Draft Revised Recovery Plan for the 

Mariana Fruit Bat or Fanihi 

(Pteropus mariannus mariannus) 

November 2009 

(Original approved 2 November 1990) 

Region 1 

U.S. Fish and Wildlife Service 

Portland, Oregon 

Approved: XXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXX

 Regional Director, U.S. Fish and Wildlife Service 

Date: XXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXXX
Draft Revised Recovery Plan for the Mariana Fruit Bat 

ii

DISCLAIMER

Recovery plans delineate reasonable actions that are believed to be 

required to recover and/or protect listed species. We, the U.S. Fish and Wildlife 

Service, publish recovery plans, sometimes preparing them with the assistance of 

recovery teams, contractors, State agencies, Tribal agencies, and others. The 

necessary funds to attain objectives identified in a recovery plan are subject to 

budgetary and other constraints affecting the parties involved, as well as the need 

to address other priorities. Costs indicated for action implementation and time for 

achievement of recovery are only estimates and subject to change. Recovery 

plans do not necessarily represent the views nor the official positions or approval 

of any individuals or agencies involved in plan formulation, other than the U.S. 

Fish and Wildlife Service. Recovery plans represent the official position of the 

U.S. Fish and Wildlife Service only after they have been signed by the Director or 

Regional Director as approved. Approved recovery plans are subject to 

modification as dictated by new findings, changes in species status, and the 

completion of recovery actions. 

Literature citation of this document should read as follows: 

U.S. Fish and Wildlife Service. 2009. Draft Revised Recovery Plan for the 

Mariana Fruit Bat or Fanihi (Pteropus mariannus mariannus). U.S. Fish and 

Wildlife Service, Portland, Oregon. xiv + 83 pp. 

Electronic copies of this document will be made available at: 

• http://pacific.fws.gov/ecoservices/endangered/recovery/default.htm 

• http://endangered.fws.gov/recovery/index.html 
Draft Revised Recovery Plan for the Mariana Fruit Bat 

iii

ACKNOWLEDGMENTS 

This draft revised recovery plan was written with extensive input and assistance 

from the Fanihi Recovery Team. A recovery team is a formal advisory group that 

provides information and advice about recovery needs and opportunities for 

species listed as threatened or endangered. The recovery team focuses on 

development of the recovery plan and may be involved in implementation of the 

plan. Recovery teams are convened at the discretion of the Regional Director and 

are not required. The U.S. Fish and Wildlife Service has administrative 

responsibility for preparing recovery plans and approving their content. 

Members of the Fanihi Recovery Team are identified below. Many other 

individuals also contributed their time and expertise to the preparation of this 

document, including Fred Amidon, Nathan Hawley, Curt Kessler, Patrick 

Leonard, Gina Shultz, and Marilet Zablan of the U.S. Fish and Wildlife Service’s 

Pacific Islands Fish and Wildlife Office, George Phocas of the U.S. Fish and 

Wildlife Service’s Office of Law Enforcement, Scott Vogt of the U.S. Navy, and 

Dustin Janeke of e2M Environmental and Engineering Management. 
Draft Revised Recovery Plan for the Mariana Fruit Bat 

iv

Members of the Fanihi Recovery Team: 

Herman A. Apatang 

Commonwealth of the Northern 

Mariana Islands 

Department of Lands and Natural 

Resources 

P.O. Box 924 

Songsong Village 

Rota, MP 96951 

Anne Brooke 

Joint Region Marianas 

PSC 455, Box 195 

FPO AP 96540-2937 

Joseph T. Lizama 

CNMI Department of Lands and Natural 

Resources 

P.O. Box 172 

Tinian, MP 96952 

Dana T. Lujan 

USDA/APHIS/Wildlife Services 

233 Pangelinan Way 

Barrigada, GU 96913 

Tammy Mildenstein 

Department of Ecosystem and 

Conservation Sciences 

University of Montana 

Missoula, MT 59812 

Laura Williams 

CNMI Division of Fish and Wildlife 

P.O. Box 10007 

Saipan, MP 96950 

Jeffrey Quitugua 

Guam Division of Aquatic and Wildlife 

Resources 

192 Dairy Road 

Mangilao, GU 96923 

Chris Tidemann 

Australia National University 

School of Resources, Environment and 

Society 

Canberra 0200 Australia 

Gary Wiles 

Washington Department of Fish and 

Wildlife 

521 Rogers St. SW 

Olympia, WA 98502 

Julia Boland 

CNMI Division of Fish and Wildlife 

PO Box 1397 

Rota, MP 96951 

Dave Worthington 

Capitol Reef National Park 

HC 70 Box 15 

Torrey, UT 84775 

Holly Freifeld (Team Liaison)

U.S. Fish and Wildlife Service 

300 Ala Moana Blvd., Rm.3-122 

PO Box 50088 

Honolulu, HI 96850
Draft Revised Recovery Plan for the Mariana Fruit Bat 

v

EXECUTIVE SUMMARY 

Current Species Status 

The Mariana fruit bat or fanihi (Pteropus mariannus mariannus) is a 

subspecies endemic to the Mariana archipelago (the Territory of Guam and the 

Commonwealth of the Northern Mariana Islands [CNMI]), where it is known 

from 14 of the 15 major islands. Surveys on most or all islands in the archipelago 

were conducted in 1983 (Wiles et al. 1989), 2000 (Cruz et al. 2000a-f), and 2001 

(Johnson 2001). These surveys indicated that the relatively isolated northern 

islands of the CNMI (i.e., the islands north of Saipan) have supported the majority 

of fanihi in the archipelago for at least the past few decades, despite the relatively 

small land mass of these islands in comparison to the southern islands. 

Survey effort for fanihi throughout the archipelago historically has been 

uneven. Despite this, published reports indicate that fanihi numbers on the 

southern islands in the archipelago declined precipitously through the 20th century 

as a result of hunting and habitat loss and degradation (Fritz 1901, 1904; Schnee 

1911; Lemke 1984; Wiles et al. 1989; Marshall et al. 1995; Wiles 1996; 

Worthington and Taisacan 1996; Krueger and O’Daniel 1999). Habitat loss and 

degradation has resulted from agriculture; introduction of non-native ungulates, 

plants, and other organisms; economic development; and war (Baker 1946; 

Bowers 1950; Fosberg 1960; Stone 1970). In the remote northern islands of the 

archipelago, chronic habitat degradation by ungulates, hunting, and, more 

recently, volcanic eruption on Anatahan, may be responsible for declines in bat 

numbers over the past several decades (Wiles et al. 1989; Rice and Stinson 1992; 

Marshall et al. 1995; Kessler 2000a, Cruz et al. 2000a-f). 

Although variation in survey methods and effort render rigorous comparisons 

between islands and surveys periods difficult, conservative interpretation of 

existing survey data indicates a 40 percent decline in fruit bat numbers between 

1983 (Wiles et al. 1989) and 2000 (Cruz et al. 2000a-f) among the six northern 

islands surveyed in both years. The majority of this decline was recorded on two 

of the three largest northern islands, Anatahan and Pagan, which together 

harbored the majority of the archipelago’s fanihi in the 1980s (Wiles et al. 1989). 
Draft Revised Recovery Plan for the Mariana Fruit Bat 

vi

Habitat Requirements 

Fanihi habitat use is influenced by several characteristics of the species. 

Fanihi typically are highly colonial, and can form large, dense roosts in multiple 

adjacent trees. Fruits, nectar, pollen, and some leaves comprise the majority of 

the bats’ diet; rapid digestion and metabolism of such foods makes these animals 

reliant on forest habitat containing diverse food resources that are available 

throughout the year. Fanihi are strong fliers and highly mobile; although the 

pattern and frequency of interisland movements is unknown, fruit bats have been 

observed flying over the ocean between islands. Connectivity of the 

archipelago’s islands for fanihi depends on the presence of enough suitable forest 

for roosting and foraging to sustain resident and in-transit bats. 

