A REVIEW OF EMPIRICAL DATA IN RESPECT OF...

June 2017 – Version 4

A REVIEW OF EMPIRICAL

DATA IN RESPECT OF

EMERGENCE AND RETURN

TIMES REPORTED FOR

THE UK’s 17 NATIVE

BAT SPECIES

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CONTENTS

1. RATIONALE …………………………………………….......................…... 1

2. BARBASTELLE Barbastella barbastellus ………….........……………….. 2

3. SEROTINE Eptesicus serotinus …………………………………..……….. 3

4. BECHSTEIN’S BAT Myotis bechsteinii …………………………......……. 5

5. ALCATHOE’S BAT Myotis alcathoe ……...................................………... 7

6. BRANDT’S BAT Myotis brandtii …………………….............……………. 8

7. DAUBENTON’S BAT Myotis daubentonii ………………………………... 10

8. WHISKERED BAT Myotis mystacinus ……………………..............…….. 13

9. NATTERER’S BAT Myotis nattereri ……….........................................….. 15

10. LEISLER’S BAT Nyctalus leisleri ………………………….......…………. 17

11. NOCTULE Nyctalus noctula ………………………………………......…... 19

12. NATHUSIUS’ PIPISTRELLE Pipistrellus nathusii ……………………... 21

13. COMMON PIPISTRELLE Pipistrellus pipistrellus ……………………... 22

14. SOPRANO PIPISTRELLE Pipistrellus pygmaeus ……………………..... 24

15. GREY LONG-EARED BAT Plecotus austriacus ……………...……….... 26

16. BROWN LONG-EARED BAT Plecotus auritus ……………...………..... 28

17. GREATER HORSESHOE-BAT Rhinolophus ferrumequinum ….……... 30

18. LESSER HORSESHOE-BAT Rhinolophus hipposideros ……………...... 32

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Report authors: Henry Andrews MSc CEcol MCIEEM & Louis Pearson BSc MSc GradCIEEM.

Final proof: Leanne Butt BSc MSc ACIEEM.

No part of this report may be reproduced without prior written approval of AEcol. Legal action may be taken

in the event of copyright infringement.

No liability is accepted in respect of the use of data, conclusions or other material contained in this report.

© AEcol 2016

AEcol & Andrews Ecology Ltd are registered Trademarks.

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AECOL LTD REVIEW OF EMERGENCE AND RETURN EMPIRICAL DATA

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- 1 - © AEcol 2017

A REVIEW OF EMPIRICAL DATA IN RESPECT OF EMERGENCE AND

RETURN TIMES REPORTED FOR THE UK’s 17 NATIVE BAT SPECIES

1. RATIONALE

1.1.1 To date, no publication or guidance document has reviewed empirical evidence in

respect of emergence times, nor has any document attempted to provide the times of

returns.

1.1.2 Emergence times quoted within grey literature may comprise approximations based

on the author’s own experiences, or cite either mean or median averages in white

literature. All emergence times cited are therefore individual figures. Where the time

given is an approximation the number of observations is unknown to the reader, and

where they are mean averages the figure may be entirely artificial.

1.1.3 Although the mean average and standard deviation (SD) may be of scientific interest,

in order to perform robust emergence and return surveillance in the context of an

Ecological Impact Assessment, it is necessary to understand the temporal range

across which the target species might emerge and return in that particular season.

This is because it informs the duration over which each survey should be performed.

1.1.4 This review of empirical data in respect of emergence and return times reported for

the UK’s 17 native bat species was performed. White-papers and scientific reports

were reviewed and the data collated for each species. Where appropriate, sunset and

sunrise times were factored in to assess the data reported against sunset in the

specific locality. Where more than one publication exists for an individual bat

species, studies in the UK were preferentially selected, as were those that span both

sexes and the greatest number of periods. However, in order that the reader is aware

of alternative texts, other studies that were of note are included within the reference

lists as ‘see also’.

1.1.5 It will be immediately apparent to the reader that many of the published accounts do

not include all the salient data, there is no standardised format for the presentation of

data in order that studies may be compared, and much of the data is presented in a

format that confounds interpretation and how the data might be put to practical use

does not appear to be considered.

1.1.6 The conclusion is that there is still room for improvement in the understanding of the

temporal behaviour of all species. Immediate gains would result from the

presentation of range data and SD within white literature.

1.1.7 The review was performed by Henry Andrews MSc CEcol MCIEEM and Louis

Pearson BSc MSc GradCIEEM of AEcol. We would be pleased to hear from anyone

who has or knows of conflicting or more detailed accounts, or has data of their own

they would be prepared to share.


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2. BARBASTELLE Barbastella barbastellus

2.1 Emergence times

2.1.1 Table 1 details the emergence times for the barbastelle Barbastella barbastellus

reported in empirical data (Zeale et al. 2012).

Table 1. Emergence times for the barbastelle Barbastella barbastellus reported

in empirical data (Zeale et al. 2012).

SEX

AVERAGE

EMERGENCE TIME

(minutes before/after

sunset)

± RANGE*


sunset)

ACTUAL RANGE


sunset)

F Mean: 24 minutes after

(Zeale et al. 2012)

SD: 17.1-30.9 minutes

after (Zeale et al. 2012)

12-36 minutes after

(Zeale et al. 2012)

M No data No data No data * range in which 95% of observations occurred.

