Flexible Foraging of the Hawaiian Hoary Bat on Maui

An update of the H. T. Harvey & Associates ecological research on Opeapea

Dave Johnston, Kristin Jonasson, and Brad Yuen

ESRC Meeting Honolulu, 5 March 2020 Strategy for Recovery of the Species

• Which habitats do they use?

• How much land does a bat use?

• What do they eat? Study Area & Bat Detector Locations

General Random Tessellation Stratified survey design Acoustic Monitoring - Methods

Bi-monthly 9 habitats 3 nights 5 replicates / month

315 total deployments SM4 bat detectors 35 bat detector sites per each habitat Relationship between Precipitation and the Nine Habitats in the Study Area. Modelling of Acoustic Data - Methods

• Differences between habitat activity - generalized linear model fit by maximum likelihood with a negative binomial distribution date and site as random factors, and habitat as fixed effect of interest.

• Tested for differences between months within each habitat, habitats within each month, using pairwise contrasts with a Tukey adjustment for comparing among estimates Methods: Determining Core Use Areas

Mist-netting at “hot spots” of bat activity

Radio-telemetry of foraging bats using triangulation by mobile antennae and fixed stations

Analysis of data to determine core use area (CUA) 50% Kernel and foraging range (FR) 95% kernel. Reanalized data using methods described in Bonaccorso et al. 2015. Prey Availability

Bi-monthly UV light collection trap

9 habitats

Extraordinary sample set Identification of from Light Traps

Moths ID by Matt Medeiros

Slide preparations of Male Genitalia of Darna pallivitta and abydata Used to Identify the Species. Modelling of Dry Weights of per Habitat and Month

• We fit a negative binomial generalized linear model with a log link function from the MASS package (Venables and Ripley 2002)

• We used the estimated marginal means (emmeans) (Searle et al. 1980) function from the emmeans software package in R (Lenth 2009) to generate estimated means for each group

• We tested for differences between months and habitats using pairwise contrasts with a Tukey adjustment (Lenth 2009) from Haarsma 2016 DNA Extraction, Amplification, Polymerase Chain Reaction, Library Prep and Sequencing

Targeted CO1 region with primers from Zeale et al. (2010)

Targeted the 16s region with primers from Epp et al. (2012) Results: Acoustic Monitoring

Habitat Use – All Habitats Results: Acoustic Monitoring

Calls per Night by Habitat

alpha = 0.05 (P < 0.01)

Schlaepfer MA (2018) Do non-native species contribute to biodiversity? Calls per Night over Time by Habitat

Low Density Development

Grassland

Gulch Calls per Night by Habitat

Lower Forest Upper Forest Calls per Night by Habitat

Lower Forest Upper Forest Results: Capture and Tracking Success

20 bats caught; 11 ranges mapped

Mean number days tracked : 5.3 +/- 1.5 Core Use Area – 50% Legend Kernel Mean = 3,991 Hectares 50% Kernel Foraging Areas Bat ID - Gender, Age Range = 2.8 to 19,830 Hectares 2 - Male, Subadult Haiku 3 - Female, Subadult 4 - Male, Adult Paia 5 - Male, Adult Kahului 6 - Male, Adult 7 - Male, Adult 9 - Male, Adult 10 - Male, Adult 11 - Male, Adult 12 - Female, Adult 20 - Male, Adult

Maalaea Kihei Maalaea Bay u K Haleakala National Park Maui

4 2 0 4

± Miles Home Range – 95% Kernel Legend Mean = 17,362 Hectares 95% Kernel Core Areas Bat ID - Gender, Age Range = 14.6 to 81,341 Hectares 2 - Male, Subadult Haiku 3 - Female, Subadult 4 - Male, Adult Paia 5 - Male, Adult 6 - Male, Adult Kahului 7 - Male, Adult 9 - Male, Adult 10 - Male, Adult 11 - Male, Adult 12 - Female, Adult 20 - Male, Adult

Maalaea Kihei Maalaea Bay u K Haleakala National Park Maui

4 2 0 4

± Miles Legend Paia Capture Haiku Paia i Location Kahului Kernel Areas (HT Harvey) Kihei 95% 50% Bat ID Makawao 2 3 4 Kihei 5 6 Haleakala 7 National Park Kernel Areas (USGS)