 

Fanihi forage and roost primarily in native forest and forage occasionally 

in agricultural forests composed primarily of nonnative plants (Wiles 1987b; 

Worthington and Taisacan 1996). Wiles (1987b) described six bat roost sites on 

Guam, all within native limestone forest. On Rota, fanihi used primary and 

secondary limestone forest for roosting and foraging; at least eight native and one 

introduced tree species were used for roosting (Glass and Taisacan 1988). A 

small colony also was observed roosting in Casuarina equisetifolia (ironwood) 

trees on Aguiguan (Worthington and Taisacan 1996). On Sarigan, fanihi were 

observed roosting and foraging in the small patch of native forest, and foraging to 

a lesser extent in the large area dominated by coconut (Wiles and Johnson 2004). 

Limiting Factors

Five factors are considered in decisions to list, delist, or reclassify a species. 

These factors are: 

A – The present or threatened destruction, modification, or curtailment of its 

habitat or range; 

B – Overutilization for commercial, recreational, scientific, or educational 

purposes; 

C – Disease or predation; 

D – Inadequacy of existing regulatory mechanisms; and 

E – Other natural or man-made factors affecting its continued existence. 
Draft Revised Recovery Plan for the Mariana Fruit Bat 

vii

Factor A: The degradation and loss of primary and other forest habitats resulting 

from ungulate damage, invasion by alien plant species, and economic 

development has substantially diminished the extent of habitat for fruit bats in the 

Mariana archipelago since human settlement of the islands; all of these processes 

accelerated during the 20th century. These sources of habitat degradation and loss 

all are still present in the islands and may lead to further reduction in the 

availability of resources critical for the survival and reproduction of fanihi and 

thus to a potential reduction in the number of bats that the remaining habitat is 

able to support. 

Factor B: Fanihi have been used as food since humans first arrived on the islands, 

and consumption of bats represents a significant cultural tradition. Overhunting, 

however, is cited as a causal factor in the initial fanihi declines on Guam, Saipan, 

and Tinian. In response to plummeting fanihi numbers, hunting of bats became 

illegal under local law in both Guam and the CNMI in the 1970s, and commercial 

trade in fanihi largely has been curbed. However, illegal hunting for local 

consumption remains a chronic, population-level threat that precludes the 

recovery of the species to healthy, self-sustaining numbers. 

Factor C: The brown treesnake (Boiga irregularis), which has caused the 

extinction or extirpation of most native landbird species on Guam, is considered 

capable of preying on non-volant young bats, and may contribute to the lack of 

recruitment of young bats on Guam. 

Factor D: Although current CNMI hunting regulations (Part 4, Section 10.7.i) 

prohibit the hunting, killing, or possessing of protected species, including fanihi, 

and the CNMI Department of Fish and Wildlife (DFW) has statutory authority to 

promulgate and enforce these regulations to protect bats and impose fines for 

violations, it has been reported that there is little enforcement of the hunting ban, 

and few investigations or convictions have taken place. The Mariana fruit bat is 

listed as an endangered species by the Government of Guam and take is 

prohibited under this designation, but it is widely believed that illegal hunting of 

Guam’s few remaining bats occurs opportunistically. 

Factor E: The small number of fanihi remaining on some islands (e.g., Guam and 

Saipan) may face significant risk of extirpation from natural disturbances, 
Draft Revised Recovery Plan for the Mariana Fruit Bat 

viii

environmental changes, and other chance events to which small populations 

typically are vulnerable. Fanihi do fly between islands, although the frequency of 

these events throughout the Mariana archipelago is unknown. Recolonization of 

islands where bats were extirpated likely has occurred in the past, but the 

likelihood of recolonizations probably is declining with the decline in total 

numbers and with gaps of one or more islands in the bats’ distribution. Although 

this subspecies has evolved in the presence of natural disturbance, today a 

declining population and anthropogenic threats such as illegal hunting erode the 

resilience of the population and reduce the likelihood of complete recovery in the 

wake of typhoons and volcanic eruptions. Typhoons, in particular, could 

eliminate bats that persist in small numbers on one or more of these islands (see 

Wiles and Brooke in press for a discussion of density-dependent typhoon effects). 

Military training activities such as live fire and aircraft overflight exercises in 

areas used by fanihi could disrupt the behavior of these bats and may impede 

recolonization if military activities create a gap in distribution. Farallon de 

Medinilla may provide an example of this situation: the use of this island as a 

bombing range has rendered it unsuitable habitat for resident fanihi (although a 

transiting bat may occasionally use the island as stopover), thus creating a 

disjunction between the northern and southern islands in the archipelago. An 

increase in air traffic at Andersen Air Force Base, which harbors the single 

remaining fruit bat colony on Guam, is likely to occur in conjunction with a 

proposed base expansion. 

Recovery Priority Number 

The Mariana fruit bat is assigned a recovery priority number of 9 on a scale of 1C 

(highest) to 18 (lowest; the “C” indicates the potential for conflict with human 

economic activities), based on the moderate degree of threat, a high potential for 

recovery as stated above, and its status as a subspecies. 

Recovery Criteria 

The recovery criteria discussed below address the major threats to the species, 

including general criteria for population distribution and post-delisting 

monitoring, as well as additional criteria organized by the five Listing Factors to 
Draft Revised Recovery Plan for the Mariana Fruit Bat 

ix

be used in a delisting analysis. The fanihi may be considered for delisting when 

the following criteria have been met: 

1. Population and Distribution 

The total size of the fanihi population has increased, based on data from 

scientifically defensible monitoring protocols, and stable or increasing 

subpopulations of sufficient size to avoid genetic and demographic risks 

associated with very small populations are distributed among the Mariana 

Islands so that the probability of the fanihi’s persistence over 100 years is high 

(at least 90 percent). 

To meet this criterion, we propose that stable or increasing fanihi 

subpopulations should be distributed as follows: three of the five southern 

islands (Saipan, Tinian, Aguiguan, Rota, and Guam), and six of the eight 

islands north of Saipan where fanihi have persisted historically (Anatahan, 

Sarigan, Guguan, Alamagan, Pagan, Agrihan, Asuncion, and Maug). Of the 

six northern islands with stable or increasing fanihi numbers, two of these 

must include Pagan, Anatahan, or Agrihan, the three largest of the northern 

islands. This distribution may be modified as appropriate based on future 

monitoring data or population viability modeling (Recovery Action 5.2). 

2. Post-delisting monitoring 

A post-delisting monitoring program for the species has been developed to 

reliably detect population trends and is ready for implementation. 

3. Listing Factor A: Habitat Loss and Degradation 

Specific actions to restore habitat (including ungulate control and control of 

invasive non-native plants as appropriate for specific islands – see Table 3) 

have been identified and management plans developed as necessary for 

recovery under Criterion 1, and these actions and plans are being successfully 

implemented so that habitat loss and degradation no longer endanger the 

survival of the fanihi. 
Draft Revised Recovery Plan for the Mariana Fruit Bat 

x

4. Listing Factor B: Hunting 

Specific actions to reduce illegal hunting have been identified and 

management plans developed as necessary for recovery under Criterion 1, and 

these actions and plans are being successfully implemented so that illegal 

hunting no longer endangers the survival of the fanihi. 

5. Listing Factor C: Brown Treesnakes 

Long term measures are being successfully implemented to control the 

incipient brown treesnake population on Saipan and to prevent the 

introduction of the brown treesnake from Guam and Saipan to other islands in 

the CNMI. 

6. Listing Factor E: Development and Military Training Activities 

Impacts of urban development and military training on the fanihi are 

successfully being avoided, minimized, or mitigated so that they do not 

endanger the survival of the fanihi. 

Date of Recovery: Delisting could occur by 2030 if all of the criteria have 

been met. 

Important Recovery Actions (for details, see the Narrative Outline and 

Implementation Schedule): 

Recovery of the fanihi focuses on the following actions: 

1. Immediate management to reduce risks and stabilize the existing population. 

2. Specific actions to reduce or eliminate illegal hunting to allow increase in 

fanihi numbers throughout the archipelago. 

3. Protection of the best existing habitat and enhancement of additional suitable 

habitat. 

