roost boxes as a tool in the conservation of microbats ... · native habitat since european...

20
Roostboxes as a tool in the conservation of tree roosting microbats (Microchiroptera) in a highlymodified agricultural landscape Quarry Life Award Research Project, Australia Project Topic: Biodiversity and Rehabilitation Stephen Griffiths July 2012 Department of Zoology The University of Melbourne Victoria 3010 Australia Telephone: +61 3 8344 6244 Fax: +61 3 8344 7909

Upload: vanngoc

Post on 24-Aug-2019

216 views

Category:

Documents


0 download

TRANSCRIPT

Page 1: Roost boxes as a tool in the conservation of microbats ... · native habitat since European settlement has resulted in significant negative pressure on microbats (Microchiroptera),

Roost‐boxes as a tool in the conservation of 

tree roosting microbats (Microchiroptera) in 

a highly‐modified agricultural landscape 

Quarry Life Award Research Project, Australia 

Project Topic: Biodiversity and Rehabilitation 

 

Stephen Griffiths   

July 2012 

 

 

 

 

 

 

 

 

 

 

     

Department of Zoology 

The University of Melbourne 

 Victoria 3010 Australia 

Telephone: +61 3 8344 6244   

Fax: +61 3 8344 7909 

Page 2: Roost boxes as a tool in the conservation of microbats ... · native habitat since European settlement has resulted in significant negative pressure on microbats (Microchiroptera),

Roost-boxes as a tool in the conservation of tree roosting microbats

1

Abstract

The loss, modification and fragmentation of natural habitats constitute a major threat to bat

populations worldwide. This is particularly relevant in Australia where extensive loss of

native habitat since European settlement has resulted in significant negative pressure on

microbats (Microchiroptera), the majority of which rely on tree hollows for roosting.

However, because microbats are small, nocturnal and remain hidden in roosts during the

day, few people are aware of the diversity or abundance of bats that roost in trees in both

urban and rural areas. I used bat detectors to examine activity of microbats at two sites in a

highly-modified agricultural matrix. The species assemblage recorded comprised the

majority of microbat species known to occur in the Western Volcanic Plains region of

Victoria. A total of 6,214 echolocation call sequences were recorded over 39 consecutive

nights at the two sites. Considerable variation in activity was documented both within and

between nights. The echolocation data revealed minimal levels of foraging activity. At one

site, social calls were recorded in the hour prior to civil twilight (i.e. prior to nightfall) on

17 nights, providing indirect evidence of bats roosting diurnally in tree hollows within an

isolated stand of mature Eucalyptus cladocalyx. Artificial roosting boxes (bat-boxes) were

installed to provide supplementary roosting habitat as an initial step toward investigating

the efficacy of bat-boxes as a targeted conservation tool within the framework of a long-

term ecological management and restoration plan. The presence and activity of microbats

recorded in this study, and specifically the detection of social calls produced by diurnally

roosting bats, has implications for the management and rehabilitation of isolated patches of

land within highly-modified agricultural landscapes. Increasing the awareness of land

managers and decision makers to the benefits of these species (e.g. regulation of

herbivorous insect abundance), will enhance knowledge of this faunal group in the

community and provide further reasons for prioritising the management and conservation

of scattered trees within urban and rural environments.

Page 3: Roost boxes as a tool in the conservation of microbats ... · native habitat since European settlement has resulted in significant negative pressure on microbats (Microchiroptera),

Roost-boxes as a tool in the conservation of tree roosting microbats

2

Introduction

A wide variety of vertebrate taxa use tree hollows (also referred to as tree holes or cavities)

for shelter and as breeding sites (Goldingay 2009). Worldwide there are approximately 260

species of bird and 360 species of non-flying mammals that use tree cavities (Newton 1994;

Novak 1999), while in Australia, as many as 300 native vertebrate species (birds, bats,

arboreal marsupials, reptiles and frogs) use tree hollows (Gibbons and Lindenmayer 2002;

Kunz and Lumsden 2003; Goldingay and Stevens 2009).

Bats are among the most threatened fauna in Australia, with 36 species listed under The

Action Plan For Australian Bats (Duncan et al. 1999). Day-roost sites have been identified

as a key limiting resource for microbat conservation, but we know almost nothing about the

causes and consequences of roosting in different types of tree hollows or artificial nesting

boxes (Turbill 2006; Goldingay 2011). The dramatic decline of Australian native habitats

since European settlement has led to a growing awareness of the impact of habitat loss on

native fauna, and revegetation efforts, both in farmland (Kavanagh et al. 2007) and in

suburbia (Harper et al. 2005a,b), are increasingly common. However, trees planted now

may take up to 100 years to develop hollows suitable for use by wildlife (Lindenmayer et

al. 1993; Weinberg et al. 2011). Therefore, while revegetated areas can provide foraging

habitat for bats soon after planting, the lack of hollows may limit colonisation of these areas

as roosting habitat.

