1

American Burying Beetle (Nicrophorus americanus) Survey on Nantucket

Island, Massachusetts, 2015

Andrew McKenna-Foster1, Lou Perrotti2, Brendon Donohue1 Elisabeth Sorrows1

1. Maria Mitchell Association, 4 Vestal Street, Nantucket MA 02554

2. Roger Williams Park Zoo, 1000 Elmwood Avenue, Providence, RI 02907

Report submitted to the U.S. Fish and Wildlife Service on October 22, 2015

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Contents ABSTRACT ...................................................................................................................................................... 3

INTRODUCTION ............................................................................................................................................. 3

METHODS ...................................................................................................................................................... 4

Early Summer Trapping ............................................................................................................................. 4

Late Summer Trapping .............................................................................................................................. 4

Temperature Data Collection .................................................................................................................... 4

2015 Trap Site Locations ........................................................................................................................... 4

Measurements and Markings ................................................................................................................... 6

Provisioning ............................................................................................................................................... 6

Abundance of other Carrion Beetle Species ............................................................................................. 6

RESULTS ........................................................................................................................................................ 6

Early Summer Trapping ............................................................................................................................. 6

Recapture of 2014 Teneral Beetles ........................................................................................................... 9

Provisioned Broods ................................................................................................................................... 9

Mass and Pronotal Width ....................................................................................................................... 10

Ground Temperature Data ..................................................................................................................... 11

Late Summer Trapping ............................................................................................................................ 13

DISCUSSION ................................................................................................................................................. 13

ACKNOWLEDGMENTS ................................................................................................................................. 14

LITERATURE CITED ...................................................................................................................................... 15

APPENDIX A ................................................................................................................................................. 17

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ABSTRACT

The American burying beetle (Nicrophorus americanus, ABB) is a federally listed endangered beetle.

Once common throughout the eastern half of the United States, N. americanus now survives in a small

number of isolated or undisturbed habitats in eight states. Since 1993, multiple organizations have

collaborated to reintroduce the ABB to Nantucket Island, Massachusetts. The project is now in a

monitoring phase and, in 2015, we trapped and marked 32 adult N. americanus as part of this effort. We

provisioned 14 pairs and four individual ABB’s each with a piece of carrion on conservation land for a

total of 18 broods. The provisioned broods had a 86% success rate, producing an estimated 357 larvae.

During our late summer trapping we captured 80 teneral beetles in four trap intervals. The first late

summer trap interval was specifically timed to look for teneral beetles from wild broods (not

provisioned by us) and we did not capture any beetles. Our results indicate that the minimal

provisioning effort is possibly the only thing maintaining the ABB population on Nantucket. We make

recommendations for changes to the monitoring protocol to be more efficient in terms of person hours

and to allow the collection of information that is not possible with baited pitfall traps.

INTRODUCTION

The American burying beetle (ABB, Nicrophorus americanus Oliver (1790)) is the largest of North

America’s carrion beetles (Family Silphidae). Its historical range covers 35 states in the eastern

temperate areas of North America (Raithel 1991). However, the species is extirpated across much of its

native range in the eastern U.S. Today, there are only scattered populations in Arkansas, Kansas,

Massachusetts, Missouri, Nebraska, Ohio, Oklahoma, Rhode Island, South Dakota, and Texas (TESS). The

only naturally occurring population east of the Mississippi River is located on Block Island, RI, and is

comprised of an estimated 1,000-2,000 individuals (USFWS 2008).

The last wild N. americanus on Nantucket Island, Massachusetts was recorded in 1926 (Johnson 1930).

Nantucket is approximately 42 km south of Cape Cod and is one of the more remote islands off the

Northeast coast. Extensive trapping surveys in 1992 and 1993 were unsuccessful in catching any N.

americanus on Nantucket (Amaral 1993, Northrup 1994). In the summer of 1994, 22 pairs of beetles

and four individuals were provisioned on the island in the Massachusetts Audubon Society’s Sesachacha

Heathland Wildlife Sanctuary, hereafter referred to as site 6(USFWS Memorandum 1998). Releases

continued until 2006; to date, a total of 2,923 beetles have been released on Nantucket. Annual reports

from Nantucket have been submitted to USFWS since 2004 (Mckenna-Foster et al. 2004, 2005, 2006a,

2006b, 2007, 2009, 2010; Shuster et al. 2008, LoPresti et al. 2011, Morse et al. 2012, Hawkins et al.