4. Effective control and interdiction of the brown treesnake. 

5. Research to address gaps in our knowledge of fanihi life history and ecology 

and improve our ability to model the population, assess its sensitivity to 

specific threats and management actions, and forecast its persistence. This 

research is essential to eventually determining thresholds for bat numbers and 

other conservation criteria on each island that could indicate when the fanihi 
Draft Revised Recovery Plan for the Mariana Fruit Bat 

xi

population has recovered sufficiently for some legal hunting to resume. The 

fanihi population is clearly too precarious at present to support a hunting 

season. However, if illegal hunting and other threats are reduced so the 

species’ status can improve substantially and our knowledge of population 

dynamics and threats allows us to identify a limited, sustainable hunting 

regime consistent with recovery of the species, such a hunt might be allowed 

under the Endangered Species Act either by delisting the species as indicated 

above or through a special rule under section 4(d) of the Act. 

Total Estimated Cost of Recovery 

The estimated cost for recovery actions over the next 5 years is $12,578,000, 

distributed as follows: 

Year Priority 1 Priority 2 Priority 3 Total 

2010 163,000 2,849,000 253,000 $ 3,265,000 

2011 170,000 2,379,000 200,000 $ 2,749,000 

2012 170,000 2,329,000 101,000 $ 2,600,000 

2013 163,000 2,159,000 60,000 $ 2,382,000 

2014 163,000 1,979,000 71,000 $ 2,213,000 

Total $ 829,000 $ 11,695,000 $ 685,000 $ 13,209,000 
Draft Revised Recovery Plan for the Mariana Fruit Bat 

xii

TABLE OF CONTENTS 

ACKNOWLEDGMENTS ................................................................................................. iii

EXECUTIVE SUMMARY .................................................................................................v

TABLE OF CONTENTS .................................................................................................. xii

I. BACKGROUND............................................................................................................1

A. Status Overview and Structure of the Recovery Plan ...................................................1

B. The Mariana Islands......................................................................................................2

C. Species Description and Taxonomy..............................................................................4

D. Historical and Current Population Status and Distribution...........................................5

1. Guam ........................................................................................................................6

2. CNMI Southern Islands............................................................................................8

3. CNMI Northern Islands..........................................................................................11

E. General Biology and Ecology .....................................................................................14

1. Habitat Use: Roosting and Foraging ......................................................................14

2. Life History ............................................................................................................15

3. Interisland Movement.............................................................................................17

F. Reasons for Decline and Current Threats....................................................................19

1. Hunting [Listing Factor B].....................................................................................19

2. Habitat Loss and Degradation [Listing Factor A] ..................................................26

3. Predation by Brown Treesnakes [Listing Factor C] ..............................................30

4. Natural Disturbance, Development, and Military Training Activities 

[Listing Factor E]....................................................................................................31

G. Conservation Efforts ...................................................................................................34

II. RECOVERY ...............................................................................................................37

A. Recovery Goal and Objective .....................................................................................37

B. Recovery Strategy .......................................................................................................37

C. Recovery Criteria ........................................................................................................39

III. RECOVERY ACTIONS..............................................................................................42

A. Outline of Recovery Actions.......................................................................................42

B. Recovery Action Narrative..........................................................................................45

IV. RECOVERY IMPLEMENTATION SCHEDULE FOR 2010-2014........................62

V. REFERENCES............................................................................................................72

A. Literature Cited ............................................................................................................72

B. Personal Communications............................................................................................81

C. In Litt. References ........................................................................................................82
Draft Revised Recovery Plan for the Mariana Fruit Bat 

xiii

LIST OF FIGURES 

Figure 1. The Mariana Islands. ...................................................................................................... 3

LIST OF TABLES 

Table 1. Summary of results from two archipelago-wide fanihi surveys ........................................ 7

Table 2. Plant parts commonly eaten by fanihi............................................................................. 16

Table 3. Island summary of historical factors and current threats affecting the fanihi................. 20

Table 4. Estimated extent of fanihi habitat in the Marianas ......................................................... 28
Draft Revised Recovery Plan for the Mariana Fruit Bat 

1

I. BACKGROUND 

A. Status Overview and Structure of the Recovery Plan 

 The Mariana fruit bat, or fanihi (Pteropus mariannus mariannus), was 

Federally listed as endangered on Guam in 1984 (USFWS 1984). In January 

2005 we published a final rule listing P. m. mariannus as threatened throughout 

its range (the Mariana archipelago: Guam and the Commonwealth of the Northern 

Mariana Islands (CNMI)) (USFWS 2005). This rule represented a reclassification 

of the fanihi on Guam from endangered to threatened, a change reflecting our 

determination that the fruit bats on Guam are a subset of a single population 

distributed throughout the archipelago. We determined that the fanihi faces 

threats that should be managed rangewide, and in this expanded geographic 

context, the fanihi merited listing as threatened throughout the islands. 

 We prioritize listed species for recovery, assigning recovery priority to a 

species based on degree of threat, recovery potential, taxonomic status, and 

conflict with human activities. Numerical ranks range from 1 to 18, with a letter 

designation of “C” indicating conflict with human economic activities. The 

highest priority is 1C; the lowest priority is 18 (USFWS 1983). The fanihi has a 

recovery priority number of 9, indicating that it faces a moderate degree of threat 

(i.e., extinction is not imminent, but threats are ongoing), has a high potential for 

recovery, and has taxonomic status as a subspecies. Conservation of the fanihi 

generally is not in conflict with economic activities. A final rule designating 

critical habitat for the fanihi on Guam was published in the Federal Register on 

October 28, 2004 (USFWS 2004). Critical habitat refers to areas occupied by the 

species at the time of listing under the Endangered Species Act that contain 

resources essential for the species’ survival and reproduction, and that may 

require special management so that the species may recover. No critical habitat 

was designated in the CNMI because at that time the fanihi was not listed there. 

Therefore, the critical habitat designated for fanihi on Guam does not reflect the 

true extent of habitat required for the survival of the fanihi throughout the 

archipelago. Recovery planning is not constrained by critical habitat designations 

and this recovery plan includes all those actions and areas we deem necessary for 

the recovery of the fanihi. 
Draft Revised Recovery Plan for the Mariana Fruit Bat 

2

The original recovery plan for the fanihi on Guam was issued in 1990 

(USFWS 1990). In the subsequent 18 years, knowledge of this species has 

improved. In addition to providing a recovery strategy, this recovery plan 

represents a revision of the original plan for Guam and incorporates significant 

new information about the Mariana fruit bat. The plan is divided into four main 

sections. Part I, Background, provides an overview of the biology of the 

subspecies, the reasons for its decline, current threats to its persistence, and 

current conservation measures. Part II, Recovery, describes the recovery strategy, 

goals, and objectives for the fanihi, and includes discussion of the criteria 

established to determine recovery. Part III, Recovery Actions, describes the 

conservation and research activities required to achieve the recovery goals. Part 

IV, the Implementation Schedule, provides in tabular form a list of the 

recommended recovery actions, with emphasis on actions needed to meet interim 

recovery objectives within the next five years. This structure reflects the need for 

effective adaptive management in advancing the recovery of the Mariana fruit bat, 

as many variables remain unknown and long-term planning without inherent 

flexibility is unlikely to succeed. These short-term implementation plans will be 

prepared every five years to reflect the knowledge gained and to refine the 

management program to maximize the success of the Mariana fruit bat recovery 

program. 

B. The Mariana Islands 

The Mariana Islands consist of the 15-island Commonwealth of the Northern 

Mariana Islands (CNMI) and the Territory of Guam, both within the jurisdiction 

of the United States (Figure 1). This archipelago lies in the Western Pacific 

approximately 1,500 kilometers (900 miles) east of the Philippine Islands, and 

extends 750 kilometers (470 miles) from 13o14' north latitude (N), 144 o45' west 

longitude (W) to 20o3' N, 144o54' W. The nine northernmost islands (Anatahan, 

Sarigan, Guguan, Alamagan, Pagan, Agrihan, Asuncion, Maug, and Uracas) are 

volcanic in origin, and the six southernmost islands (Guam, Rota, Aguiguan, 

Tinian, Saipan, and Farallon de Medinilla) are uplifted limestone plateaus, some 

of which have volcanic outcrops. Fanihi historically have inhabited all of these 

islands except Uracas, the northernmost island (Wiles et al. 1989). The climate in 

the archipelago is tropical, with daily mean temperatures of 75o to 90o Fahrenheit 
Draft Revised Recovery Plan for the Mariana Fruit Bat 

3

Ogasawara

Isls. (JAPAN)

PHILIPPINES

JAPAN

ASIA

NEW GUINEA

AUSTRALIA

MARIANA ISLS.