One short to medium term method of compensating for limited roosting sites is to install

nesting boxes (bat-boxes) as substitutes for natural hollows (Arnett and Hayes 2000;

Brittingham and Williams 2000; Flaquer et al. 2005; Lesinski et al. 2009). There have been

few studies of bat-box use in Australia, particularly in agricultural regions, and

consequently relatively little is known of the use and preferred designs of bat-boxes by

Australian tree roosting microbats (Irvine and Bender 1995; Smith and Agnew 2002;

Goldingay and Stevens 2009). My objective in the present study was to examine microbat

activity in a highly-modified agricultural landscape and conducted a preliminary

assessment of the efficacy of bat-boxes as a targeted conservation tool within the

framework of a long-term ecological management and restoration plan. Specifically, I

investigated: (1) temporal patterns (hourly and nightly) in bat activity; (2) the diversity of

Page 4: Roost boxes as a tool in the conservation of microbats ... · native habitat since European settlement has resulted in significant negative pressure on microbats (Microchiroptera),

Roost-boxes as a tool in the conservation of tree roosting microbats

3

species present; (3) the level of foraging activity; and (4) the potential value of deploying

bat-boxes as a short to medium term conservation tool.

Methods

Study sites

I conducted fieldwork at two former sheep grazing properties located in the Western

Volcanic Plains of Victoria, Australia. The region is characterised by temperate grassland

and grassy eucalypt woodland, now dominated by introduced grassland species, with

sparsely scattered paddock trees and occasional trees planted along fence lines as

windbreaks (DEWHA 2008). In 2008, the Australian government upgraded the listing of

the natural temperate grassland and the grassy eucalypt woodland of the Volcanic Plains of

Victoria to critically endangered, the most protected conservation status under the Federal

Environment Protection and Biodiversity Conservation Act 1999 (DSEWPC 2011). The

climate of the Western Volcanic Plains is temperate with warm to hot summers (average

summer maximum 24.6°C) and cool to cold winters (average winter maximum 12.6°C).

Rainfall on the plains ranges from 600 to 800 mm per annum and occurs mostly between

April and November. Annual average rainfall at the Australian Bureau of Meteorology

Hamilton Airport monitoring station (37°38'39.04'' S, 142°00'47.74'' E) is 685.7 mm

(Australian Bureau of Meteorology 2012).

The Warrayure Conservation Offset Site (37°44'05.65'' S, 142°12'47.93'' E) is located 230

km west of Melbourne, 14 km south-west of Dunkeld and 17 km east of Hamilton. The

Warrayure site has an area of 58 hectares, comprised predominantly of degraded grassland

with small patches of mature Eucalyptus cladocalyx. The site is on the north shore of Lake

Linlithgow, and it adjoins the Lake Linlithgow Nature Reserve. The Strathkellar

Conservation Offset Site (37°43'24.48'' S, 142°07'50.02'' E) is located 10 km west of the

Warrayure site, 20 km south-west of Dunkeld and 12 km east of Hamilton. The Strathkellar

site has an area of 80 hectares, comprised of highly degraded grassland and creek line

tussock grassland. Native tree species are limited to a small patch of Acacia melanoxylon

along the southeast boundary. The property is bordered by the Glenelg Highway to the

south and Rhooks Road to the northeast. Both the Warrayure and Strathkellar Conservation

Page 5: Roost boxes as a tool in the conservation of microbats ... · native habitat since European settlement has resulted in significant negative pressure on microbats (Microchiroptera),

Offse

the im

Acou

I use

Cons

bat d

Elect

levels

Detec

heigh

(Figu

time

FigurWarr

I dow

manu

www

echol

2008

file (

softw

ident

et Sites are

mprovement

ustic sampli

ed echoloca

servation Of

detectors (A

tronics, Law

s against an

ctors were p

ht of approx

ure 1). Reco

the detector

e 1. AnaBatayure Conser

wnloaded e

ually analy

w.hoarybat.c

location pul

). Each call

(Hayes 2000

ware (Gibso

ify call seq

Roo

covered by

t of native g

ing of bat c

ation calls a

ffset Sites. I

naBat II ba

wnton, Quee

n ultrasound

placed insid

ximately 1.

ordings bega

rs were trigg

t II bat detervation Offse

echolocation

ysed calls

com). A sing

lses (frequen

l sequence,

0). Files tha

on and Lum

quences to

ost-boxes as

a conservat

grasslands, p

calls

as an index

I surveyed b

at detector c

ensland). De

d frequency

de weatherp

.5 m, with

an 60 minut

gered autom

ector and At Site and b) t

n call data

s using

gle call sequ

ncy sweeps

separated b

at passed th

msden 2003

taxa by ex

s a tool in th

tion covena

plus restorat

x of bat act

bat activity a

connected to

etectors wer

y generator

roof plastic

the microp

tes before su

matically by

naBat ZCAIthe Strathkel

to a comp

AnalookW

uence was d

) identified

by more tha

hese criteria

). AnaSche

xtracting a r

he conservat

ant and have

tion of degr

tivity at the

at both sites

o an AnaBa

re calibrated

(AnaBat C

c boxes secu

phone facing

unset and e

ultrasonic n

IM storage ullar Conserva

puter using

version

defined as a

as bat echo

an 5 second

a were then

eme uses re

range of ca

tion of tree

e manageme

raded areas.

e Warrayur

s concurrent

at ZCAIM s

d by adjustin

Chirper 2, T

ured to the t

g upward a

ended at sun

noise.

units securedation Offset S

g CFCread

3.8s (C.

a file contai

olocation cal

s, was desig

processed

egional iden

all paramete

roosting mi

ent plans re

re and Strat

tly using ult

storage unit

ng their sen

Titley Electr

trunk of a tr

at an angle

nrise, during

d to trees atite.