0 8 Miles 0 4 Miles 0 5 Miles Bat 2 Bat 3 Bat 4 u

Kihei

Kihei e g an R d x e m m . o 7 o H t \ s 2 l e a u ur d g i i v F nd i I 01 \ -

8 H 7 od _ 9 h t 3 \

e Na 0 m S 39 0 G

S 0 4 Miles 0 4 Miles 0 4 Miles cts

U 4 2 0 4 e \

j Bat 5 Bat 6 Bat 7 t o r po r P \ e : R N Miles 95% and 50% Kernel Foraging and Core Areas per Individual Bat Using HT Harvey and USGS Methodology Ecology of the Hawaiian Hoary Bat (3978-01) March 2020 Paia Haiku Paia

0 4 Miles 0 4 Miles Bat 11 Bat 9 0 2 Miles Bat 10

lui Haiku Paia Kahului

Legend Capture Location ei Kihei Kernel Areas (HT Harvey) 95% 50% Bat ID e g

an Haleakala 9 R e National Park m 10 o H \ s e 11 ur g i d F x N

m 12 . 01 \ - 0 8 2 7 o t 9 20 9 3 l \ 0 du a

i Kernel Areas (USGS) 39 0 v i

d 0 5 Miles 0 8 Miles cts n

I 4 2 0 4 e \

j Bat 12 Bat 20 t o r po r P \ e : R N Miles 95% and 50% Kernel Foraging and Core Areas per Individual Bat Using HT Harvey and USGS Methodology Ecology of the Hawaiian Hoary Bat (3978-01) March 2020 Mean Number of Telemetry Fixes within Habitat Types from Pooled Data from all Bats

16

14

12

10

8

6

4

2

0 AV DevH DevL FWL FWU Grass Gulch Shrub SV

-2 Number of telemetry fixes in habitat types per bat and by season

Fall 100

90

80

70

60 Summer Spring 50 Fall

40 Winter 30

20

10

0 AV DevH DevL FWL FWU Grass Gulch Shrub SV Bat 2 Bat 3 Bat 4 Bat 5 Bat 6 Bat 7 Bat 9 Bat 10 Bat 11 Bat 12 Bat 20 Legend Study Area (34226 ha) Kahului Bat #5 Telemetry Fixes Kernel Areas (HT Harvey) Bat #5 50 95 September 2017 Acoustic Data Means Calls each Night, Habitat Type 1.6, Forest Upper 9.3, Agriculture 10, Forest Lower 20.4, Gulch Maalaea 21, Shrubland Bay 21.3, Sparsely vegetated 28.9, Grassland Kihei 29.7, Developed Low Int 32.2, Developed High Haleakala Landcover Basemap National Park (outside of Areas with Acoustic Monitoring Data) Alien Forest/Tree Plantation Agriculture c i st Developed, high-intensity u o

Ac Developed, low-intensity er v o 5 t Grassland a B \ t Native forest po r e R e

g Shrubland an R e Sparsely vegetated m o H \ s e ur g i F 01 \ - 8 7 9 3 \ 0 2.75 1.375 0 2.75 39 0 d cts x e j m o r s. Miles b P \ a : H N Bat #5 Telemetry Over September 2017 Acoustic Monitoring Data Ecology of the Hawaiian Hoary Bat (3978-01) February 2020 Legend Study Area (34226 ha) Kahului Bat #9 Telemetry Fixes Kernel Areas (HT Harvey) Bat #9 50 95 March 2018 Acoustic Data Mean Calls Each Night, Habitat Type 0.53, Forest Lower 0.8, Grassland 0.93, Agriculture 2.27, Developed (High) Maalaea 3.93, Sparsely vegetated Bay 4, Gulch 5.6, Developed (Low) Kihei 7.5, Forest Upper 18.3 Shrubland Haleakala Landcover Basemap National Park (outside of Areas with Acoustic Monitoring Data) Alien Forest/Tree Plantation Agriculture c i Developed, high-intensity st u o

Ac Developed, low-intensity er v o Grassland 9 t a B \ t Native forest po r e R

e Shrubland g an R Sparsely vegetated e m o H \ s e ur g i F 01 \ - 8 7 9 3 \ 0 2.75 1.375 0 2.75 39 0 d x cts m e . j 2 o r s Miles b P \ a : H N Bat #9 Telemetry Over March 2018 Acoustic Monitoring Data Ecology of the Hawaiian Hoary Bat (3978-01) February 2020 Detail of Bat 9 Fixes over Habitat Types N:\Projects3900\3978-01\Figures\Home Range Report\Bat2 over Acoustic Habs.mxd a Maalaea Bay 2 . 7 5 Ka 1 . 3 7 hu 5 l ui Miles 0 Kihei 2 . 7 5