PALAU

0 250 500 750 1000 Miles

0 250 500 750 1000 Kilometers

Uracas

(Farallon de Pajaros)

Maug

Asuncion

Agrihan

Pagan

Alamagan

Guguan

Sarigan

Anatahan

Farallon

de Medinilla

Saipan

Tinian

Aguiguan

Rota

Guam

0 25 50 75 100 Miles

0 25 50 75 100 Kilometers

15° N

0°

15° S

30° N

130° E 145° E

Figure 1. The Mariana Islands.
Draft Revised Recovery Plan for the Mariana Fruit Bat 

4

(24o to 32o Celsius), high humidity, and average annual rainfall of 200 to 260 

centimeters (80 to 100 inches). The natural vegetation of the islands is 

predominantly various types of low-stature tropical and coastal forest, but these 

have been extensively altered, particularly over the past century (Fosberg 1960; 

Stone 1970; Engbring et al. 1986; Falanruw et al. 1989). Typhoons may strike 

the Mariana Islands during any month of the year, but are most frequent between 

July and October. The large southern islands (Guam, Rota, Tinian, and Saipan) 

support nearly the entire human population of the archipelago, with most people 

on Guam (155,000) and Saipan (62,000). The northern islands (north of Saipan) 

are either unoccupied or support only small human settlements (as many as 40 

people total, seasonally). 

C. Species Description and Taxonomy 

 

The fanihi is a medium-sized bat in the family Pteropodidae that weighs 330 

to 577 grams (0.66 to 1.15 pounds) and has a forearm length ranging from 13.4 to 

15.6 centimeters (5.3 to 6.1 inches). The underside (abdomen) is colored black to 

brown, with gray hair interspersed, creating a grizzled appearance. The shoulders 

(mantle) and sides of the neck are usually bright golden brown, but may be paler 

in some individuals. The head varies from brown to dark brown. The wellformed and rounded ears and large eyes give the face a canine appearance; 

medium and large members of the genus Pteropus are often called flying foxes. 

 The paleotropical genus Pteropus is represented by 67 species distributed 

across the Indian Ocean, southern Asia, Australia, and Oceania as far east as the 

Cook Islands (Simmons 2005). Thirty-nine species (58 percent) are considered 

critically endangered, endangered, threatened, near threatened, or vulnerable 

under the definitions of the IUCN (IUCN 2008). Most of these face the risk of 

extinction because of a combination of habitat loss and hunting (Wiles and 

Brooke in press). 

Excluding the possibly extinct little Mariana fruit bat (Pteropus tokudae), the 

fruit bats of the Mariana Islands consistently have been treated as one or two 

endemic subspecies of the single species P. mariannus; that is, the taxon (or taxa) 

occurring in the Marianas occurs nowhere outside the archipelago (e.g., Andersen 

1912; Kuroda 1938; Corbet and Hill 1980, 1986, 1991; Koopman 1982, 1993; 
Draft Revised Recovery Plan for the Mariana Fruit Bat 

5

Flannery 1995; Simmons 2005). Other subspecies of P. mariannus have been 

described from Palau (pelewensis), Kosrae (ualanus), Yap (yapensis), Ulithi 

(ulthiensis), and the Ryukyu archipelago (loochoensis) (Pierson and Raney 1992, 

Koopman 1993); some or all of these taxa are treated as separate species by other 

authors (e.g., Anderson 1912, Corbet and Hill 1980, Flannery 1995). Flannery 

(1995) and Simmons (2005) recognize two subspecies restricted to the Mariana 

Islands: the Mariana fruit bat or fanihi (P. mariannus mariannus) and the socalled Pagan fruit bat (P. mariannus paganensis). The slight morphological 

differences used to distinguish P. mariannus paganensis from P. mariannus 

mariannus, based on four specimens, are likely attributable to natural variation 

that occurs not only between islands but within individual island subpopulations 

(Tom Lemke, Montana Department of Fish, Wildlife, and Parks, in litt. 1986; 

Gary Wiles, Washington Department of Fish and Wildlife, pers. comm. 2005). 

Thus, the Pagan fruit bat is likely not distinct from the Mariana fruit bat (Pierson 

and Rainey 1992; Worthington and Taisacan 1996; G. Wiles, pers. comm. 2007), 

particularly in light of the strong evidence of movement and probable gene flow 

between islands of the archipelago (e.g., Wiles et al. 1989; Wiles and Glass 1990; 

Wiles and Johnson 2004). Future studies may elucidate the genetic relationships 

among the fanihi on different islands in the Marianas; at this time we do not 

consider Pteropus mariannus paganensis as distinct from Pteropus mariannus 

mariannus. Furthermore, recent analysis of tissue from P. mariannus collected in 

the Mariana Islands and Palau found that fruit bats in the Marianas were not 

distinct from one another when compared to Palau bats (Gary McCracken, 

University of Tennessee, pers. comm. 2007). 

D. Historical and Current Population Status and Distribution 

 Similar to other animal species, obtaining accurate estimates of fruit bat 

populations in Pacific archipelagos depends on regular monitoring, standardized 

survey methods, and consideration of the unique ecology and physiographic 

environment of bat populations in various island groups. In addition, monitoring 

of population size and trends depends on accurate assessment of detectability, 

variation in skill among observers conducting surveys, and the statistical power of 

survey data. The difficult terrain of the Mariana Islands, remote location of the 

northern islands of the CNMI, and the high cost of interisland transportation by 
Draft Revised Recovery Plan for the Mariana Fruit Bat 

6

sea and aerial surveys of individual islands introduce additional complications 

that have hindered the establishment of a standard monitoring program for fanihi. 

 No known historical records exist to document the status of the fanihi prior to 

the 20th century. The history of bat surveys and changes in numbers summarized 

in Table 1 and subsequent text represent a variety of methods and analyses. 

Furthermore, the existing data do not reflect consideration of detection problems 

peculiar to fruit bats or variation among observers or seasons, nor have they been 

collected in a manner appropriate for rigorous analysis to generate statistically 

meaningful confidence intervals (around individual points) or trends. 

 Prior to recent volcanic activity on Anatahan, the relatively isolated northern 

islands (i.e., the islands north of Saipan) supported the majority of the fruit bats in 

the archipelago, but because of their remote location, these islands have been 

surveyed much less frequently than the southern islands. Individual surveys have 

been conducted on several of the larger and more accessible northern islands at 

relatively frequent intervals, and surveys of all or most islands in the archipelago 

were conducted in 1983 (Wiles et al. 1989), 2000 (Cruz et al. 2000a-f), and 2001 

(Johnson 2001). 

 

 The survey methods used in the northern islands in 2001 were significantly 

different from those used in 1983 and 2000; we therefore consider only Wiles et 

al. (1989) and Cruz et al. (2000a-f) for purposes of comparison (Table 1). A 

conservative interpretation of this comparison indicates a decline in the northern 

islands between 1983 and 2000, especially on Anatahan and Pagan, which 

supported the largest numbers of fruit bats in the archipelago 20 years ago (Table 

1). Bat numbers in the southern islands also have declined during this time period 

(Esselstyn et al., 2006; Wiles et al. 2006; Anne Brooke, U.S. Navy, pers. comm. 

2007), although this is not evident in Table 1. 

1. Guam 

On Guam, the sighting of fanihi was considered to be “not... uncommon” in 

1920 (Crampton 1921). However, by 1931, bats were uncommon on Guam, 

possibly because the introduction of firearms led to more hunting (Coultas 1931). 

Woodside (1958) reported that in 1958 fanihi on Guam were estimated to number 
Draft Revised Recovery Plan for the Mariana Fruit Bat 

7

Table 1. Summary of results from two archipelago-wide fanihi surveys: 

minimum estimates. Two of the northern islands are not included in this table: 

Uracas, the most northerly, where fruit bats are not known to occur; and Farallon 

de Medinilla (FDM), where fanihi have been observed on only one occasion. 