version 4.3

Corben

ining at lea

lls (Houriga

gnated as a

using AnaS

ntification k

ers (Lumsd

icrobats

4

equiring

thkellar

trasonic

, Titley

nsitivity

ronics).

ree at a

of 45º

g which

t a) the

3s then

2011,

st three

an et al.

unique

Scheme

keys to

den and

Page 6: Roost boxes as a tool in the conservation of microbats ... · native habitat since European settlement has resulted in significant negative pressure on microbats (Microchiroptera),

Roost-boxes as a tool in the conservation of tree roosting microbats

5

Bennett 2005; Adams et al. 2010). I used an existing identification key developed to

identify the species in the Grampians region of Victoria (Arthur Rylah Institute for

Environmental Research 2012). Using the Grampians key, AnaScheme identifies call

sequences to species, except for Nyctophilus and Mormopterus, which are grouped by

genus. A list of microbat species known to occur in the Western Volcanic Plains of Victoria

is provided in Appendix 1.

Call sequences for all taxa recorded were grouped to investigate activity patterns of the

entire microbat community. Hourly activity patterns were recorded as the mean number of

call sequences recorded per hour after civil twilight for all species during the survey period.

Civil twilight times were accessed from Geoscience Australia (2012). Call sequences were

screened manually for buzz calls associated with pursuit and capture of prey. Foraging buzz

calls are characterised by a rapid increase in pulse repetition rate, slope, frequency and

speed (Griffin et al. 1960; Schnitzler et al. 1987). Call sequences were also examined for

social calls: distinctive audible contact calls made by bats from within roost sites before

leaving for the night to forage (Aldridge et al. 1990; Pfalzer and Kusch 2003; Arnold and

Wilkinson 2011).

Results

Bat activity

Over 39 consecutive nights from 20 April to 28 May 2012 (78 detector-nights) I identified

6,214 bat call sequences from 11,531 files (53.9%). A total of 5,474 and 740 call sequences

were recorded at the Warrayure and Strathkellar sites respectively. Bat activity was

recorded on every night at Warrayure, and on the majority of nights at Strathkellar (zero

call sequences recorded on 4 nights (10.3%)). Considerable variation in the nightly number

of bat call sequences was documented at both sites. An average (± standard error) of

140.4±26.7 call sequences per night was recorded at Warrayure, with a nightly maximum

of 775 calls (20 April 2012) and minimum of 15 (21 May 2012). At Strathkellar an average

of 19±5.2 call sequences per night was recorded, with a nightly maximum of 163 calls (20

April 2012) (Figure 2).

Page 7: Roost boxes as a tool in the conservation of microbats ... · native habitat since European settlement has resulted in significant negative pressure on microbats (Microchiroptera),

FigurStrath Hour

Hour

hour

more

sites.

hour

Figurthe Wcivil tw

e 2. Numbehkellar Conse

rly activity

rly activity p

after civil tw

e than 60%

A total of

prior to civi

e 3. Mean (±sWarrayure an

wilight.

Roo

er of echoloervation Offse

patterns wer

wilight (Wa

of all calls

45 call seq

il twilight, i

s.e.) number d Strathkella

ost-boxes as

cation call set Sites.

re similar at

arrayure 18.

were record

quences (0.8

i.e. prior to n

of bat call sear Conservati

s a tool in th

sequences re

t both sites w

.9% and Str

ded in the f

8% of total

nightfall (Fi

equences per ion Offset Sit

he conservat

ecorded each

with greates

rathkellar 41

first four ho

calls) were

igure 3).

night detectetes. The -1 ca

tion of tree

h night at t

st activity re

1.6% of call

ours after civ

recorded at

ed each hour ategory refers

roosting mi

the Warrayu

ecorded in t

l sequences)

vil twilight

t Warrayure

after civil tws to the hour

icrobats

6

ure and

the first

), while

at both

e in the

wilight at prior to

Page 8: Roost boxes as a tool in the conservation of microbats ... · native habitat since European settlement has resulted in significant negative pressure on microbats (Microchiroptera),

Spec

Using

level

ident

to me

not b

were

tasma

timor

austr

flaviv

(16.2

Figurof calgouldiNyctopV. regSacco

Fora

One h

(2% o

ies diversit

g the AnaS

(or to genu

ified either

eet the mini

be distinguis

identified

aniensis, M

riensis, Vesp

ralis, and

ventris. Vesp

2%), while V

e 4. Echolocall sequences ii; Cm, C. mphilus geoffrogulus; Vv, V

olaimus flavive

aging activit

hundred and

of total calls

Roo

ty

Scheme prog

us for Nycto

as a result

imum criter

shed betwee

d: Vespert

Minopteurs

padelus dar

Mormopter

padelus dar

V. vulturnus

ation call seqwere not ide

morio; Ft, Faoyi/N. gouldi/V. vulturnus;entris.

ty

d fifteen fee

s) and 13 at

ost-boxes as

gram, 2828

ophilus sp.,

of being a

ria, or havin

en two speci

tilionidae:

schreibers

rlingtoni, V

rus species

rlingtoni wa

dominated

uences identientified at Stalsistrellus ta/N. timoriensi; Ta, Tadarid

eding buzze

t Strathkella

s a tool in th

8 (45.5%) c

and Mormo

short seque

ng the majo

ies (Gibson

Chalinolob

sii bassani

V. regulus,

s 4/M. spe

as the most c

calls at Stra

ified to specietrathkellar anasmaniensis; is; Sb, Scotoreda australis;

es were reco

ar (1.8% of t

he conservat

call sequenc

opterus sp.)