2.2 Return times

2.2.1 Table 2 details the return times for the barbastelle reported in empirical data (Zeale

et al. 2012).

Table 2. Return times for the barbastelle Barbastella barbastellus reported in

empirical data (Zeale et al. 2012).

SEX SEASON

AVERAGE

RETURN TIME

(minutes

before/after

sunrise)

± RANGE*

(minutes

before/after

sunrise)

ACTUAL

RANGE

(minutes

before/after

sunrise)

F Pregnancy –

May/June Mean: 194 minutes

before (3 hours and 14 minutes)

(Zeale et al. 2012)

SD: 254-135

minutes before (4 hours and 14 minutes up

to 2 hours and 15 minutes)

(Zeale et al. 2012)

No data F Lactation – July

F Weaning – August

M All No data No data No data * range in which 95% of observations occurred.

2.3 References

Zeale M, Davidson-Watts I & Jones G 2012. Home range use and habitat selection by

barbastelle bats (Barbastella barbastellus): implications for conservation. Journal of

Mammalogy 93(4): 1110-1118

See also: Russo D, Cistrone L, Jones G 2007. Emergence time in forest bats: the influence

of canopy closure. Acta Oecologica. 31: 119-126


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3. SEROTINE Eptesicus serotinus

3.1 Emergence times

3.1.1 Table 3 details the emergence times for the serotine Eptesicus serotinus reported in

empirical data (Catto et al. 1995, Petrželkova & Zákul 2001).

Table 3. Emergence times for the serotine Eptesicus serotinus reported in

empirical data (Catto et al. 1995, Petrželkova & Zákul 2001).

SEX

AVERAGE

EMERGENCE TIME


sunset)

± RANGE*


sunset)

ACTUAL RANGE


sunset)

F Mean: 11.6 minutes after

(Catto et al. 1995)


after (Catto et al. 1995)

15 minutes 45 seconds

before up to 42

minutes after

(Petrželkova & Zákul

2001)


3.2 Return times

3.2.1 Table 4 details the return times for the serotine reported in empirical data (Catto et

al. 1995).

Table 4. Return times for the serotine Eptesicus serotinus reported in empirical

data (Catto et al. 1995).

SEX SEASON

AVERAGE

RETURN TIME

(minutes

before/after

sunrise)

± RANGE*

(minutes

before/after

sunrise)

ACTUAL RANGE


sunrise)

F Pregnancy –

May/June No data No data

309-9 minutes before†

(5 hours and 9 minutes up to 9 minutes)

(Catto et al. 1995)

F Lactation –

July No data No data

199-66 minutes before† (3 hours and 19 minutes up to 1 hour six

and minutes)

(Catto et al. 1995)

F Weaning –

August No data No data

66 minutes before† (1 hour and six minutes)

(Catto et al. 1995)


† Inferred from data presented in figures.


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3.3 References

Catto C, Racey P & Stephenson P 1995. Activity patterns of the serotine bat (Eptesicus

serotinus) at a roost in southern England. J. Zool., Lond. 235:635-644

Petrželkova K & Zákul J 2001. Emergence behaviour of the serotine bat (Eptesicus

serotinus) under predation risk. Netherlands Journal of Zoology 51: 395-414

See also: Jones G & Rydell J 1994. Foraging strategy and predation risk as factors

influencing emergence time in echolocating bats. Phil. Trans. R. Soc. Lond. B 346: 445-455

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4. BECHSTEIN’S BAT Myotis bechsteinii

4.1 Emergence times

4.1.1 Table 5 details the emergence times for Bechstein’s bat Myotis bechsteinii reported

in empirical data (Jones & Rydell 1994, Dietz & Pir 2011).

Table 5. Emergence times for Bechstein’s bat Myotis bechsteinii reported in

empirical data (Jones & Rydell 1994, Dietz & Pir 2011).

SEX

AVERAGE

EMERGENCE TIME


sunset)

± RANGE*

(minutes

before/after

sunset)

ACTUAL RANGE

(minutes before/after sunset)

F

June – Median: 33

minutes after

(Jones & Rydell 1994)

No data Earliest starting 30 minutes after


Mean: 47 minutes after

(Dietz & Pir 2011) No data

2 minutes before up to 92

minutes after (2 minutes before up to 1 hour and 32 minutes

after)

(Dietz & Pir 2011)

M Mean: 56 minutes after

(Dietz & Pir 2011) No data

25-87 minutes after (25 minutes to 1 hour and 27 minutes)

(Dietz & Pir 2011) * range in which 95% of observations occurred.

4.2 Return times

4.2.1 Table 6 details the return times for Bechstein’s bat reported in empirical data (Dietz

& Pir 2011).

Table 6. Return times for Bechstein’s bat Myotis bechsteinii reported in

empirical data (Dietz & Pir 2011).

SEX SEASON

AVERAGE RETURN

TIME


sunrise)

± RANGE*

(minutes

before/after

sunrise)

ACTUAL RANGE


sunrise)

F Pregnancy –

May/June

60 minutes before

(Fitzsimons et al. 2002) No data

77-9 minutes before (1 hour and 17 minutes up to 9

minutes before)

(Dietz & Pir 2011)

F Lactation –

July

60 minutes before


F Weaning –

August

60 minutes before


M All No data No data

413 minutes before up

to 2 minutes after (6 hours and 53 minutes before

up to 2 minutes after) (Dietz & Pir 2011)

* range in which 95% of observations occurred.