K u

k Telemetry Telemetry #2 Bat National Haleakala Park Over Over Kernel Areas (HT Areas Kernel September September Legend (outside of of (outside each Habitat MeansNight, Calls Bat Landcover July #2 Ecology Ecology Developed, high Developed, Agriculture Alien 10, 9.3, 1.6, Forest 95 50 Bat (34226 ha)Area Study Sparsely Shrubland forest Native Grassland 32.2, Developed 29.7, Developed 28.9, 21.3, Sparsely 21, 20.4, 2017 Shrubland Forest Agriculture #2 Areas withAreas Gulch Grassland Forest/Tree Plantation Forest/Tree Plantation Basemap 2017 Telemetry Hawaiian Hoary Bat Hoary Hawaiian the of Acoustic Acoustic vegetated Lower Upper Harvey) Acoustic low vegetated Acoustic Monitoring Monitoring Acoustic - intensity - High Low Low Fixes intensity intensity Data Monitoring Int Int Type February Data) (3978 Data 2020 - 01) Legend Study Area (34226 ha) Kahului Bat #7 Telemetry Fixes Kernel Areas (HT Harvey) Bat #7 50 95 January 2018 Acoustic Data Mean Calls Each Night, Habitat Type 0.13, Forest Lower 1.2, Grassland 3.7, Agriculture 3.73, Sparsely vegetated Maalaea 10.7, Developed (High) Bay 11.7, Developed (Low) 12.5, Gulch Kihei 13.5, Shrubland 45, Forest Upper Haleakala Landcover Basemap National Park (outside of Areas with Acoustic Monitoring Data) Alien Forest/Tree Plantation Agriculture c i Developed, high-intensity st u o

Ac Developed, low-intensity er v o Grassland 7 t a B \ t Native forest po r e R

e Shrubland g an

R Sparsely vegetated e m o H \ s e ur g i F 01 \ - 8 7 9 3 \ 0 2.75 1.375 0 2.75 39 0 d x cts m e . j 2 o r s Miles b P \ a : H N Bat #7 Telemetry Over January 2018 Acoustic Monitoring Data Ecology of the Hawaiian Hoary Bat (3978-01) February 2020 Model Estimates for Dry Weight by Habitat Percentage of Prey Items Based on Number of Sequences Detected by Order for 11 Bats Coleoptera, 0.04

Diptera, 0.12

Isoptera, 0.09

Hemiptera, 0.02 , 0.68 Orthoptera, 0.05 Important prey species in ~ ½ of guano samples

Herpetogramma licarsisalis Tropical grass webworm

Marion Friedrich

Peridroma saucia Variegated cutworm Dwaine Wagoner

Athetis thoracica moth

Kendrick/C&R Wildlife Summary Bats from Maui vs. Hawaii used very different habitats! Mostly in Gulches, Low density developed and Grasslands Most forests on Maui are non-native that have few insects and are not productive Maui Bats’ core use areas were highly variable in size, and averaged many times the size of the Hawaii bats’ core use areas

Bats moved up and down elevation gradient year round. Bats tended to forage in lower altitudes in Summer, Fall, and Spring, and in higher altitudes in Winter

Bats ate primarily moths and very few beetles. They ate native and non-native insects and appeared somewhat selective

Bats of each season used very similar habitats except for during winter when each of 3 bats used different habitats Discussion

• Bats should be managed on an Island by Island basis. Different habitats and different CUA sizes

• CUA sizes may not be as important as the quality of the habitat.

• Bats foraged in patchy habitats so conservation efforts may not necessarily need be contiguous.

• We advocate enhancing or improving habitat as opposed to the purchase of a specific amount of forest as mitigation for take. Mahalo!

Frank Bonaccorso and Corinna Pinzari, Native Nursery, Kama'ole Ranch, 'Ulupalakua Ranch, Haleakala Ranch, Maui Nui Farms, Maui Bees, Ali'i Kula Lavender Farm Deb Bauer, Bazie Brandt, Danny and Jenta Brictson, Kelly Bryce, Monroe Bryce, Andrea Buckman, Keahi Bustamente, Lois Campbell, Terry Chang, Mark Damon, Kerri Fay, Dominic Gambino, Geoff Haines, Joseph Imhoff, Hadley Luis, Debra Lordan, Mary Jane Gapero, Linsey Manuel, Wisa Miller, Molly O’Grady, Evan Ryan, Zach Pezzillo, Michelle Smith, Andrei Stanescu, Ruth-Marie Stecker, Monte Tudor-long, Doc Vitale, Heather Wade, Ian Wade, John Wilson, Jamie Woodburn, Bonnie Young Terry Zinn and many more. Questions?