nce with in

ority of puls

n and Lumsd

bus gouldi

ii, Nyctoph

and V. vult

ecies 2; E

commonly r

athkellar (43

es or genus bnd WarrayurMsb, Miniopepens balston

M, Mormop

orded during

total calls) (

tion of tree

ces were ide

). The rema

nsufficient g

ses with par

den 2003). T

ii, C. mo

hilus geoff

turnus; Mol

Emballonuri

recorded sp

3.9%) (Figu

by AnaSchemre respectivepterus schreibni; Vd, Vespadpterus specie

g this study,

(Figure 5).

roosting mi

entified to

ainder could

good quality

rameters tha

The followi

orio, Falsi

ffroyi/N. go

lossidae: Ta

idae: Sacco

ecies at Wa

ure 4).

me; 25.8% andely. Cg, Chalbersii oceanedelus darlingtes2/M. specie

, 112 at Wa

icrobats

7

species

d not be

y pulses

at could

ng taxa

istrellus

ouldi/N.

adarida

olaimus

arrayure

d 58.4% linolobus ensis; N, toni; Vr,

es 4; Sf,

arrayure

Page 9: Roost boxes as a tool in the conservation of microbats ... · native habitat since European settlement has resulted in significant negative pressure on microbats (Microchiroptera),

FigurStrath

Socia

Socia

provi

matur

separ

Strath

speci

regio

Warr

FigurWarr

e 5. Numberhkellar Conse

al calls

al calls wer

iding indire

re E. clado

rate occasio

hkellar site.

ies level, as

on are not

rayure are pr

e 6. Number ayure Conser

Roo

r of echolocaervation Offse

re recorded

ect evidence

ocalyx at th

ons at Wa

It was not p

reference s

available. E

resented in

of microbat rvation Offse

ost-boxes as

ation foraginet Sites.

in the hour

e of bats ro

his site. A m

arrayure (F

possible to

social calls p

Examples o

Appendix 2

social calls ret Site.

s a tool in th

ng buzz calls

r prior to c

osting diurn

maximum o

igure 6). N

identify the

produced by

of two soci

2.

ecorded each

he conservat

recorded ea

civil twiligh

nally in tree

of 16 socia

No social

e social calls

y species in

ial calls rec

h night in the

tion of tree

ach night at

ht on 17 nig

e hollows w

al calls was

calls were

s recorded d

n the Weste

corded on

hour prior t

roosting mi

the Warray

ghts at War

within the s

s recorded

e recorded

during this s

rn Volcanic

20 April 2

to civil twiligh

icrobats

8

yure and

rrayure,

stand of

on two

at the

study to

c Plains

2012 at

ht at the

Page 10: Roost boxes as a tool in the conservation of microbats ... · native habitat since European settlement has resulted in significant negative pressure on microbats (Microchiroptera),

Roost-boxes as a tool in the conservation of tree roosting microbats

9

Discussion

Considerable variation in activity was documented both within and between nights during

this study, however bats were present (as detected via echolocation calls) on every night at

Warrayure, and on the majority of nights at Strathkellar, albeit at lower levels (Figure 2).

The overall hourly pattern of bat activity featured a peak immediately after dusk, followed

by a steady decrease throughout the night and only minimal activity in the second half of

the night (Figure 3). Studies by Law et al. (1998) and Milne et al. (2005b) examining bat

activity in southeastern and northern Australia respectively have reported similar patterns.

In both studies, the majority of activity occurred in the first half of the night, with

considerably less activity from midnight to sunrise. Several lines of evidence suggest this

pattern is driven primarily by activity of nocturnal insects which also exhibit the greatest

peak of activity at twilight, with activity dropping off during the remainder of the night

until dawn (Rautenbach et al. 1988; Vaughaun et al. 1996; Jetz et al. 2003;

Wickramasinghe et al. 2003a,b; Weinbeer et al. 2006).

High levels of variation in bat activity between nights has also been identified in previous

studies (Hayes 1997; Milne et al. 2005b). A range of factors have been shown to influence

variation in nightly bat activity, including climate variables (Bullen and McKenzie 2005;

Turbill 2008; Brooks 2009), insect availability (Bell 1980; Avila-Flores et al. 2005;

Akasaka et al. 2009), and lunar phase (Fenton et al. 1977; Milne et al. 2005a). While

specific factors influencing activity patterns were not examined here, the findings clearly

show bats were consistently present in this highly-modified agricultural landscape. The

assemblage recorded comprised the majority of species known to occur in the Western

Volcanic Plains region of Victoria (Figure 4, Appendix 1). However, a conservative

approach should be taken when drawing inferences from a survey based solely on passively

collected echolocation data, as detection rates can differ significantly between species

(Hayes 2000; Sherwin et al. 2000). To provide greater confidence in recording an accurate

species inventory, capture techniques (e.g. harp trapping or mist netting) and physical

identification of animals should be employed in conjunction with bat detector sampling

(Australasian Bat Society 2006).