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4.3 References

Dietz M & Pir J 2011. Distribution, Ecology and Habitat Selection by Bechstein’s bat

(Myotis bechsteinii) in Luxembourg. Ökologie der Säugetiere 6

Fitzsimons P, Hill D & Greenaway F 2002. Patterns of habitat use by female Bechstein’s

bats (Myotis bechsteinii) from a maternity colony in a British woodland. School of

Biological Sciences, University of Sussex

Jones G & Rydell J 1994. Foraging strategy and predation risk as factors influencing

emergence time in echolocating bats. Phil. Trans. R. Soc. Lond. B 346: 445-455

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5. ALCATHOE’S BAT Myotis alcathoe

5.1 Emergence times

5.1.1 Table 7 details the emergence times for Alcathoe’s bat Myotis alcathoe reported in

empirical data (observations provided by R. Baker & L. Whitfield 2016 pers comm.

16th October, and D. Whitby 2016 pers comm. 18th October).

Table 7. Emergence times for Alcathoe’s bat Myotis alcathoe reported in

empirical data (observations provided by R. Baker & L. Whitfield 2016 pers

comm. 16th October, and D. Whitby 2016 pers comm. 18th October).

SEX

AVERAGE

EMERGENCE TIME


sunset)

± RANGE*


sunset)

ACTUAL RANGE


sunset)

F

Mean: 1.1 minutes before

(calculated from 11

observations provided by

R. Baker & L. Whitfield

2016 pers comm. 16th

October, and D. Whitby


October)

SD: 16.4 minutes before

to 14.3 minutes after

(calculated from 11

observations provided by

R. Baker & L. Whitfield


October, and D. Whitby


October)

33 minutes before to 17

minutes after (calculated

from 11 observations

provided by R. Baker &

L. Whitfield 2016 pers

comm. 16th October, and

D. Whitby 2016 pers

comm. 18th October)


5.2 Return times

5.2.1 Table 8 details the return times for Alcathoe’s bat reported in empirical data.

Table 8. Return times for Alcathoe’s bat Myotis alcathoe reported in empirical

data.

SEX SEASON

AVERAGE

RETURN TIME

(minutes

before/after

sunrise)

± RANGE*

(minutes

before/after

sunrise)

ACTUAL

RANGE

(minutes

before/after

sunrise)

F Pregnancy –

May/June No data No data No data

F Lactation – July No data No data No data

F Weaning – August No data No data No data


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6. BRANDT’S BAT Myotis brandtii

6.1 Emergence times

6.1.1 Table 9 details the emergence times for Brandt’s bat Myotis brandtii reported in

empirical data (Berge 2007).

Table 9. Emergence times for Brandt’s bat Myotis brandtii reported in empirical

data (Berge 2007).

SEX

AVERAGE

EMERGENCE TIME


sunset)

± RANGE*


sunset)

ACTUAL RANGE


sunset)

F Mean: 43.3 minutes after

(Berge 2007) No data No data

F

May/June – Mean: 27.3

minutes after (Schmidt

2007)

23.7-30.9 minutes after

(Schmidt 2007)

20-35 minutes after

(Schmidt 2007)

F

July – Mean: 21.4


2007)

18-24.8 minutes after

(Schmidt 2007)

16-27 minutes after

(Schmidt 2007)

F

August – Mean: 24.9


2007)

20.8-29 minutes after

(Schmidt 2007)

21-34 minutes after

(Schmidt 2007)


6.2 Return times

6.2.1 Table 10 details the return times for Brandt’s bat reported in empirical data (Berge

2007).

Table 10. Return times for Brandt’s bat Myotis brandtii reported in empirical

data (Berge 2007).

SEX SEASON

AVERAGE RETURN

TIME


sunrise)

± RANGE*

(minutes

before/after

sunrise)

ACTUAL

RANGE

(minutes

before/after

sunrise)

F Pregnancy –

May/June Mean: 135.5 minutes

before (2 hours and 15.5 minutes)

(Berge 2007)

No data No data

F Lactation – July No data No data

F Weaning – August No data No data



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6.3 References

Berge L 2007. Resource partitioning between the cryptic species Brandt's bat (Myotis

brandtii) and the whiskered bat (M. mystacinus) in the UK. A thesis submitted to the

University of Bristol in accordance with the requirements of the degree of PhD in the

Faculty of Science

Schmidt C 2007. Emergence behaviour of a nursery colony of Myotis brandtii (Eversmann,

1845) in Saxony. Myotis 43: 55-62

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7. DAUBENTON’S BAT Myotis daubentonii

7.1 Emergence times

7.1.1 Table 11 details the emergence times for Daubenton’s bat Myotis daubentonii

reported in empirical data (Encarnação et al. 2006, Dietz & Kalko 2007, Lučan

2009).

Table 11. Emergence times for Daubenton’s bat Myotis daubentonii reported in

empirical data (Encarnação et al. 2006, Dietz & Kalko 2007, Lučan 2009).

