanish summary. V. Krantz as-

sisted in the preparation of the figures. A. L. Gardner

Tents clearly function to shield roosting bats

and D. E. Wilson also commented on the manuscript.

from sun, wind, and rain (Timm & Lewis 1991)

The field work was supported in part by the Alexander

and to conceal them from predators overhead and

Wetmore Fund of the Smithsonian Institution. I thank

to the side. Ground predators may be deterred by

all these individuals and institutions for their generous

the rachial spines on the Astrocaryum fronds and assistance.

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454 Foster

LITERATURE CITED

ASCORRA, C. F., D. E. WILSON, AND M. RoMo. 1991. Lista anotada de los quir6pteros del parque nacional Manu,

Peru. Publ. Mus. Hist. Nat. UNMSM (A) 42: 1-14.

BARBOUR, T. 1932. A peculiar roosting habit of bats. Q. Rev. Biol. 7: 307-312.

BOINSKI, S., AND R. M. TIMM. 1985. Predation by squirrel monkeys and double-toothed kites on tent-making bats.

Am. J. Primatol. 9: 121-127.

BROOKE, A. P. 1987. Tent construction and social organization of Vampyressa nymphaea (Chiroptera: Phyllostomidae)

in Costa Rica. J. Trop. Ecol. 3: 171-175.

1990. Tent selection, roosting ecology and social organization of the tent-making bat, Ectophylla alba,

in Costa Rica. J. Zool. (Lond.) 221: 11-19.

CHAPMAN, F. M. 1932. A home-making bat. Nat. Hist. 32: 555-556.

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