Page 11: Roost boxes as a tool in the conservation of microbats ... · native habitat since European settlement has resulted in significant negative pressure on microbats (Microchiroptera),

Roost-boxes as a tool in the conservation of tree roosting microbats

10

Minimal levels of foraging activity were recording during this study, predominantly at the

Warrayure site (Figure 5). Microbats are known to forage in a range of forested and rural

landscapes of southeastern Australia (Law et al. 1998; Lumsden and Bennett 1995), as well

as in highly-modified urban landscapes (Threlfall et al. 2012). Lumsden and Bennett (2000)

suggested bats are likely to play an important role in regulating populations of herbivorous

invertebrates around sparsely scattered trees, because insectivorous birds dependent on

good quality forest cover are often scarce in agricultural landscapes. Many land managers

are familiar with the beneficial role of birds as insectivores or pollinators (Reid and

Landsberg 1999; Lindemayer et al. 2011; Schirmer et al. 2012). However, because

insectivorous bats are small, nocturnal and remain hidden in roosts during the day, few

people are aware of the ecosystem services they provide by eating insects in rural

landscapes (Lumsden and Bennet 2005).

The presence and activity of microbats recorded in this study, and specifically the detection

of social calls produced by diurnally roosting bats (Figure 6), has implications for the

management and rehabilitation of isolated patches of land within highly-modified

agricultural landscapes. Across much of southeastern Australia, the amount of land

dedicated to conservation of biodiversity is generally small in size compared with the

extensive areas transformed for urbanisation, industrialisation and agricultural development

(Gobbons and Lindenmayer 2002; Lumsden and Bennett 2005; Rhodes et al. 2006).

Consequently, relatively small sites dedicated to conservation and restoration often exist

within a matrix of natural, semi-natural and highly modified habitats (Lindemayer et al.

2011). The mobility of bats means they can traverse the rural landscape to exploit multiple,

isolated habitat patches (Lumsden and Bennett 2000). Therefore, the management of

sparsely scattered and even single trees in the agricultural landscape can provide significant

conservation outcomes for bats (Lumsden and Bennett 1995; Law et al. 1998).

In areas with limited roosting sites, bat-boxes can be installed as substitutes for natural

hollows. In Australia, relatively little is known of the use and preferred designs of bat-

boxes (Goldingay and Stevens 2009), however studies in the Northern Hemisphere provide

evidence that internal microclimate, height off the ground, construction material and design

can affect the attractiveness and suitability of a bat-box for the target species (Arnett and

Page 12: Roost boxes as a tool in the conservation of microbats ... · native habitat since European settlement has resulted in significant negative pressure on microbats (Microchiroptera),

Roost-boxes as a tool in the conservation of tree roosting microbats

11

Hayes 2000; Brittingham and Williams 2000; Kerth et al. 2001; Flaquer et al. 2005;

Bartonicka and Rehak 2007; Lesinski et al. 2009). Therefore, the design and placement of

roost boxes is likely to influence their usefulness to bats, and other wildlife, and their

effectiveness in allowing the maintenance of sustainable populations of native wildlife in

lower-quality, fragmented habitats.

Conclusions

The loss, modification and fragmentation of natural habitats constitute a major threat to bat

populations worldwide (Racey and Entwistle 2003). This is particularly relevant in

Australia where the majority of microbats rely on tree hollows to roost in during the day

(Churchill 2008). Echolocation data collected in this study provide indirect evidence of bats

roosting diurnally in a small, isolated patch of trees located within a highly-modified

agricultural matrix. Bat-boxes were installed at the Warrayure Conservation Offset Site as

part of an ongoing ecological management and restoration plan, with the intention of

providing supplementary microbat roosting sites (Appendix 3). Further work is planned to

incorporate 6-monthly manual checks of bat-box occupancy at Warrayure, paired with

additional bat detector surveys to examine seasonal patterns of activity at both the

Warrayure and Strathkellar Conservation Offset Sites. This research will provide insight

into the efficacy of bat-boxes as a management tool for the conservation of tree-hollow

roosting microbats in highly-fragmented rural landscapes.

Page 13: Roost boxes as a tool in the conservation of microbats ... · native habitat since European settlement has resulted in significant negative pressure on microbats (Microchiroptera),

Roost-boxes as a tool in the conservation of tree roosting microbats

12

References

Adams, M. D., Law, B. S., and Gibson, M. S. (2010). Reliable automation of bat call identification for eastern New South Wales, Australia, using classification trees and AnaScheme software. Acta Chiropterologica 12(1): 231-245.

Akasaka, T., Nakano, D., and Nakamura, F. (2009). Influence of prey variables, food supply, and river restoration on the foraging activity of Daubenton's bat (Myotis daubentonii) in the Shibetsu River, a large lowland river in Japan. Biological Conservation 142(7): 1302-1310.

Aldridge, H. D. J. N., Obrist, M., Merriam, H. G., and Fenton, M. B. (1990). Roosting, vocalizations, and foraging by the African Bat, Nycteris thebaica. Journal of Mammalogy 71(2): 242-246.

Arnett, E. B. and Hayes, J. P. (2000). Bat use of roosting boxes installed under flat-bottom bridges in western Oregon. Wildlife Society Bulletin 28(4): 890-894.