Additional and more recent surveys and observations have been made on the 

southern islands, and on FDM, Anatahan, Sarigan, and Pagan (see text; Table 3). 

Island Area 

Sq. mi (Sq. km) 

Estimated 

minimum 

number of bats 

19831

Estimated 

number of bats 

20002

Maug 0.8 (2.0) <25 not surveyed 

Asuncion 2.9 (7.4) 400 not surveyed 

Agrihan 18.3 (47.4) 1,000 1,000 

Pagan 18.4 (47.7) 2,500 1,500 

Alamagan 4.3 (11.0) 03

 200 

Guguan 1.5 (4.0) 400 350 

Sarigan 1.9 (5.0) 125 200 

Anatahan 12.5 (32.3) 3,000 1,000 

Subtotal (Northern Islands) 

[Subtotal six islands] 

7,450 

[7,025] [4,250] 

Saipan 47.5 (122.9) <50 not surveyed 

Tinian 39.3 (101.8) <25 not surveyed 

Aguiguan 2.7 (7.0) <10 150-200 

Rota 32.9 (85.2) 800-1,000 not surveyed 

Guam 212.0 (549.0) 425-500 119-179 

Subtotal (Southern Islands) 1,310-1,585 insufficient data 

TOTAL (All Islands) 8,760-9,035 insufficient data 

1

 Wiles et al. 1989. Dates: August 17-September 10, 1983; 1-4 days/island. 

Count methods: Evening dispersal counts at colonies; evening station counts of 

solitary fruit bats. All counts considered to be minimum estimates. 

2 Cruz et al. 2000a-f. Dates: June 4-August 16, 2000; 7-9 days/island. Count 

methods: Evening dispersal counts at colonies, evening and morning station 

counts of solitary fanihi. Data for Guam represents the range of 10 counts 

conducted in a separate effort in 2000 (A. Brooke, pers. comm. 2007). 

3 Alamagan was inadequately surveyed in 1983 and may have held some fanihi. 

Some were observed there in 1988 (Wiles et al. 1989). 
Draft Revised Recovery Plan for the Mariana Fruit Bat 

8

no more than 3,000, although the method used to make this estimate is not known 

(Utzurrum et al. 2004). The abundance of fanihi on the island continued to 

decline during the 1960s and 1970s so that by 1978, fewer than 50 bats were 

believed to remain (Perez 1972; Wiles 1987b). Fanihi numbers increased to an 

estimated 850-1,000 in 1982 through movement from Rota (Wiles 1987b), but 

have been in decline on Guam overall since then. 

During the 1990s, numbers on Guam remained in the range of several 

hundred animals with occasional spikes to nearly a thousand bats; 

presumably these spikes reflected temporary immigration events from 

Rota (Wiles et al. 1995; Wiles 1996, 1998, 1999). Surveys conducted 

between 2000 and 2006 indicated further decline to fewer than 100 

individuals (Jeff Quitugua, Guam DAWR, pers. comm. 2006), and at this 

writing the number of fanihi on Guam may have slipped to fewer than 50 

(J. Quitugua, pers. comm., 2009). Predation by brown treesnakes on nonvolant young probably prevents recruitment of juvenile bats on Guam 

(Wiles 1987b, Wiles et al. 1995; Wiles 1996), and losses from illegal 

hunting are also likely to be a factor. 

2. CNMI Southern Islands 

 

Fruit bats on the southern islands of the CNMI (Rota, Tinian, Aguiguan, and 

Saipan) were not surveyed prior to the 1970s, but historical accounts indicate that 

fruit bats once were common on these islands (Fritz 1904). 

Rota

Fanihi from Rota are believed to move episodically among the southern 

islands, and this island thus is considered to be important to the long-term stability 

of the species in the southern part of the Mariana archipelago and to the existence 

of the colony on Guam (Wiles and Glass 1990; Wiles et al. 1995). Fanihi surveys 

have been conducted on Rota regularly by numerous workers since 1986, using 

methods described by Stinson et al. (1992): primarily evening dispersal counts, 

with some station counts of solitary or extracolonial bats and direct counts of 

colonial roosts (Glass and Taisacan 1988; Stinson et al. 1992; Worthington and 

Taisacan 1995, 1996; Johnson 2001; Esselstyn et al. 2006). This monitoring 

effort has yielded numbers that vary widely both intra- and interannually (e.g., 
Draft Revised Recovery Plan for the Mariana Fruit Bat 

9

Glass and Taisacan 1988; Worthington and Taisacan 1995, 1996; CNMI-DFW 

unpublished data, 2007). Owing to variation in methods, observers, and coverage, 

a significant trend cannot be extracted from these data. However, visual 

inspection of the data suggests a decline over the past seven years (CNMI-DFW, 

unpublished data 2007). 

Fanihi numbers on Rota and perhaps consistency in survey methods and effort 

have fluctuated sharply through time (CNMI-DFW unpublished data, 2007); in 

2008 methods were modified in an effort to yield improved information about the 

status of this subpopulation. Island-wide estimates from 1999 to 2008 were based 

on departure counts at seven historically occupied roost sites. However, the 

disturbance of bats caused by illegal hunting results in unpredictable and 

relatively frequent turnover of roost sites used by fruit bats on Rota. In 2008, it 

was determined that colonies were no longer occupying most of the roost sites 

that had been monitored in previous years (Julia Boland, CNMI Department of 

Fish and Wildlife, pers. comm., 2009). Therefore, an intensive search of the 

island for currently occupied roost sites took place, and count and monitoring 

methods were revised. As of 2009, new methods consist of direct counts of 

colonies and of extra-colonial bats at currently occupied roost sites. Colonies are 

observed weekly to monitor possible movements and determine the cause. In the 

event of roost abandonment, colonies are tracked to new locations where counts 

resume. Where possible, pictures of roosting colonies are taken and bats are 

counted from the pictures to confirm direct counts made in the field. Data 

generated using revised count methods suggests that approximately 1200-1400 

fruit bats occur on Rota (in early 2009; J. Boland, pers. comm., 2009). We do not 

think this estimate represents an increase in numbers; rather, it reflects new count 

stations and more accurate methods. A reliable estimate of the trend cannot be 

generated now, owing to the short period that the new methods have been in 

effect. 

Severe storms (and associated hunting) at short intervals combined with low 

and fluctuating numbers could erode what resilience exists in this population. 

Fanihi numbers on Rota declined following Typhoon Roy in 1988 from an 

estimated 2,400 animals to just under 1,000 (Stinson et al. 1992). Prior to 

Typhoon Pongsona in 2002, fanihi numbers on had risen to about 1,300-2,000 

bats (Esselstyn et al. 2006). However, in the months following the storm, 
Draft Revised Recovery Plan for the Mariana Fruit Bat 

10

repeated surveys indicated that numbers had again declined sharply to perhaps 

500 bats (Esselstyn et al. 2006). Numbers remained relatively low through April 

2004, when about 700 bats were counted (Esselstyn et al. 2006). It is unlikely 

that the naturally low reproductive rate of this species can sustain the level of 

hunting pressure observed on Rota. 

Saipan

Schnee (1911) reported that fanihi were commonly seen and heard on Saipan, 

where they were heavily hunted by local residents. The Navy restricted civilian 

access to the northern part of Saipan until the early 1970s, effectively providing 

the bats with protected roost sites. Fanihi on Saipan were observed to decline 

rapidly after the U.S. Navy turned over control to the CNMI government, and 

access to the whole island became unrestricted (Wiles et al. 1989). Observations 

made between the late 1970s and 2007 suggested that Saipan harbored a small 

number of bats; typically 50 bats or fewer (Wheeler 1980; Lemke 1984; Glass and 

Taisacan 1988; Wiles et al. 1989; Worthington and Taisacan 1996; Johnson 2001; 

Ann Marshall, U.S. Fish and Wildlife Service, pers. comm. 2007). 