SEX

AVERAGE

EMERGENCE TIME


sunset)

± RANGE*


sunset)

ACTUAL RANGE


sunset)

F

May – Mean: 46.1

minutes after

(Dietz & Kalko 2007)

No data

17-94 minutes after


Lime kiln: 8-64 minutes

after (Lučan 2009)

Trees: 26-61 minutes

after (Lučan 2009)

June – Mean: 58.1

minutes after


No data

July/August – Mean: 43.1

minutes after


No data

Lime kiln – Mean: 28

minutes after

(Lučan 2009)

Lime kiln – SD: 16-40

minutes after

(Lučan 2009)

Trees – Mean: 45 minutes

after (Lučan 2009)

Trees – SD: 34-56

minutes after

(Lučan 2009)

M

May – Mean: 47.5

minutes after


No data No data

June – Mean: 67.2

minutes after


No data No data

July/August – Mean: 39.4

minutes after


No data No data


(Encarnação et al. 2006)

SD: 22-85 minutes after

(Encarnação et al. 2006)

25-220 minutes after

(Encarnação et al. 2006) * range in which 95% of observations occurred.

7.2 Return times

7.2.1 Table 12 on the following page details the return times for Daubenton’s bat reported

in empirical data (Encarnação et al. 2006, Dietz & Kalko 2007).


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Table 12. Return times for Daubenton’s bat Myotis daubentonii reported in

empirical data (Encarnação et al. 2006, Dietz & Kalko 2007).

SEX SEASON

AVERAGE

RETURN TIME


sunrise)

± RANGE*

(minutes

before/after

sunrise)

ACTUAL

RANGE

(minutes

before/after

sunrise)

F Pregnancy – May 46.3 minutes before

(Dietz & Kalko 2007) No data

c. 70-30 minutes

before† (1 hour and 10 minutes

up to 30 minutes)

(Dietz & Kalko

2007)

F Lactation – June 40.5 minutes before

(Dietz & Kalko 2007) No data

90-10 minutes


up to 10 minutes before) (Dietz & Kalko

2007)

F Weaning –

July/August No data No data

c. 450-10 minutes

before† (7 hours and 30 minutes

up to 10 minutes)

(Dietz & Kalko

2007)

M

May

May – Mean: 110.8

minutes before


No data

c. 330-30 minutes

before † (5 hours and 30 minutes

up to 30 minutes)

(Dietz & Kalko

2007)

June No data No data

c. 180-20 minutes

before† (3 hours up to 20

minutes)

(Dietz & Kalko

2007

July/August No data No data

c. 80-20 minutes


up to 20 minutes)

(Dietz & Kalko

2007)

All

Mean: 152 minutes

before (2 hours and 32 minutes) (Encarnação et al.

2006)

SD: 299-5

minutes before (4 hours and 59 minutes

up to 5 minutes)

(Encarnação et al.

2006)

945-2 minutes

before (9 hours and 5 minutes

up to 2 minutes)

(Encarnação et al.

2006) * range in which 95% of observations occurred.

† figures cited are actually the range of means of returns on individual nights, but this was the only

data found.


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7.3 References

Dietz M & Kalko E 2007. Reproduction affects flight activity in female and male

Daubenton’s bat, Myotis daubentonii. Can. J. Zool. 85: 653-664

Encarnação J, Kierdorf U & Wolters V 2006. Seasonal variation in nocturnal activity of

male Daubenton’s bats, Myotis daubentonii (Chiroptera: Vespertilionidae). Folia Zool

55(3): 237-249

Lučan R 2009. Effect of colony size and reproductive period on the emergence behaviour of

a maternity colony of Daubenton’s bats (Myotis daubentonii) occupying an artificial

roost (Chiroptera: Vespertilionidae). Lynx 40: 71-81



See also: Nyholm E 1965. The ecology of Myotis mystacinus (Leisl.) and Myotis

daubentonii. Annales Zoologici Fennici 2: 77-123

See also: Shirley M, Armitage V, Barden T, Gough M, Lurz P, Oatway D, South A &

Rushton S 2001. Assessing the impact of a music festival on the emergence behaviour of a

breeding colony of Daubenton’s bats (Myotis daubentonii). J. Zool., Lond. 254: 367-373

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8. WHISKERED BAT Myotis mystacinus

8.1 Emergence times

8.1.1 Table 13 details the emergence times for the whiskered bat Myotis mystacinus

reported in empirical data (Jones & Rydell 1994, Berge 2007).

Table 13. Emergence times for the whiskered bat Myotis mystacinus reported in

empirical data (Jones & Rydell 1994, Berge 2007).

SEX

AVERAGE EMERGENCE

TIME


± RANGE*


sunset)

ACTUAL RANGE


sunset)

F

June – Median: 32 minutes after

(Jones & Rydell 1994) No data

Earliest starting 28

minutes after


Mean: 33.3 minutes after


M Mean: 33.3 minutes after



8.2 Return times

8.2.1 Table 14 details the return times for the whiskered bat reported in empirical data

(Berge 2007).

Table 14. Return times for the whiskered bat Myotis mystacinus reported in

empirical data (Berge 2007).

SEX SEASON

AVERAGE RETURN

TIME


sunrise)

± RANGE*

(minutes

before/after

sunrise)

ACTUAL

RANGE

(minutes

before/after

sunrise)

F Pregnancy –


before (2 hours and 6.9 minutes)

(Berge 2007)

No data No data



M All No data No data * range in which 95% of observations occurred.