Arnold, B. D. and Wilkinson, G. S. (2011). Individual specific contact calls of pallid bats (Antrozous pallidus) attract conspecifics at roosting sites. Behavioral Ecology and Sociobiology 65(8): 1581-1593.

Arthur Rylah Institute for Environmental Research (2012). AnaScheme microbat echolocation key: Grampians region, Victoria. Department of Sustainability and Environment Victoria, Heidelberg.

Avila-Flores, R. and Fenton, M. B. (2005). Use of spatial features by foraging insectivorous bats in a large urban landscape. Journal of Mammalogy 86(6): 1193-1204.

Australasian Bat Society (2006). Recommendations of the Australasian Bat Society Inc for reporting standards for insectivorous bat surveys using bat detectors. The Australasian Bat Society Newsletter 27: 6-9.

Bartonicka, T. and Rehak, Z. (2007). Influence of the microclimate of bat boxes on their occupation by the soprano pipistrelle Pipistrellus pygmaeus: possible cause of roost switching. Acta Chiropterologica 9(2): 517-526.

Bell, G. P. (1980). Habitat use and response to patches of prey by desert insectivorous bats. Canadian Journal of Zoology 58(10): 1876-1883.

Brittingham, M. C. and Williams, L. M. (2000). Bat boxes as alternative roosts for displaced bat maternity colonies. Wildlife Society Bulletin 28(1): 197-207.

Brooks, R. T. (2009). Habitat-associated and temporal patterns of bat activity in a diverse forest landscape of southern New England, USA. Biodiversity and Conservation 18(3): 529-545.

Page 14: Roost boxes as a tool in the conservation of microbats ... · native habitat since European settlement has resulted in significant negative pressure on microbats (Microchiroptera),

Roost-boxes as a tool in the conservation of tree roosting microbats

13

Bullen, R. D. and McKenzie, N. L. (2005). Seasonal range variation of Tadarida australis (Chiroptera: Molossidae) in Western Australia: the impact of enthalpy. Australian Journal of Zoology 53(3): 145-156.

Churchill, S. K. (2008). Australian Bats. Allen and Unwin, NSW.

Department of the Environment, Water, Heritage and the Arts (2008). Natural Temperate Grassland of the Victorian Volcanic Plain: A Nationally Threatened Ecological Community. Australian Government, Canberra.

Department of Sustainability, Environment, Water, Population and Communities (2011). Nationally Threatened Ecological Communities of the Victorian Volcanic Plain: Natural Temperate Grassland and Grassy Eucalypt Woodland. Australian Government, Canberra.

Duncan, A., Baker, B., Hall, L, Richards, G., Tideman, C., Clague, C., and Law, B (1999). The Action Plan for Australian Bats: Threatening processes and conservation priorities. Threatened Species & Ecological Communities. Environment Australia, Canberra.

Fenton, M. B., Boyle, N. G. H., Harrison, T. M., and Oxley, D. J. (1977). Activity patterns, habitat use, and prey selection by some African insectivorous bats. Biotropica 9(2): 73-85.

Flaquer, C., Torre, I., and Ruiz-Jarillo, R (2005). The value of bat-boxes in the conservation of Pipistrellus pygmaeus in wetland rice paddies. Biological Conservation 128(2): 223-230.

Gibson, M. S. and Lumsden, L. F. (2003). The AnaScheme automated bat call identification system. The Australasian Bat Society Newsletter 20: 24-27.

Gibbons, P. and Lindenmayer, D. (2002). Tree hollows and wildlife conservation in Australia. CSIRO Publishing, Melbourne.

Goldingay, R. L. (2009). Characteristics of tree hollows used by Australian birds and bats. Wildlife Research 36(5): 394-409.

Goldingay, R. L. (2011). Characteristics of tree hollows used by Australian arboreal and scansorial mammals. Australian Journal of Zoology 59(5): 277-294.

Goldingay, R. L. and Stevens, J. R. (2009). Use of artificial tree hollows by Australian birds and bats. Wildlife Research 36(2): 81-97.

Griffin, D. R., Webster, F. A., and Michael, C. R. (1960). The echolocation of flying insects by bats. Animal Behaviour 8(1): 141-154.

Harper, M. J., McCarthy, M. A., and van der Ree, R. (2005a). The abundance of hollow-bearing trees in urban dry sclerophyll forest and the effect of wind on hollow development. Biological Conservation 122(2): 181-192.

Page 15: Roost boxes as a tool in the conservation of microbats ... · native habitat since European settlement has resulted in significant negative pressure on microbats (Microchiroptera),

Roost-boxes as a tool in the conservation of tree roosting microbats

14

Harper, M. J., McCarthy, M. A., and van der Ree, R. (2005b). The use of nest boxes in urban natural vegetation remnants by vertebrate fauna. Wildlife Research 32(6): 509-516.

Harvey, C. A., Villanueva, C., Esquivel, H., Gomez, R., Ibrahim, M., Lopez, M., Martinez, J., Munoz, D., Restrepo, C., Saenz, J. C.,Villacis, J., and Sinclair, F. L. (2011). Conservation value of dispersed tree cover threatened by pasture management. Forest Ecology and Management 261(10): 1664-1674.

Hayes, J. P. (1997). Temporal variation in activity of bats and the design of echolocation-monitoring studies. Journal of Mammalogy 78(2): 514-524.

Hayes, J. P. (2000). Assumptions and practical considerations in the design and interpretation of echolocation-monitoring studies. Acta Chiropterologica 2(2): 225-236.