2013, Mckenna-Foster et al. 2014). Between 2007-2010, every beetle caught was provisioned with a

quail carcass and as many as possible with mates. In 2011, provisioning was reduced to 25 broods to

assess the viability of a self-sustaining population. We have continued this level of provisioning since

then.

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METHODS

Early Summer Trapping

We set traps on the night of 15 June 2015, and kept traps open continuously until 30 June 2015 with the

exception of Site 3, where traps were removed on 28 June (Table 1). At all sites we placed five traps

approximately 20 meters apart in a line. Trapping followed the procedure described by Kozol (1991).

Each trap consisted of a 946 mL mason jar buried with the top flush with ground level and covered with

a square piece of hardware cloth (with a 3cm x 3cm hole cut from the middle) and a two-sided “tent” of

thick aluminum to keep out the rain and sun. Inside each jar was a plastic container with a screw-on

screen lid filled to three quarters volume with small pieces of rotted chicken. We prepared the chicken

beforehand in plastic containers left it at room temperature for 7-8 days. Each trap also contained a wet

~3 cm3 sponge to provide moisture. We checked traps every morning between 0500 and 1000 hours to

ensure that all ABB were removed before traps became lethally warm.

While in captivity beetles were stored in a dark cooler chilled with ice packs to an average of 18.3 + 2.3

C. We housed beetles in groups of one to four individuals, separated by sex, in plastic containers with a

moist paper towel and we provided mealworms ad libitum for food.

Late Summer Trapping

During the late summer we trapped for three intervals: nights of 12 August – 14 August (three nights),

21 August – 29 August (nine nights), 31 August – 2 September (three nights) and 8 September – 9

September (two nights). During the first interval we trapped at all sites (except site, five traps at each

site) and during the next three intervals we only trapped at the provisioning site with 7 to 20 traps.

Trapping protocol was identical to the early summer protocol.

Temperature Data Collection

We obtained air temperature, humidity, and wind speed data from the Weather Underground

(www.wunderground.com). We used iButtons at trap sites 1, 4, 6, 7, and 14 to collect ground-level

temperatures from 16 June to 30 June at 30 minute intervals. We calculated mean nightly temperature

from approximately 2000 (+ 10 min) to 0730 (+ 10 min) from these data.

2015 Trap Site Locations

We set traps for ABB at 12 sites on the eastern side of the island (Fig. 1). We used the same grid system

set up in 2013 and used in 2014 to assess the distribution of the population across the eastern side of

the island. Trap sites 2 and 13 were abandoned after 2013 and are not listed. The average distance

between a site and its four closest neighbors is 1.96 ± 0.571km. Assuming each trap line has an effective

range of 0.8 Km (USFWS 2013), the grid system covers 22.7 Km2.

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1: Located on Nantucket Land Bank (NLB) property, this site is along a trail bordered by pine, oak, and

shrubs. The coordinates for the start of the trap line are 41.2851N 70.0523W

3: Previously Squam Farm, This site is on NCF property three miles north of the Eastern Moors site.

Located on the edge of a tupelo (Nyssa sylvatica) and scrub oak (Quercus ilicifolia) forest with

interspersed wetlands, the trap site is in a field just west of the parking lot. The coordinates for the start

of the trap line are 41.3072N 70.0009W

4: On NLB property, this site runs along a trail surrounded by pine, oak, and shrubs. It is near a wetland

and lies along the boundary of outwash plain and ice contact deposits. The coordinates for the start of

the trap line are 41.2898N 69.9926W

5: Located on NCF property, this site runs along an old road and is surrounded by scrub oak. The

coordinates for the start of the trap line are 41.2836N 69.9780W

6: Previously Eastern Moors, this is the location of most previous releases on the eastern side of the

island and was referred to in previous reports as the Middle Moors or Release site. Located in the

Massachusetts Audubon Society’s Sesachacha Heathland Wildlife Sanctuary, it is a large area composed

of sandplain grasslands and heathlands. The dominant plant species are bearberry (Arctostaphylos uva-

ursi), lowbush blueberry (Vaccinium angustifolium), scrub oak, and sweetfern (Comptonia peregrina).