8.3 References

Berge L 2007. Resource partitioning between the cryptic species Brandt's bat (Myotis

brandtii) and the whiskered bat (M. mystacinus) in the UK. A thesis submitted to the

University of Bristol in accordance with the requirements of the degree of PhD in the

Faculty of Science



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9. NATTERER’S BAT Myotis nattereri

9.1 Emergence times

9.1.1 Table 15 details the emergence times for Natterer’s bat Myotis nattereri reported in

empirical data (Jones & Rydell 1994, Swift 1997).

Table 15. Emergence times for Natterer’s bat Myotis nattereri reported in

empirical data (Jones & Rydell 1994, Swift 1997).

SEX

AVERAGE EMERGENCE

TIME


± RANGE*


sunset)

ACTUAL RANGE


sunset)

F

June – Median: 75 minutes

after (Jones & Rydell 1994) No data


minutes after


Median: 55.9 minutes after

(Swift 1997)


after (Swift 1997) No data

July – Mean: 31 minutes after

(Siemers et al. 1999)

SD: 22-41.2 minutes

after (Siemers et al.

1999)

No data


9.2 Return times

9.2.1 Table 16 details the return times for Natterer’s bat reported in empirical data (Swift

1997).

Table 16. Return times for Natterer’s bat Myotis nattereri reported in empirical

data (Swift 1997, Siemers et al. 1999).

SEX SEASON

AVERAGE

RETURN TIME

(minutes

before/after

sunrise)

± RANGE*

(minutes

before/after

sunrise)

INDIVIDUAL

OBSERVATIONS


sunrise)

F Pregnancy –

May/June No data No data

135 minutes before† (2 hours and 15 minutes)

(Swift 1997)

F Lactation –

July No data No data

110 minutes before† (1 hour and 50 minutes)

(Swift 1997)

F Weaning –

August

No data No data 60 minutes before†

(1 hour)

(Swift 1997)

40 minutes before

(Siemers et al.

1999)

50-30 minutes

before (Siemers

et al. 1999)

No data




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9.3 References



Siemers B, Kaipf I & Schnitzler H 1999. The use of day roosts and foraging grounds by

Natterer’s bats (Myotis nattereri Kuhl, 1818) from a colony in southern Germany.

International Journal of Mammalian Biology 64: 241-245

Swift S 1997. Roosting and foraging behaviour of Natterer's bats (Myotis nattereri) close to

the northern border of their distribution. J. Zool., Lond. 242: 375-384

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10. LEISLER’S BAT Nyctalus leisleri

10.1 Emergence times

10.1.1 Table 17 details the emergence times for Leisler’s bat Nyctalus leisleri reported in

empirical data (Jones & Rydell 1994, Shiel & Fairley 1999, Waters et al. 1999).

Table 17. Emergence times for Leisler’s bat Nyctalus leisleri reported in

empirical data (Jones & Rydell 1994, Shiel & Fairley 1999, Waters et al. 1999).

SEX

AVERAGE

EMERGENCE TIME


sunset)

± RANGE*


sunset)

ACTUAL RANGE


sunset)

F

June – Median: 18 minutes

after


No data

Earliest starting 3

minutes after



(Waters et al. 1999)


after (Waters et al. 1999) No data


(Shiel & Fairley 1999) No data

23 minutes 38 seconds

before up to 77 minutes

40 seconds after

(Shiel & Fairley 1999)


10.2 Return times

10.2.1 Table 18 details the return times for Leisler’s bat reported in empirical data (Waters

et al. 1999).

Table 18. Return times for Leisler’s bat Nyctalus leisleri reported in empirical

data (Waters et al. 1999).

SEX SEASON

AVERAGE RETURN

TIME


sunrise)

± RANGE*

(minutes

before/after

sunrise)

ACTUAL

RANGE

(minutes

before/after

sunrise)

F Pregnancy –

May/June

Mean: 320 minutes before† (5 hours and 20 minutes)

(Waters et al. 1999) No data No data

F Lactation – July 12 minutes before‡

(Waters et al. 1999)

24-0 minutes before

(Waters et al. 1999) No data

F Weaning –

August

Mean: 320 minutes before† (5 hours and 20 minutes)

(Waters et al. 1999) No data No data


† assuming one foraging bout.

‡ assuming two foraging bouts.


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10.3 References



Shiel C & Fairley 1999. Evening emergence of two nursery colonies of Leisler’s bat

(Nyctalus leisleri) in Ireland. J. Zool., Lond 247: 439-447

Waters D, Jones G & Furlong M 1999. Foraging ecology of Leisler’s bat (Nyctalus leisleri)

at two sites in southern Britain. J. Zool., Lond 249: 173-180

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11. NOCTULE Nyctalus noctula


11.1.1 Table 19 details the emergence times for the noctule Nyctalus noctula reported in

empirical data (Kronwitter 1988, Jones 1995, Kaňuch 2007).

Table 19. Emergence times for the noctule Nyctalus noctula reported in

empirical data (Kronwitter 1988, Jones 1995, Kaňuch 2007).