Hourigan, C. L., Catterall, C. P., Jones, D., and Rhodes, M. (2008). A comparison of the effectiveness of bat detectors and harp traps for surveying bats in an urban landscape. Wildlife Research 35(8): 768-774.

Irvine, R., and Bender, R. (1995). Initial results from bat roosting boxes at Organ Pipes National Park. Victorian Naturalist 112: 212-217.

Jetz, W., Steffen, J., and Linsenmair, K. E. (2003). Effects of light and prey availability on nocturnal, lunar and seasonal activity of tropical nightjars. Oikos 103(3): 627-639.

Kavanagh, R. P., Stanton, M. A., and Herring. M. W. (2007). Eucalypt plantings on farms benefit woodland birds in south-eastern Australia. Austral Ecology 32(6): 635-650.

Kerth, G., Weissmann, K., and Konig, B. (2001). Day roost selection in female Bechstein's bats (Myotis bechsteinii): a field experiment to determine the influence of roost temperature. Oecologia 126(1): 1-9.

Kunz, T. H. and Lumsden, L. F. (2003). Ecology of cavity and foliage roosting bats. In: Kunz, T. H., Fenton, M. B. (Eds.), Bat Ecology. The University of Chicago Press, Chicago, pages 3-89.

Law, B., Anderson, J., and Chidel, M. (1998). A bat survey in state forests on the south-west slopes region of New South Wales with suggestions of improvements for future surveys. Australian Zoologist 30:467-479.

Lesinski, G., Skrzypiec-Nowak, P., Janiak, A., and Jagnieszczak, Z. (2009). Phenology of bat occurrence in boxes in central Poland. Mammalia 73(1): 33-37.

Lindenmayer, D. B., Archer, S., Barton, P., Bond, S., Crane, M., Gibbons, P., Kay, G., MacGregor, C., Manning, A., Michael, D., Montague-Drake, R., Munro, N., Muntz, R., and Stagoll, K. (2011). What makes a good paddock for biodiversity? In: Lindenmayer, D. B. (Ed.), What Makes a Good Farm for Wildlife? CSIRO Publishing, Collingwood, pages 71-95.

Page 16: Roost boxes as a tool in the conservation of microbats ... · native habitat since European settlement has resulted in significant negative pressure on microbats (Microchiroptera),

Roost-boxes as a tool in the conservation of tree roosting microbats

15

Lindenmayer, D. B., Cunningham, R. B., Donnelly, C. F., Tanton, M. T., and Nix, H. A. (1993). The abundance and development of cavities in Eucalyptus trees - a case-study in the Montane forests of Victoria, southeastern Australia. Forest Ecology And Management 60(1-2): 77-104.

Lumsden, L. F. and Bennett, A. F. (1995). Bats of a semi-arid environment in south-eastern Australia: biogeography, ecology and conservation. Wildlife Research 22(2): 217-239.

Lumsden, L.F. and Bennett, A.F. (2000). Bats in rural landscapes: a significant but largely unknown faunal component. In: Barlow, T., Thorburn, R. (Eds.), Balancing Conservation and Production in Grassy Landscapes. Proceedings of the Bushcare Grassy Landscapes Conference, Clare, South Australia, 19-21 August 1999. Environment Australia, Canberra, pages 42-50.

Lumsden, L. F. and Bennett, A. F. (2005). Scattered trees in rural landscapes: foraging habitat for insectivorous bats in south-eastern Australia. Biological Conservation 122(2): 205-222.

Milne, D. J., Armstrong, M., Fisher, A., Flores, T., and Pavey, C. R. (2005a). Structure and environmental relationships of insectivorous bat assemblages in tropical Australian savannas. Austral Ecology 30(8): 906-919.

Milne, D. J., Fisher, A., Rainey, I., and Pavey, C. R. (2005b). Temporal patterns of bats in the Top End of the Northern Territory, Australia. Journal of Mammalogy 86(5): 909-920.

Newton, I. (1994). The role of nest sites in limiting the numbers of hole-nesting birds: a review. Biological Conservation 70(3): 265-276.

Nowak, R. M. (1999). Walker’s Mammals of the World. 6th Edition. John Hopkins University Press, Baltimore & London.

Pfalzer, G. and Kusch, J. (2003). Structure and variability of bat social calls: implications for specificity and individual recognition. Journal of Zoology 261(1): 21-33.

Racey, P. A. and Entwistle, A. (2003). Conservation ecology of bats. In: Kunz, T. H., Fenton, M. B. (Eds.), Bat Ecology. The University of Chicago Press, Chicago, pages 680-743.

Rautenbach, I. L., Kemp, A. C., and Scholtz, C. H. (1988). Fluctuations in availability of arthropods correlated with microchiropteran and avian predator activities. Koedoe 31:77-90.

Reid, N. and Landsberg, J. (1999). Tree decline in agricultural landscapes: what we stand to lose. In: Hobbs, R.J., Yates, C.J. (Eds.), Temperate Eucalypt Woodlands in Australia: Biology, Conservation, Management and Restoration. Surrey Beatty & Sons, Chipping Norton, pages 127-166.