Provisioning sites are clustered west of a dirt road in the center of a large area of low grass and

bearberry. The trap line runs south roughly parallel with the dirt road. The coordinates for the start of

the trap line are 41.2760N 69.9891W

7: Near sites formerly referred to as Gibbs Pond, this trap line runs along Barnard Valley Road and is

surrounded by scrub oak. The coordinates for the start of the trap line are 41.2747N 70.0124 W

8: On NCF property this site is relatively high in elevation and runs along the side of a road surrounded

by heath shrubs and scrub oak. The coordinates for the start of the trap line are 41.2843N 70.0215W

9: Near Altar Rock, this site is on NCF property and is along the road surrounded by open heathland. The

coordinates for the start of the trap line are 41.2798N 70.0314W

10: : This site was formerly referred to as Pout Ponds and is just east of the road next to the southern-

most pond in the Pout Pond area on NCF property. It is on the edge of scrub oak, bearberry and pitch

pine (P. rigida) habitat. The coordinates for the start of the trap line are 41.2753N 70.2753W

11: Previously ‘Sconset Dump, This site is on NCF property and runs along the northern edge of the

Sconset Dump just south of Milestone Road. The surrounding habitat is mostly scrub oak with

occasional open grassy areas. The coordinates for the start of the trap line are 41.2621N 69.9884W

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12: Previously referred to as Serengeti, this site is on NCF property. It is located 300 meters north of

milestone four on Milestone Road in an area that is mowed regularly and is predominately scrub oak

and bearberry. The coordinates for the start of the trap line are 41.2655N 70.0320W

14: This site is on NCF property near the airport. The traps are surrounded by pine and broom

crowberry (Corema conradii). The coordinates for the start of the trap line are 41.2612N 70.0478W

Provisioning Site: Centrally located among the trap sites, this area is predominated by lowbush

blueberry (Vaccinium angustifolium) and small scrub oak (Quercus ilicifolia). This is the same site used in

2014.

Measurements and Markings

During the early-summer trapping, we sexed, massed, and measured pronotal width on all captured

beetles, notched them according to trap site (Appendix A), and glued a bee tag (www.beeworks.com)

with a unique number/color combination to each individual.

For newly emerged beetles trapped in the late summer, we sexed, measured pronotal width, notched

each individual and glued on a bee tag while in the field and released the beetles on-site.

Provisioning

We provisioned 14 pairs of beetles and four individual beetles (two male, two female) each with piece of

carrion buried approximately 20 centimeters underground. Quail, rats, and rabbit were used as carrion.

We kept all captured beetles in captivity until 25 June 2015, at which time we paired and provisioned

seven pairs and two individual males. We provisioned seven more pairs and two individual females on 2

July 2015. Carrion weight averaged 151.2 ± 17.18g SD (range 126.2-177.8 g). To protect the carrion and

beetles from scavengers, we stapled hardware cloth to the ground over the filled-in hole. Provisioned

broods were parented by paired male and female ABB captured at the same site when possible, and

adjacent sites in some cases. All broods were provisioned at the central provisioning site (Fig. 1).

Abundance of other Carrion Beetle Species

We recorded the abundance of other carrion beetle species in traps during the early-summer trapping

interval and can make that data available upon request.

RESULTS

Early Summer Trapping

We trapped 32 beetles between 16 June 2015 and 30 June, 2015 at 12 traps sites (Table 1). Sixteen of

these individuals were female and sixteen were male. This ratio fits exactly with the expected 1:1 male

to female ratio observed in previous years. We caught the greatest number of beetles at site 9 (Fig. 1).

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Table 1: Summary of early-summer trapping. Lost nights are nights when bait was stolen by scavengers

and no beetles were captured, or nights when a trap was not set. Values represent new beetle captures

and not recaptured beetles.

Trap Site

Trapping Dates

# Traps Lost Trap Nights

Total Trap Nights

Total Females

Total Males

Total Beetles

New Beetles/Trap Night

1 16 June- 30 June 5 0 75 1 0 1 0.013 3 16 June- 28 June 5 0 65 1 0 1 0.015 4 16 June- 30 June 5 0 75 0 0 0 0 5 16 June- 30 June 5 0 75 1 0 1 0.013 6 16 June- 30 June 5 0 75 4 4 8 0.107 7 16 June- 30 June 5 4 71 0 0 0 0 8 16 June- 30 June 5 0 75 2 0 2 0.027 9 16 June- 30 June 5 2 73 5 8 13 0.178 10 16 June- 30 June 5 2 73 0 2 2 0.027 11 16 June- 30 June 5 0 75 0 0 0 0 12 16 June- 30 June 5 2 73 1 2 3 0.041 14 16 June- 30 June 5 0 75 1 0 1 0.013 Overall 10 880 16 16 32 0.036

Figure 1: Trap rates for trap sites. The size of the circle indicates its success. The transparent circles

show the assumed 0.8km effective trap site radius.