SEX

AVERAGE EMERGENCE

TIME


± RANGE*


sunset)

ACTUAL RANGE


sunset)

F

May & August – Median: 7.6

minutes after (Jones 1995)

May & August – SD:

0.1 of a minute before

to 15.3 minutes after

(Jones 1995)

May: c. 12 minutes

before to 46 minutes

after† (Jones 1995)

July – Median: 0.2 of a

minute before (Jones 1995)

July – SD: 4.18

minutes before to 4.58

minutes after (Jones

1995)

No data

Median


(Kronwitter 1988)

SD: 16 minutes before

to 31 minutes after

(Kronwitter 1988)

No data


(Kaňuch 2007) No data

7-26 minutes after

(Kaňuch 2007)

M Mean: 7 minutes after

(Kronwitter 1988)

SD: 16 minutes before

to 31 minutes after

(Kronwitter 1988)

No data


† Inferred from data presented in figure.

11.2 Return times

11.2.1 Table 20 details the return times for the noctule reported in empirical data (Kaňuch

2007).

Table 20. Return times for the noctule Nyctalus noctula reported in empirical

data (Kaňuch 2007).

SEX SEASON

AVERAGE

RETURN TIME

(minutes

before/after

sunrise)

± RANGE*

(minutes

before/after

sunrise)

ACTUAL RANGE


sunrise)

F Pregnancy –

May/June No data No data Onset of civil twilight

up to 3 minutes before

sunrise (Kaňuch 2007) F Lactation – July No data No data




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11.3 References

Jones G 1995. Flight performance, echolocation and foraging behaviour in noctule bats

Nyctalus noctula. J. Zool., Lond. 237: 303-312

Kaňuch P 2007. Evening and morning activity schedules of the noctule bat (Nyctalus

noctula) in Western Carpathians. Mammalia: 126-130

Kronwitter F 1988. Population structure, habitat use and activity patterns of the noctule bat,

Nyctalus noctula Schreb., 1774 (Chiroptera: Vespertilionidae) revealed by radio-tracking.

Myotis 26: 23-85



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12. NATHUSIUS’ PIPISTRELLE Pipistrellus nathusii


12.1.1 Table 21 details the emergence times for Nathusius’ pipistrelle Pipistrellus nathusii

reported in empirical data (Gelhaus & Zahn 2010).

Table 21. Emergence times for Nathusius’ pipistrelle Pipistrellus nathusii

reported in empirical data (Gelhaus & Zahn 2010).

SEX

AVERAGE

EMERGENCE TIME


sunset)

± RANGE*


sunset)

ACTUAL RANGE


sunset)


(Gelhaus & Zahn 2010) No data

11-50 minutes after

(Gelhaus & Zahn 2010)


12.2 Return times

12.2.1 Table 22 details the return times for Nathusius’ pipistrelle reported in empirical data

(Gelhaus & Zahn 2010).

Table 22. Return times for Nathusius’ pipistrelle Pipistrellus nathusii reported

in empirical data (Gelhaus & Zahn 2010).

SEX SEASON

AVERAGE

RETURN TIME

(minutes

before/after

sunrise)

± RANGE*

(minutes

before/after

sunrise)

ACTUAL

RANGE

(minutes

before/after

sunrise)

F Pregnancy – May/June No data No data 60-sunrise (1 hour up to sunrise)

(Gelhaus & Zahn

2010)




12.3 References

Gelhaus M & Zahn A 2010. Roosting ecology, phenology and foraging habitats of a nursery

colony of Pipistrellus nathusii in the southwestern part of its reproduction range.

Vespertilio 13-14: 93-102

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13. COMMON PIPISTRELLE Pipistrellus pipistrellus


13.1.1 Table 23 details the emergence times for the common pipistrelle Pipistrellus

pipistrellus reported in empirical data (Davidson-Watts & Jones 2006).

Table 23. Emergence times for the common pipistrelle Pipistrellus pipistrellus

reported in empirical data (Davidson-Watts & Jones 2006).

SEX

AVERAGE

EMERGENCE TIME


sunset)

± RANGE*


sunset)

ACTUAL RANGE


sunset)

F


(Davidson-Watts & Jones

2006)

SD: 6.9-42.7 minutes after


2006)

No data


13.2 Return times

13.2.1 Table 24 details the return times for the common pipistrelle reported in empirical

data (Davidson-Watts & Jones 2006).

Table 24. Return times for the common pipistrelle Pipistrellus pipistrellus


SEX SEASON

AVERAGE

RETURN TIME

(minutes

before/after

sunrise)

± RANGE*


sunrise)

ACTUAL

RANGE

(minutes

before/after

sunrise)

F Pregnancy –


before (2 hours and 57.8 minutes

before) (Davidson-Watts &

Jones 2006)


before (4 hours and 49.5 minutes up to 1

hour and 6.1 minutes before) (Davidson-Watts & Jones

2006)

No data

F Lactation –

July No data

F Weaning –

August No data


13.3 References

Davidson-Watts I & Jones G 2006. Differences in foraging behavior between Pipistrellus

pipistrellus (Schreber, 1774) and Pipistrellus pygmaeus (Leach, 1825). J. Zool 268: 55-

62

See also: Rydell J, Entwistle A & Racey P 1996. Timing of foraging flights of three species


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of bats in relation to insect activity and predation risk. Oikos 76: 243-252

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14. SOPRANO PIPISTRELLE Pipistrellus pygmaeus


14.1.1 Table 25 details the emergence times for the soprano pipistrelle Pipistrellus

pygmaeus reported in empirical data (Davidson-Watts & Jones 2006, Swift 1980).

Table 25. Emergence times for the soprano pipistrelle Pipistrellus pygmaeus

reported in empirical data (Davidson-Watts & Jones 2006, Swift 1980).