Tinian

Fritz (1901) reported a “large number” of bats on Tinian in 1900. Since the 

late 1970s fruit bats have been seen rarely and only in small numbers, with 

estimates of fewer than 25 animals usually given for the island (Wheeler 1980; 

Glass and Taisacan 1988; Wiles et al. 1990; Marshall et al. 1995; Krueger and 

O’Daniel 1999; Johnson 2001). Observations during the 1990s suggested that the 

presence of bats on Tinian was intermittent, and their numbers were low 

(Worthington and Taisacan 1996). Brief surveys on Tinian conducted in 2001 

found no fruit bats (Johnson 2001), and between 2002 and 2007 fruit bats have 

been observed only once during forest surveys conducted on Tinian each month 

by U.S. Navy biologists (Scott Vogt, U.S. Navy, pers. comm. 2007). 

Aguiguan

Since the early 1980s fewer than 50 fanihi typically have been observed on 

the island, but bats have been observed on most survey trips to the island (Wiles 

et al. 2006). Typically, bats are dispersed and roost alone or in small groups 

(Wiles et al. 2006). Higher estimates of 75 to 300 animals are sometimes made 

(Glass and Taisacan 1988; Wiles et al. 1989; Stinson et al. 1992; Wiles 1996; 
Draft Revised Recovery Plan for the Mariana Fruit Bat 

11

Esselstyn et al. 2006), suggesting arrivals from Rota (Wiles and Glass 1990). 

Bats on Aguiguan are observed anecdotally to be wary of people, suggesting that 

illegal hunting occurs there (A. Brooke, pers. comm. 2007). 

3. CNMI Northern Islands 

 

The 1983 survey of the northern islands resulted in a minimum estimate of 

7,450 fanihi for Anatahan, Sarigan, Guguan, Alamagan, Pagan, Agrihan, 

Asuncion, and Maug (Wiles et al. 1989; see Table 1). Because fanihi are 

gregarious and often roost in large colonies during the day, this and subsequent 

surveys focused on locating colonies. Wiles et al. (1989) located colonies by 

circumnavigating islands by boat, traversing portions of each island on foot, and 

interviewing residents on islands with human inhabitants. Count methods used 

included evening dispersal counts at colonies and evening counts of solitary or 

extra-colonial bats made from vantage points determined to overlap least with the 

apparent dispersal trajectory of colony bats. Island-wide estimates were based on 

the number of fruit bats recorded, island size, extent of forest cover and 

abundance and diversity of food-plant species (Wiles et al. 1989). 

Surveys of the northern islands undertaken in 2000 (Cruz et al. 2000a-f) 

employed a combination of the same methods used by Wiles et al. (1989) in 1983 

and, on Anatahan, by Worthington et al. (2001) in 1995: land- and sea-based 

colony searches, evening dispersal counts, station-counts of extra-colonial bats, 

and direct day-time counts at roosts. On each island they visited, Cruz et al. 

(2000a-f) spent periods conducting fruit bat surveys equal to or greater than 

periods spent by Wiles et al. (1989) on the same six islands. The individual 

island-wide estimates of Cruz et al. (2000a-f) thus are comparable to those of 

Wiles et al. (1989), but owing to logistical and fiscal constraints, Cruz et al. 

(2000a-f) did not visit Asuncion and Maug. The 2000 surveys yielded an estimate 

of 4,250 fruit bats for the 6 northern islands they visited (Cruz et al. 2000a-f). 

The 1983 surveys yielded an estimate of 7,025 fruit bats for the same six islands 

(Wiles et al. 1989). A conservative interpretation of these data indicates roughly 

a 40 percent decline in fruit bat numbers between 1983 and 2000 among these six 

northern islands. 
Draft Revised Recovery Plan for the Mariana Fruit Bat 

12

The majority of this decline was recorded on two of the three largest northern 

islands, Anatahan (32.3 square kilometers [12.5 square miles]) and Pagan (47.7 

square kilometers [18.4 square miles]), which together harbored roughly 70 

percent of the archipelago’s fruit bats in the 1980s (Wiles et al. 1989). These two 

islands, which were estimated to support a total of 5,500 fruit bats in 1983, were 

estimated to have only 2,500 fruit bats in 2000, approximately a 45 percent 

decline since 1983 (Cruz et al. 2000d, 2000e). Declines on both islands may be 

related to hunting and severe habitat damage caused by ungulates (Cruz et al. 

2000d, 2000e; Kessler 2000a). 

Anatahan

On Anatahan, surveys suggested a minimum estimate of 3,000 fanihi in 1983 

(Wiles et al. 1989), about 1,902 to 2,136 individuals in 1995 (Marshall et al. 

1995; Worthington et al. 2001), and roughly 1,000 individuals in 2000 (Cruz et 

al. 2000d; Kessler 2000a). In conjunction with an ungulate eradication project, 

fanihi on Anatahan have been surveyed by helicopter at least four times each year 

since 2002. These surveys are performed over 2 days with 4 hours spent over the 

island each day. Coverage of the island during each survey is complete. Fanihi 

colonies are rapidly reconnoitered to verify known roost sites and identify new 

ones, colonies are counted and mapped, and individual bats in flight also are 

counted. After the volcanic eruption in May 2003, the island’s devegetated state 

facilitated accurate location of all colonies (C. Kessler, in litt. 2003, pers. comm. 

2004). In 2002 and early 2003, estimates of the island’s bat numbers ranged from 

950 to 1,250 (USFWS unpublished data 2007). Following Anatahan’s volcanic 

eruption in May 2003, aerial surveys conducted in 2003 yielded estimates of 350 

to 700 bats. Four surveys conducted between March and November of 2004 

documented increasing numbers, with a high estimate of 1,570 to 1,805 in 

November (USFWS unpublished data 2007). This localized increase in fruit bat 

numbers over a short period of time (1 to 1.5 years) was concomitant with some 

vegetation recovery and indicates that Anatahan’s fanihi numbers may have been 

approaching their pre-eruption level, whether the source of the additional bats was 

immigration, recruitment of newly volant (flying) young, or both (see Summary 

of Factors Affecting the Species section). However, Anatahan erupted in January, 

April, and August of 2005. Aerial surveys conducted after these eruptions have 

documented new declines, with numbers fluctuating but remaining low. 
Draft Revised Recovery Plan for the Mariana Fruit Bat 

13

Estimates from counts between December 2005 and August 2007 have been 160 

or fewer (USFWS unpublished data, 2007). 

Pagan and Agrihan

On Pagan, fanihi numbers were estimated at a minimum of 2,500 in 1983 

(Wiles et al. 1989), and at roughly 1,500 in 1999 and 2000 (Cruz et al. 2000e). A 

rapid reconnoiter of Pagan in August 2007 yielded a very rough estimate of 500 to 

800 bats (C. Kessler, pers. comm. 2007). On the second-largest northern island, 

Agrihan (47.4 square kilometers [18.3 square miles]), surveys have yielded very 

rough estimates of about 1,000 animals in both 1983 and 2000, suggesting that 

abundance may have been stable during that interval (Wiles et al. 1989; Cruz et 

al. 2000f). Pagan and Agrihan harbor feral ungulate populations, and both islands 

have a long history of intermittent human settlements of variable size. Fanihi on 

these islands likely are under pressure at least intermittently from illegal hunting, 

and their forest habitats are suffering degradation by goats and pigs, and, on 

Pagan, cattle. 

Maug, Asuncion, Alamagan, Guguan, and Sarigan

The remaining northern islands that harbor fanihi are all smaller than 13 

square kilometers (5 square miles) (see Table 1), and each has harbored from 100 

to 500 fanihi (Wiles et al. 1989; Cruz et al. 2000a, b, c). Sarigan, the island 

immediately north of Anatahan, has been surveyed the most frequently in recent 

years in conjunction with the ungulate eradication there. Four surveys between 

1983 and 2000 yielded fairly consistent estimates of 125 to 235 bats (Wiles et al. 