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The total number of beetles captured in the summer of 2015 was the lowest since 2004 (Table 2). We

accumulated beetles at a slightly lower rate than 2014 but were on par with 2013 (Fig. 2).

Table 2: Total early-summer trap nights and ABB captures with trap rates since 2004.

Year Total Nights Total Wild ABBs Number ABBs/Trap

Night

2004 360 33 0.092 2005 480 38 0.079 2006 640 50 0.078 2007 1,022 97 0.095 2008 1,348 112 0.083 2009 1,791 150 0.083 2010 1,663 191 0.115 2011 1,011 212 0.210 2012 1,237 115 0.093 2013 1,386 52 0.038 2014 778 36 0.046 2015 880 32 0.036

Figure 2: Accumulation curves of total ABB captures as a function of trap nights for years 2007 to 2015.

0

50

100

150

200

250

0 500 1000 1500 2000 Cu

mu

lati

ve N

um

be

r o

f N

ew B

eet

les

Trap Nights

2007

2008

2009

2010

2011

2012

2013

2014

2015

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Recapture of 2014 Teneral Beetles

We captured three beetles marked during the late summer trapping season of 2014 for an overwinter

survival rate of 8.5% (Table 3).

Table 3: The number of beetles marked one year and recaptured the following year.

Year of recapture # Marked Previous year # Recaptured % Recaptured

2007 82 12 14.6

2008 87 9 10.3

2009 112 21 18.8

2010 372 51 13.8

2011 434 63 14.5

2012 139 31 22.3

2013 77 15 19.5 2014 2015

42 35

6 3

14.3 8.5

This year had the lowest recapture rate since 2007. Of the three recaptures from 2014, two (67%) were

readable and one (33%) was missing its tag. These beetles moved an average of 0.95 ± 0.83 Km (SD)

between fall and spring.

Provisioned Broods

We excavated seven broods (39%) between 11-13 days after provisioning. The broods provisioned on 25

July were excavated 11 days after provisioning, and the broods provisioned on 2 July were excavated 13

days after provisioning. Mean brood size for all seven broods exhumed in 2015 was 20 ± 9.3 SD larvae

(range 0-27). Mean successful brood size was 23.3 ± 3.3 SD. We counted 85 3rd instar larvae, 27 2nd

instar larvae, and 28 1st instar larvae in the broods we exhumed.

Eighty-six percent of the broods were successful and we estimate that they produced a total of 357

larvae (Table 4). We estimated total ABB larvae production by multiplying the number of un-checked

broods by mean successful brood size, and then multiplying this figure by the brood success rate

observed in the seven broods that we exhumed.

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Table 4: Descriptive data for brood provisioned from 2009-2015. Means are reported (± SD).

Year # Broods

Provisioned

# Broods

Exhumed

Brood

Success

Rate

Mean

Successful

Brood Size

Est. Larvae Produced

Counted

Larvae

Est. Total

Production

2009 104 37 70% 17 (±3) 750 416 1,166 2010 153 58 43% 18 (±1) 694 445 1,139 2011 25 8 63% 16 (±2) 171 82 253

2012 25 8 63% 21 (±3) 225 105 330

2013 25 10 30% 18 (±5) 79 53 132 2014 2015

22 18

8 7

50% 86%

12 (±11) 23 (±3)

84 217

46 140

130 357

Of the three broods we checked that were provisioned with a single beetle, two were successful. Both

adults were captured in one of the successful broods and the individually provisioned male had

attracted a previously uncaptured female.

Mass and Pronotal Width

Beetle size and mass were similar among sexes, adults and tenerals, and previous years (Tables 5 and 6,

Fig. 3).

Table 5: Mean pronotal width (±SE) for males and female beetles for the 2015 early (adult) and late

(teneral) summer trapping intervals.

Trapping Period Summer (n =32) Late Summer (n = 80)

Sex Male (n = 16) Female (n = 16) Male (n = 45) Female (n = 35)

Average (mm) 11.7 ± 0.2 11.6 ± 0.2 11.6 ± 0.2 11.5 ± 0.2

Range (mm) 10.0-13.0 10.0-12.6 9.2-13.0 8.7-12.8

Overall Average (mm) 11.7 ± 0.2 11.5 ± 0.1

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Table 6: Mean mass (±SE) for males and female beetles for the 2015 early summer trapping. Teneral

beetles in the late summer were not weighed.