SEX

AVERAGE

EMERGENCE TIME


sunset)

± RANGE*


sunset)

ACTUAL RANGE


sunset)

F

May/June – Mean: 35

minutes after†

(Swift 1980)

July – Mean: 27 minutes

after† (Swift 1980)

August – Mean: 29

minutes after†

(Swift 1980)

May/June – SD: 23- 47

minutes after†

(Swift 1980)

July – SD: 21-33 minutes


August – SD: 25-34

minutes after†

(Swift 1980)

May/June: 15-56 minutes


July: 19-35 minutes after†

(Swift 1980)

August: 24-35 minutes




2006)



2006)

No data



14.2 Return times

14.2.1 Table 26 on the following page details the return times for the soprano pipistrelle


14.3 References

Davidson-Watts I & Jones G 2006. Differences in foraging behavior between Pipistrellus

pipistrellus (Schreber, 1774) and Pipistrellus pygmaeus (Leach, 1825). J. Zool 268: 55-

62

Swift S 1980. Activity patterns of Pipistrelle bat (Pipistrellus pipistrellus) in north-east

Scotland. J. Zool 190: 285-295


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Table 26. Return times for the soprano pipistrelle Pipistrellus pygmaeus


SEX SEASON

AVERAGE

RETURN TIME

(minutes

before/after sunrise)

± RANGE*


sunrise)

ACTUAL

RANGE

(minutes

before/after

sunrise)

F Pregnancy –


before (4 hours and 28 minutes)

(Davidson-Watts &

Jones 2006)

378-159.6 minutes

before (6 hours and 18 minutes up to 2 hours and 39.6 minutes before)

(Davidson-Watts &

Jones 2006)

No data

F Lactation – July No data

F Weaning –

August No data


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15. GREY LONG-EARED BAT Plecotus austriacus


15.1.1 Table 27 details the emergence times for the grey long-eared bat Plecotus austriacus

reported in empirical data (Razgour et al. 2011).

Table 27. Emergence times for the grey long-eared bat Plecotus austriacus

reported in empirical data (Razgour et al. 2011).

SEX

AVERAGE

EMERGENCE TIME


sunset)

± RANGE*


sunset)

ACTUAL RANGE


sunset)


(Razgour et al. 2011)


(Razgour et al. 2011) No data

M Mean: 39 minutes after†



(Razgour et al. 2011) No data


15.2 Return times

15.2.1 Table 28 details the return times for the grey long-eared bat reported in empirical

data (Scheunert et al. 2010, Razgour et al. 2011).

Table 28. Return times for the grey long-eared bat Plecotus austriacus reported

in empirical data (Scheunert et al. 2010, Razgour et al. 2011).

SEX SEASON AVERAGE RETURN TIME

(minutes before/after sunrise)

± RANGE*

(minutes

before/after

sunrise)

ACTUAL

RANGE

(minutes

before/after

sunrise)

F Pregnancy –

May/June

Mean: 140 minutes before† (2 hours and 20 minutes before)

(Scheunert et al. 2010) No data No data

F Lactation –

July

Mean: 0 minutes before† (at dawn)

(Razgour et al. 2011) No data No data

F Weaning –

August

Mean: 55 minutes before†

(Razgour et al. 2011) No data No data

M All

July – Mean: 18 minutes before†


August – Mean: – 114 minutes

before† (1 hour and 54 minutes before)


No data No data


† Inferred from data presented.


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15.3 References

Razgour O, Hanmer J & Jones G 2011. Using multi-scale modelling to predict habitat

suitability for species of conservation concern: The grey long-eared bat as a case study.

Biological Conservation 144: 2922-2930

Scheunert A, Zahn A & Kiefer A 2010. Phenology and roosting habits of the Central

European grey long-eared bat Plecotus austriacus (Fischer 1829). Eur. J. Wildl. Res. 56:

435-442

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16. BROWN LONG-EARED BAT Plecotus auritus


16.1.1 Table 29 details the emergence times for the brown long-eared bat Plecotus auritus

reported in empirical data (Jones & Rydell 1994, Entwistle et al. 1996, Rydell et al.

1996).

Table 29. Emergence times for the brown long-eared bat Plecotus auritus

reported in empirical data (Jones & Rydell 1994, Entwistle et al. 1996, Rydell et

al. 1996).

SEX

AVERAGE EMERGENCE

TIME


± RANGE*


sunset)

ACTUAL RANGE

(minutes

before/after sunset)

F




minutes before

(Jones & Rydell

1994)

Mean: 61.7 minutes after (c. 1

minute before the end of civil

twilight)

(Entwistle et al. 1996)

SD: 57.4-66 minutes

after


No data


(Rydell et al. 1996)


(Rydell et al. 1996) No data


16.2 Return times

16.2.1 Table 30 details the return times for the brown long-eared bat reported in empirical

data (Entwistle et al. 1996).

Table 30. Return times for the brown long-eared bat Plecotus auritus reported

in empirical data (Entwistle et al. 1996).

SEX SEASON

AVERAGE RETURN

TIME


sunrise)

± RANGE*

(minutes

before/after

sunrise)

ACTUAL

RANGE

(minutes

before/after

sunrise)

F Pregnancy –


before (1 hour and 22 minutes before

and typically c. 8 minutes before

the beginning of civil twilight)


SD: 91.9-73.5

minutes before (1 hour and 31 minutes up

to 1 hour and 13 minutes)

(Entwistle et al.