1989; Fancy et al. 1999; Wiles and Johnson 2004). Observed numbers increased 

to an estimated 300 to 400 bats in 2001; however, this temporary increase is 

thought to be an anomaly that likely reflects immigration to Sarigan from a 

neighboring island, probably Anatahan, which is 37 kilometers (23 miles) to the 

south (Wiles and Johnson 2004), and numbers have not remained this high. The 

most recent estimate, based on 2007 surveys, was roughly 140 bats (CNMI-DFW, 

unpublished data 2007; C. Kessler, pers. comm. 2007). The potential for increase 

in fruit bat numbers on Sarigan is thought to be limited by the island’s small size 

(4.9 square kilometers (1.9 square miles)) and the small extent of native forest 

(Wiles and Johnson 2004). 
Draft Revised Recovery Plan for the Mariana Fruit Bat 

14

Of the other four islands, Guguan, Asuncion, and Maug are designated for 

conservation, but Guguan and Maug have not been visited by biologists since 

2001 (Asuncion was visited in 2008). Therefore, the current status of fahini on 

these islands is unknown. Alamagan has a small human settlement and 

populations of non-native goats and pigs and thus the likelihood is high that fanihi 

face hunting pressure there and that the island’s forest is degraded by the 

browsing and rooting of ungulates. Fanihi have been elusive during most surveys 

on Alamagan (Wiles et al. 1989; Johnson 2001); it is thought that they roost in the 

crater in the center of the island (G. Wiles, pers. comm. 2007). 

E. General Biology and Ecology 

1. Habitat Use: Roosting and Foraging 

Fanihi inhabit several native forest types, including primary and secondary 

limestone forest, volcanic (or ravine) forest, old coconut plantations, and groves 

of Casuarina equisetifolia (gagu or ironwood) (Glass and Taisacan 1988; Wiles et 

al. 1989, 1995; Worthington et al. 2001; Wiles and Johnson 2004). Most of these 

habitats are dominated by a variety of native trees, with introduced trees present 

in lower abundance. Grasslands with isolated trees are also used by fanihi (Wiles 

and Johnson 2004) and foraging sometimes occurs at farms and suburban 

residential areas with flowering or fruiting trees (G. Wiles, unpubl. data). On 

islands inhabited by humans, bat colonies usually occur in remote sites, especially 

near or along clifflines. Fanihi often prefer to roost in locations with large, 

emergent trees. On Guam, large Ficus spp. were heavily favored as roosting sites 

in the 1980s and early 1990s (Wiles 1987a). After many of these were lost to 

typhoons, roosting shifted to Aglaia mariannensis (mapunao), Macaranga 

thompsonii (pengua), Mammea odorata (chopak), and Neisosperma oppositifolia

(fagot) (G. Wiles, unpubl. data). Glass and Taisacan (1988) reported Artocarpus

spp. (breadfruit), Elaeocarpus joga (yoga or blue marble tree), Guettarda 

speciosa (beach gardenia or zebrawood), Hernandia labyrinthica (oschal), and 

Macaranga thompsonii as common roost trees on Rota. Colonies have been 

observed in Casuarina equisetifolia, Ficus spp. (fig), and Elaeocarpus joga on 

Anatahan, Sarigan, and Aguiguan, as well as in isolated Cocos nucifera (coconut) 

trees in grasslands on Sarigan (Wiles et al. 1989; Wiles and Johnson 2004; 

Worthington et al. 2001; G. Wiles, unpublished data). 
Draft Revised Recovery Plan for the Mariana Fruit Bat 

15

Fanihi have been observed to feed on the fruits, flowers, and leaves of 39 

plant species, including 11 introduced species. Reported foods include the fruits 

of 29 species of plants, the flowers of 17 species, and the leaves of two species 

(Wiles and Fujita 1992). Favored foods (Table 2) vary somewhat by island. 

Pteropodid bats are an important component of tropical forest ecosystems because 

they disperse plant seeds and thereby help maintain forest diversity and contribute 

to plant regeneration following typhoons and other catastrophic events (Cox et al. 

1992). 

Most of the known fanihi roost sites in the Mariana Islands are located on 

public lands. On Guam, the single remaining colony and probably most foraging 

occur on U.S. military lands (Wiles 1998; Janeke 2006) that are managed as part 

of the Guam National Wildlife Refuge under a cooperative agreement between 

the U.S. Fish and Wildlife Service and the U.S. Air Force. In the CNMI, most 

roosting and foraging areas occur on public lands (belonging to the 

Commonwealth). 

2. Life History 

Fanihi are strongly colonial, forming colonies ranging from a few to as many as 

2,000 animals (Wiles 1987a; Wiles et al. 1989; Worthington and Taisacan 1995). 

Large colonies with more than 1,000 bats occur infrequently. Islands with small 

bat numbers usually feature smaller roosts with fewer than 75 animals (e.g., Wiles 

and Johnson 2004). Within colonies, fanihi group themselves into harems (one 

male and two to 15 females) or bachelor groups (predominantly males), or roost 

as single males scattered throughout (Wiles 1987a). On Guam, the average 

estimated sex ratio in a single colony varied from 37.5 to 72.7 males per 100 

females (Wiles 1982). Guam and Rota also harbor a small percentage of noncolonial animals that roost solitarily, but on some smaller islands, such 

individuals may comprise as much as half the subpopulation (Wiles 1987a, Wiles 

and Johnson 2004, Janeke 2006). 
Draft Revised Recovery Plan for the Mariana Fruit Bat 

16

Table 2. Plant parts commonly eaten by fanihi on different islands in the 

Marianas (Wiles 1987a, Glass and Taisacan 1988, Worthington et al. 2001, 

Wiles and Johnson 2004, Janeke 2006). Fl = flowers, fr = fruits, lv = leaves. 

Species Parts 

Eaten 

Guam Rota Aguiguan Northern 

Islands 

Artocarpus mariannensis fr, lv x x 

Artocarpus altilis* fr x x x 

Ceiba pentandra* fl x x 

Cestrum diurnum* fr x 

Cocos nucifera fl, sap x x 

Cycas micronesica fr x 

Elaeocarpus joga fr, fl x 

Erythrina variegata fl x x x 

Ficus spp. fr x x x 

Guettarda speciosa fl x 

Mammea odorata fr, fl x 

Mucuna gigantea fl x 

Neisosperma oppositifolia fr x x 

Ochrosia mariannensis fr x 

Pandanus tectorius fr x x 

Persea americana* fr, fl x 

Premna obtusifolia fr x 

Terminalia catappa fr, fl x x 

* = introduced species
Draft Revised Recovery Plan for the Mariana Fruit Bat 

17

Reproduction occurs throughout the year in Pteropus mariannus mariannus 

on Guam (Wiles 1987a) and in the subspecies P. m. yapensis on Yap (Falanruw 

1988). Mating and the presence of nursing young have been observed year-round 

on Guam with no consistent annual peak in births (Perez 1972; Wiles 1987a). 

Glass and Taisacan (1988) suggested a similar pattern on Rota, but also indicated 

that a peak birthing season may occur during May and June, as has been observed 

in other fruit bats (Pierson and Rainey 1992). Female bats of the family 

Pteropodidae generally have one offspring per year (Pierson and Rainey 1992), 

and observations on Guam between July 1982 and May 1985 documented a total 

of 262 female bats, each with a single young (USFWS 1990). Based on these 

reproductive traits, several authors have suggested that Pteropus bats have a low 

maximum population growth rate and thus a slow rate of recovery when a 

population is diminished (Pierson and Rainey 1992; McIlwee and Martin 2002). 

However, some Pacific island populations of Pteropus have recovered fairly 

quickly in response to reduced hunting pressure (e.g., P. mariannus pelewensis in 

Palau [Wiles et al. 1997], P. m. yapensis in Yap [Mickleburgh et al. 1992], and P. 

tonganus in American Samoa [Brooke et al. 2000; Utzurrum et al. 2003]); thus, 

reproductive rates may be higher than believed in some populations (Wiles and 

Brooke in press). 

Length of gestation, age of sexual maturity, and average lifespan are unknown 

for the fanihi, but other related bats have a gestation period of approximately 4.6 

to 6.3 months (Pierson and Rainey 1992). Many Pteropus species typically do 

not give birth before 18 months of age (Pierson and Rainey 1992; McIlwee and 

Martin 2002). Mean longevity of a substantially larger species, P. alecto, in 

Australia is four to five years, with a maximum of eight years (Vardon and 

Tidemann 2000). 