Trapping Period Summer (n =32)

Sex Male (n = 16) Female (n = 16)

Average (g) 1.8 ± 0.1 1.6 ± 0.1

Range (g) 1.2-2.7 1.0-2.3

Total Average (g) 1.7 ± 0.1

Figure 3. Pronotal widths for beetles during the reproductive season.

Ground Temperature Data

Mean nightly ground temperature between 16 June and 30 June was relatively stable after an initial cold

snap on 18 June (Fig. 4). The temperature at site 4 varied outside the iButton error range (+1 degree)

10

10.5

11

11.5

12

12.5

2007 2008 2009 2010 2011 2012 2013 2014 2015

Su

mm

er P

ron

ota

l Wid

th (m

m)

Year

Males

Females

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when compared to other sites suggesting this site was colder than the other sites. The mean nightly

ground temperature from all sites over the course of the summer was 15 C (+ 4 SD). There was fog on

the island 50% of the nights (7 of 14) during the trapping period. Two nights saw gusting winds.

In the late summer trapping interval across all trap sites, no iButtons were used but Weather

Underground reported that the average nightly temperature was 20 C (±3 SD). There was no rain but fog

was reported two of the three nights.

In the late summer trapping intervals at the provisioning site, no iButtons were used but Weather

Underground reported that the average nightly temperature was 22 C (±0.5 SD) and there was 0.87

inches of rain. Fog or mist was reported 36% (five of 14 )of the nights. The average temperature for

each interval was 21 C (±1 SD), 20C (±2 SD), and 22C (±0.4 SD) in chronological order.

Figure 4: Mean nightly above ground temperatures from iButtons at five trap sites.

0

5

10

15

20

25

Tem

pe

ratu

re (

C)

Date

1

4

6

7

14

13

Late Summer Trapping

In four separate trap intervals and in 401 total trap nights, we captured 80 teneral beetles (Table 7). No

beetles were captured in the first interval. This interval was timed before the scheduled emergence of

teneral beetles from our provisioned broods to detect teneral beetles from broods we did not provision

(i.e. wild broods). We recaptured teneral beetles 12 times representing nine beetles. One beetle was

captured three times and one was captured twice. Eight (10%) of the beetles showed damaged to their

elytra and wings. One beetle was found eaten in a trap- only part of its pronotum was identifiable.

Table 7: Trap nights, rates, and captures at sites during late summer trapping periods.

Date Night of:

Trap Nights

Trap Rate

New Males

New Females

Recaptures Total New Beetles

12 Aug – 14 Aug 180 0 0 0 0 0 21 Aug – 29 Aug 180 0.14 15 10 4 25 31 Aug – 2 Sept 27 0.89 13 11 7 24 8 Sept – 9 Sept 14 2.2 17 14 14 31 Overall 401 0.20 35 35 25 80

DISCUSSION

We captured more beetles than expected in 2015 and had the highest provisioning success rate

recorded for this project in more than ten years. Based on a decreasing population trend since 2011, we

expected to catch about half the number of beetles seen in 2014. However, it appears that our

provisioning effort is now maintaining a small population. As in 2014, we did not catch any teneral

beetles produced by natural pairing and reproduction so we assume a majority of beetles currently in

the population are direct descendents from provisioned broods.

Site 6 and site 9 again had the highest capture rates. Site 9 is closer than any other site to the

provisioning site and we expect it to have a high trap rate. However, there are five other sites closer to

the provisioning site than site 6 and yet it had the second highest trap rate. We assume this is due to

topographic and or vegetative characteristics that enhance trap effectiveness. This demonstrates how

important site selection may be when comparing trap rates.

This project has now collected five years of data on how a reduced provisioning regime affected the

established ABB population on Nantucket. It is clear that the monitoring and provisioning effort is

responsible for maintaining this population. Key points that we have learned:

1. Trapping over the entire local range of the ABBs through the reproductive season is effective in

catching most of the adults. This is partly due to the fact that ABBs disperse several kilometers

between their emergence as tenerals in the late summer and their reproductive season in the

early summer the following year.