Page 17: Roost boxes as a tool in the conservation of microbats ... · native habitat since European settlement has resulted in significant negative pressure on microbats (Microchiroptera),

Roost-boxes as a tool in the conservation of tree roosting microbats

16

Rhodes, M., Wardell-Johnson, G. W., Rhodes, M. P., and Raymond, B. (2006). Applying network analysis to the conservation of habitat trees in urban environments: a case study from Brisbane, Australia. Conservation Biology 20(3): 861-870.

Schirmer, J., Clayton, H., and Sherren, K. (2012). Reversing scattered tree decline on farms: implications of landholder perceptions and practice in the Lachlan catchment, New South Wales. Australasian Journal of Environmental Management 19(2): 91-107.

Schnitzler, H-U., Kalko, E. K. V., Miller, L. A., and Surlykke, A. (1987). The echolocation and hunting behavior of the bat, Pipistrellus kuhli. Journal of Comparative Physiology a-Sensory Neural and Behavioral Physiology 161(2): 267-274.

Sherwin, R. E., Gannon, W. L., and Haymond, S. (2000). The efficacy of acoustic techniques to infer differential use of habitat by bats. Acta Chiropterologica 2(2): 145-153.

Smith, G. C. and Agnew, G. (2002). The value of ‘bat boxes’ for attracting hollow- dependent fauna to farm forestry plantations in southeast Queensland. Ecological Management & Restoration 3(1): 37-46.

Threlfall, C. G., Law, B., and Banks, P. B. (2012). Influence of landscape structure and human modifications on insect biomass and bat foraging activity in an urban landscape. PLoS ONE 7(6): 10.

Turbill, C. (2006). Roosting and thermoregulatory behaviour of male Gould's long-eared bats, Nyctophilus gouldi: energetic benefits of thermally unstable tree roosts. Australian Journal of Zoology 54(1): 57-60.

Turbill, C. (2008). Winter activity of Australian tree-roosting bats: influence of temperature and climatic patterns. Journal of Zoology 276(3): 285-290.

Vaughan, N., Jones, G., and Harris, S. (1996). Effects of sewage effluent on the activity of bats (Chiroptera: Vespertilionidae) foraging along rivers. Biological Conservation 78(3): 337-343.

Weinbeer, M., Meyer, C. F., and Kalko, E. K. V. (2006). Activity pattern of the trawling phyllostomid bat, Macrophyllum macrophyllum, in Panamá. Biotropica 38(1): 69-76.

Weinberg, A., Gibbons, P., Briggs, S. V., and Bonser, S. P. (2011). The extent and pattern of Eucalyptus regeneration in an agricultural landscape. Biological Conservation 144(1): 227-233.

Page 18: Roost boxes as a tool in the conservation of microbats ... · native habitat since European settlement has resulted in significant negative pressure on microbats (Microchiroptera),

Roost-boxes as a tool in the conservation of tree roosting microbats

17

Appendix 1

Table A1. Microbat species known to occur in the Western Volcanic Plains region of Victoria*, with level of taxonomic identification achieved by the AnaScheme program. Scientific name Common name AnaScheme identification Vespertilionidae Chalinolobus gouldii Gould’s wattled bat Species level Chalinolobus morio chocolate wattled bat Species level Falsistrellus tasmaniensis eastern false pipistrelle Species level Miniopterus schreibersii oceanensis southern bent-winged bat Species level Nyctophilus geoffroyi lesser long-eared bat Genus: Nyctophilus sp. Nyctophilus gouldi Gould’s long-eared bat Genus: Nyctophilus sp. Nyctophilus timoriensis greater long-eared bat Genus: Nyctophilus sp. Scotorepens balstoni inland broad-nosed bat Species level Vespadelus darlingtoni large forest bat Species level Vespadelus regulus southern forest bat Species level Vespadelus vulturnus little forest bat Species level Molossidae Tadarida australis white-striped free-tailed bat Species level Mormopterus species 2 eastern free-tailed bat Genus: Mormopterus sp. Mormopterus species 4 southern free-tailed bat Genus: Mormopterus sp. Emballonuridae Saccolaimus flaviventris yellow-bellied sheath-tailed bat Species level

*Species distributions from Churchill (2008) and on advice from L. Lumsden (Department of Sustainability and Environment, Victoria).

Page 19: Roost boxes as a tool in the conservation of microbats ... · native habitat since European settlement has resulted in significant negative pressure on microbats (Microchiroptera),

Roost-boxes as a tool in the conservation of tree roosting microbats

18

Appendix 2

a)

b)

Figure A1. Sonograms of social call sequences produced by microbats roosting diurnally in mature E. cladocalyx at the Warrayure Conservation Offset Site. Call frequency (kHz) is presented on the y-axis, with time in seconds on the x-axis (call a) 15 second duration; call b) 12 second duration). Both call sequences are comprised of short pulses of sound within the range 7-10 kHz.

Page 20: Roost boxes as a tool in the conservation of microbats ... · native habitat since European settlement has resulted in significant negative pressure on microbats (Microchiroptera),

App

Figurbuilt b

pendix 3

e A2. a) Stanbat-boxes inst

Roo

d of mature Etalled on the

ost-boxes as

E. cladocalyxtrunks of E. c

s a tool in th

at the Warracladocalyx at

he conservat

ayure ConserWarrayure o

tion of tree

rvation Offseton 29 May 20

roosting mi

t Site. b,c,d) P012.

icrobats

19

Purpose-