1996)

No data

F Lactation – July No data

F Weaning –

August No data



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16.3 References

Entwistle A, Racey P & Speakman J 1996. Habitat exploitation by a gleaning bat, Plecotus

auritus. Phil. Trans. R. Soc. Lond. B 351: 921-931



Rydell J, Entwistle A & Racey P 1996. Timing of foraging flights of three species of bats in

relation to insect activity and predation risk. Oikos 76: 243-252

See also: Fuhrmann M & Seitz A 1992. Nocturnal activity of the brown long-eared bat

(Plecotus auritus L., 1758); data from radio tracking in the Lenneberg forest near Mainz

(Germany) In: Preide I & Swift S (eds.) Wildlife telemetry. Ellis Horwood, Chichester

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17. GREATER HORSESHOE-BAT Rhinolophus ferrumequinum


17.1.1 Table 31 details the emergence times for the greater horseshoe-bat Rhinolophus

ferrumequinum reported in empirical data (Jones & Rydell 1994, Robinson et al.

2000).

Table 31. Emergence times for the greater horseshoe-bat Rhinolophus

ferrumequinum reported in empirical data (Jones & Rydell 1994, Robinson et

al. 2000).

SEX

AVERAGE

EMERGENCE TIME


sunset)

± RANGE*


sunset)

ACTUAL RANGE


sunset)

F

May/June – Mean: 28

minutes after†

(Robinson et al. 2000)

May/June – SD: 11-45

minutes after†


May/June: 4 minutes

before – 64 minutes

after† (4 minutes before up to 1 hour and

4 minutes after) (Robinson et al. 2000)

June – Median: 25

minutes after


No data


minutes after


July – Mean: 26 minutes

after†


July – SD: 20-32 minutes

after†


July: 16-36 minutes after†



† calculated using raw data provided.

17.2 Return times

17.2.1 Table 32 on the following page details the return times for the greater horseshoe-bat

reported in empirical data (Robinson et al. 2000).

17.3 References



Robinson M, Webber M & Stebbings R 2000. Dispersal and foraging behaviour of greater

horseshoe bats, Brixham, Devon. No. 344 - English Nature Research Reports. English

Nature, Peterborough

See also: Duvergé P, Jones G, Rydell J & Ransome R 2000. Functional significance of

emergence timing in bats. Ecography 23: 32-40


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Table 32. Return times for the greater horseshoe-bat Rhinolophus

ferrumequinum reported in empirical data (Robinson et al. 2000).

SEX SEASON

AVERAGE

RETURN TIME

(minutes

before/after

sunrise)

± RANGE*

(minutes

before/after

sunrise)

ACTUAL

RANGE

(minutes

before/after

sunrise)

F Pregnancy –

May/June

Mean: 34 minutes

before†

(Robinson et al.

2000)

SD: 46-22 minutes†

(Robinson et al.

2000)

49-17 minutes

before†

(Robinson et al.

2000)

F Lactation – July

Mean: 42 minutes

before†

(Robinson et al.

2000)

SD: 56-28 minutes†

(Robinson et al.

2000)

77-27 minutes

before† (1 hour and 17 minutes up

to 27 minutes before)

(Robinson et al.

2000)

F Weaning –

August No data No data No data


† calculated using raw data provided.

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18. LESSER HORSESHOE-BAT Rhinolophus hipposideros


18.1.1 Table 33 details the emergence times for the lesser horseshoe-bat Rhinolophus

hipposideros reported in empirical data (Jones & Rydell 1994, Knight 2006).

Table 33. Emergence times for the lesser horseshoe-bat Rhinolophus

hipposideros reported in empirical data (Jones & Rydell 1994, Knight 2006).

SEX

AVERAGE EMERGENCE

TIME


± RANGE*


ACTUAL

RANGE

(minutes

before/after

sunset)

F

May/June – Mean: 33 minutes

after (Knight 2006)

May/June – SD: 30-36 minutes

after (Knight 2006) No data



Earliest

starting 19

minutes after

(Jones &

Rydell 1994)

July – Mean: 21 minutes after

(Knight 2006)

July – SD: 14-28 minutes after

(Knight 2006) No data

August – Mean: 37 minutes

after (Knight 2006)

August – SD: 16-58 minutes

after (Knight 2006) No data


18.2 Return times

18.2.1 Table 34 details the return times for the lesser horseshoe-bat reported in empirical

data (Knight 2006).

Table 34. Return times for the lesser horseshoe-bat Rhinolophus hipposideros

reported in empirical data (Knight 2006).

SEX SEASON

AVERAGE

RETURN TIME


sunrise)

± RANGE*

(minutes

before/after sunrise)

ACTUAL

RANGE

(minutes

before/after

sunrise)

F Pregnancy –

May/June

Mean: 36 minutes

before (Knight 2006)

SD: 39-33 minutes

before (Knight 2006) No data

F Lactation – July Mean: 31 minutes


SD: 41-24 minutes


F Weaning –

August

Mean: 26 minutes


SD: 33-19 minutes




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18.3 References



Knight T 2006. The use of landscape features and habitats by the lesser horseshoe bat

(Rhinolophus hipposideros). Dissertation submitted to the University of Bristol in

accordance with the requirements of the degree of PhD in the Faculty of Science

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