2. Selecting a successful site for traps is difficult. When possible, sites with very low capture rates

should be moved to nearby areas in case vegetation or topography are inhibiting trap

effectiveness.

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3. Careful provisioning can greatly increase provisioning success. Sites selection should avoid dry

sandy soils and provisioning should occur early enough in the season to avoid the dry conditions

of midsummer.

Adhering to these three points leads to an effective monitoring effort, providing detailed information on

the status of the population. However, it is extremely labor intensive and in some ways intrusive to the

natural scavenger reproductive cycle. We have seen increased trap rates for Nicrophorus marginatus at

the most eastern trap sites and one possible explanation for this change is our intensive trapping effort

drawing that species farther east. Finding a less intrusive way to maintain the ABB population may

facilitate further study of this species on Nantucket. Questions that should be investigated include:

1. How successful are Nantucket ABB’s at finding and utilizing carrion.

2. Is there a critical population density that increases their success at finding either mates or

carrion?

3. What is the minimum human effort needed to maintain a critical population density.

These questions are difficult to answer using baited traps. Trap effectiveness is dependent on

numerous, difficult to measure factors such as bait quality, vegetation and topography (even at the trap

level), presences of pheromones from previously captured individuals, and presence of other carrion

species.

We propose transitioning the monitoring effort to limited use of baited traps augmented with tethered

quail carcasses.

Recommendations for future monitoring:

1. Continue trapping using the grid arrangement.

2. Replace a proportion of traps in each trap line with tethered quail.

3. Provision captured beetles and also dig up any buried quail carcasses to assess success.

4. Transition over several seasons to using mostly tethered quail carcasses. A monitoring effort

could take the form of:

a. Mid June: tether quail carcasses at several sites in the core of the known ABB range.

b. Three to four days later, check status of each carcass.

c. Ten to twelve days later trap for three nights using baited pitfall traps. Dig up a

proportion of any quail carcasses buried to identify Nicrophorus species.

d. Provision any ABB’s captured in trapping.

e. Trap for five days in mid August to capture teneral beetles.

In time, one large scale trapping effort in mid August for tenerals could be the most efficient way to

monitor the population in the least intrusive way.

ACKNOWLEDGMENTS We thank many organizations and persons for supporting this project: the Roger Williams Park Zoo in

Providence, RI, for providing staff, supplies, and financial support; Cynthia Maynard (USFWS) for

providing oversight and assistance; The Massachusetts Audubon Society, Nantucket Conservation

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Foundation, and Nantucket Land Bank for allowing us to trap on their properties; and the Nantucket

Cottage Hospital for supplies. Temperature recording equipment was purchased with a grant from the

Norcross Wildlife Foundation. Maria Mitchell facilities and vehicles were used in support of this project.

And many thanks to the Maria Mitchell Association interns and other volunteers who contributed to the

project.

LITERATURE CITED Amaral, M. April 7, 1993. United States Department of the Interior report to the American Burying Beetle Group – East. Re. Recovery Coordination Summary. Bedick, J.C, B.C. Ratcliffe, W.W. Hoback, and L.G. Higley. 1999. Distribution, ecology, and population dynamics of the American burying beetle [Nicrophorus americanus Oliver (Coleoptera, Silphidae)] in south-central Nebraska, USA. Journal of Insect Conservation. 3(3): 171-181. Creighton, J.C., M.V. Lomolino, and G.D. Schnell. 1993. Survey methods for the American burying beetle (Nicrophorus americanus) in Oklahoma and Arkansas. Oklahoma Biological Survey, Norman, OK. Creighton, J.C. and G.D. Schnell. 1998. Short-term movement patterns of the endangered American burying beetle Nicrophorus americanus. Biological Conservation. 86: 281-287. Hawkins, T., A. Mckenna-Foster, L. Perrotti, and K. Macsuga. 2013. American burying beetle (Nicrophorus americanus) survey on Nantucket Island, Massachusetts 2013. Submitted to USFWS, Region 5, November 2013. LoPresti, E., A. McKenna-Foster, L. Perrotti, and J. Blyth. 2011. American burying beetle (Nicrophorus americanus) survey on Nantucket Island, Massachusetts. Submitted to USFWS, Region 5, November 2011. Johnson, C. W. 1930. A List of the Insect Fauna of Nantucket, Massachusetts. Publications of the Nantucket Maria Mitchell Association, Vol. 3, No. 20. Kozol A. 1991. Appendix 2. Survey protocol for Nicrophorus americanus, the American burying beetle [pp. 1–6]. In: U.S. Fish and Wildlife Service, American Burying Beetle (Nicrophorus americanus) Recovery Plan. Newton Corner, MA. 80 pp. Mckenna-Foster, A, W.T. Maple, and R.S. Kennedy. 2004. American burying beetle (Nicrophorus americanus) survey and reintroduction on Nantucket 2004. Submitted to USFWS, Region 5, November 2004. Mckenna-Foster, A, W.T. Maple, and R.S. Kennedy. 2005. American burying beetle (Nicrophorus americanus) survey and reintroduction on Nantucket 2005. Submitted to USFWS, Region 5, November 2005. Mckenna-Foster, A, W.T. Maple, and R.S. Kennedy. 2006a. A summary of the effort to

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reintroduce the American burying beetle (Nicrophorus americanus) to Nantucket Island, MA 2006. Submitted to USFWS, Region 5, September 2006. Mckenna-Foster, A, W.T. Maple, and R.S. Kennedy. 2006b. American burying beetle (Nicrophorus americanus) survey and reintroduction on Nantucket 2006. Submitted to USFWS, Region 5, October 2006. Mckenna-Foster, A, L. Perrotti, C. Biegler, and R.S. Kennedy. 2007. American burying beetle (Nicrophorus americanus) survey and reintroduction on Nantucket Island, Massachusetts 2007. Submitted to USFWS, Region 5, November 2007. Mckenna-Foster, A., J.D. Shuster, L. Perrotti, and R.S. Kennedy. 2009. American burying beetle (Nicrophorus americanus) survey on Nantucket Island, Massachusetts 2009. Submitted to USFWS, Region 5, October 2008. Mckenna-Foster, A, L. Perrotti. and T. Hawkins. 2014. American burying beetle (Nicrophorus americanus) survey on Nantucket Island, Massachusetts 2014. Submitted to USFWS, Region 5, October 2014. Northrup, M. M. 1994. The American Burying Beetle (Nicrophorus americanus): A field

Investigation of its status on Coskata-Coatue Wildlife Refuge on Nantucket Island, MA. Senior Thesis.

Bard College Environmental Studies Program. Annandale-on-Hudson, New York.

Raithel, C. 1991. American Burying Beetle Recovery Plan. USFWS Region 5. Coordinated through New England Office USFWS, by Michael Amaral. Raithel, C., Ginsberg, H.S., Prospero, M.L. 2006. Population trends and flight behavior of the American Burying Beetle (Coleoptera: Silphidae), on Block Island, RI. Journal of Insect Conservation 10(4): 317-322. Shuster, J.D., A. Mckenna-Foster, L. Perrotti, R.S. Kennedy. 2008. American burying beetle (Nicrophorus americanus) survey on Nantucket Island, Massachusetts 2008. Submitted to USFWS, Region 5, December 2008. Threatened and Endangered Species System (TESS). USFWS. http://ecos.fws.gov/tess_public/ Accessed December 1, 2013. USFWS Memorandum. September 22, 1998. Summary of American Burying Beetle (Nicrophorus americanus) From Michael Amaral, Sr. Endangered Species Specialist, New England Office (NEFO) to Supervisor, New England Field Office of Endangered Species Coordinator, Region 5. USFWS. 2008. American burying beetle (Nicrophorus americanus) Five-Year Review: Summary and Evaluation. Unpubl. rep. USFWS New England Field Office, Concord, New Hampshire 46 pp. USFWS. 2013. DRAFT American Burying Beetle Nicrophorus americanus Range Wide Survey Guidance. Updated 4-20-2012. Available at http://www.fws.gov/arkansas-es/docs/DRAFT%20ABB%20Range%20wide%20Survey%20Guidance%204_20_2012.pdf

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APPENDIX A 2015 ABB Notching Schemes

Site 6

Inner R

Site 9

Inner L

Inner R

Site 14

Outer L

Outer R

Site 8

Outer R

18

Site 11

Outer R

3:00 Pro

Site 3

Inner R

3:00 Pro

Site 5

Outer L

3:00 Pro

Site 1

Outer R

Inner L

Site 4

Outer L

9:00 Pro

Site 12

Inner L

19

Site 7

9:00 Pro

Late Summer 2015

Outer R

Late Summer Trap 2014

2015 Recapture

Outer R

7:00 Pro

Site 10

3:00 Pro