3. Interisland Movement 

Pteropus bats are well known to be strong fliers and traverse long distances 

(Eby 1991; Palmer and Woinarski 1999; Nelson 2003), and significant evidence 

exists that fanihi fly between islands in the archipelago (Lemke 1986; U.S. Fish 

and Wildlife Service 1990; Wiles and Glass 1990; Worthington and Taisacan 
Draft Revised Recovery Plan for the Mariana Fruit Bat 

18

1996). The geography of the archipelago, as well as the flight capability of fruit 

bats, facilitates interisland exchange. Distances between islands in the Mariana 

archipelago generally range from 29 to 100 kilometers (18 to 62 miles), and a few 

islands in the southern part of the chain are much closer (e.g., Aguiguan to Tinian 

is 9 kilometers (5.6 miles); Tinian to Saipan is 5 kilometers (3 miles). Each island 

in the chain is visible from neighboring islands (Wiles and Glass 1990). 

Temporary spikes in fruit bat numbers on Guam were observed in 1992-1993 

(from about 350 to 550) and in 1998 (from about 150 to 760 bats) (A. Brooke, in 

litt. 2003). Each of these spikes lasted for several months, with the likely 

explanation being a temporary relocation of bats from Rota, which lies 77 

kilometers (48 miles) from Guam and is visible from Guam’s north shore. More 

modest but equally sudden increases in bat numbers on Guam were noted 2 and 4 

days following typhoons Chataan and Pongsona respectively, in 2002 (D. Janeke, 

University of Guam, in litt. 2003). Several other instances of apparent 

immigrations from Rota to Guam documented in the late 1970s and 1980s are 

described in detail by Wiles and Glass (1990). The presence of fruit bats on the 

islands of Tinian and Aguiguan, which are close to one another and to Saipan, is 

ephemeral (Worthington and Taisacan 1996), indicating that interisland travel 

likely occurs among these three islands as well. 

In the northern islands of the CNMI, fanihi surveys on Sarigan documented a 

roughly stable level of approximately 125 to 235 bats between 1983 and 2000 

(Wiles et al. 1989; Fancy et al. 1999; Wiles and Johnson 2004), and a temporary 

increase to 300 to 400 bats in 2001 (Wiles and Johnson 2004). Recruitment of 

juvenile bats alone cannot account for this increase, and Wiles and Johnson 

(2004) posited Anatahan, 37 kilometers (23 miles) to the south, as the likely 

source for immigrants. Wiles et al. (1989) twice observed individual fruit bats 2 

kilometers (0.8 mile) from Guguan, flying south in the direction of Sarigan, which 

lies 63 kilometers (39 miles) away. Anecdotal observations of likely transits 

among other northern islands are described in Wiles and Glass (1990) and by 

other species experts (Worthington and Taisacan 1996; Wiles and Johnson 2004). 

Typically, observations of vertebrates flying between islands over tens of 

miles of open ocean are extremely rare. In the wider context of Pacific 

biogeography, the evidence described above of interisland movement of fanihi 
Draft Revised Recovery Plan for the Mariana Fruit Bat 

19

within the Mariana Islands is extraordinary. Immigration rates of perhaps one 

individual per generation might be necessary for an island subpopulation to 

maintain genetic homogeneity with the populations on other islands (Mills and 

Allendorf 1996; Wang 2004). The chances of witnessing such a low rate of 

immigration are slight; the evidence described above, therefore, suggests even 

greater rates of movement and probable gene flow among fruit bats on various 

islands in the Mariana archipelago. 

F. Reasons for Decline and Current Threats 

The initial declines in the Mariana fruit bat and their likely causes are outlined 

in the previous section for each island. Table 3 summarizes by island our 

understanding of these historical mechanisms and the current threats to the 

continued existence of the fanihi. Today, illegal hunting, loss of native forest, 

predation by the brown treesnake (on Guam and possibly on Saipan), and the 

increased risk of extirpation or extinction faced by small, fragmented populations 

are the most significant threats to the survival of the Mariana fruit bat. These 

current known and potential threats are discussed below. 

1. Hunting [Listing Factor B] 

A long history of hunting has contributed to archipelago-wide declines in the 

numbers of the Mariana fruit bat and its near-extirpation from all of the southern 

islands except Rota, where illegal hunting continues but bats persist in low 

numbers. Hunting also threatens bats on some of the northern islands, although 

bat numbers and the history of human settlement throughout the archipelago 

suggest that hunting pressure north of Farallon de Medinilla probably has not 

been as severe as in the southern islands (Wiles et al. 1989). Hunting of Mariana 

fruit bats has been restricted or prohibited by Territorial, Commonwealth, and/or 

Federal law since the 1970s. Nonetheless, evidence of chronic hunting is 

consistent, plentiful, and reliable; that illegal hunting of bats is ongoing and 

widespread is a generally accepted if not widely discussed reality. 

Mariana fruit bats have been used as food since humans first arrived on the 

islands (Lemke 1992a), and consumption of bats represents a significant 

Chamorro cultural tradition. Social events and cultural status in the Mariana 

Islands are often enhanced by a variety of foods, and the fruit bat is a highly 
Draft Revised Recovery Plan for the Mariana Fruit Bat 

20

Table 3. Island summary of historical factors and current threats affecting 

the fanihi. See text for full discussion. 

Island Area 

km2

(mi2

) 

Historical Factors Known and 

Potential Current 

Threats 

Known 

(K) or 

Potential 

(P) 

Threat 

Listing 

Factor* 

Fruit bat 

numbers and 

status 

(current 

estimates**, 

year) 

Guam 549.0 

(212.0) 

Habitat loss 

• development 

• agriculture 

• ungulates 

• invasive nonnative plants 

Hunting*** 

Brown treesnakes 

Habitat loss 

• degradation of 

native forests 

• ungulates 

• invasive nonnative 

invertebrates 

• invasive alien 

plants 

• development 

Hunting*** 

Brown treesnakes 

 

K 

K 

K 

K 

K 

K 

K 

A 

B 

C 

<100; 

declining, 

20071 

Rota 85.2 

(32.9) 

Habitat loss 

• development 

• agriculture 

• ungulates 

Hunting 

Habitat loss 

• development 

• ungulates 

Hunting 

K 

K 

K 

A 

B 

500; 

declining, 

20072 

Aguiguan 7.0 

(2.7) 

Habitat loss 

• agriculture 

• ungulates 

Hunting 

Habitat loss 

• ungulates 

Hunting 

Military training 

activities 

K 

K 

P 

A 

B 

E 

<50; 

fluctuating, 

20053

Tinian 101.8 

(39.3) 

Habitat loss 

• development 

• agriculture 

• invasive nonnative plants 

• ungulates 

Hunting 

Habitat loss 

• development 

• invasive nonnative plants 

• ungulates 

Hunting 

Military training 

activities 

K 

K 

K 

K

P 

A 

B 

E 

<10; no 

known 

colonies; 

rarely 

observed, 

20074
Draft Revised Recovery Plan for the Mariana Fruit Bat 

21

Island Area 

km2

(mi2

) 

Historical Factors Known and 

Potential Current 

Threats 

Known 

(K) or 

Potential 

(P) 

Threat 

Listing 

Factor* 

Fruit bat 

numbers and 

status 

(current 

estimates**, 

year) 

Saipan 

122.9 

(47.5) 

Habitat loss 

• development 

• agriculture 

• invasive nonnative plants 

Hunting 

Habitat loss 

• development 

• invasive nonnative plants 

 Hunting 

Brown treesnakes 

K 

K 

K 

P 

A 

B 

C 

Occasional, 

small 

colonies; 30-

50 

Farallon 

de 

Medinilla 

2.0 

(0.8) 

Military training 

activities 

(disturbance and 

habitat loss) 

Military training 

activities 

(disturbance and 

habitat loss) 

K E No colonies; 

2 fruit bats 

observed in 

19966

; 

Anatahan 32.3 

(12.5) 

Habitat loss 

• agriculture 

• ungulates 

Hunting 

Habitat loss 

• ungulates 

• invasive nonnative plants 

Hunting 

Volcanic activity 

K 

P 

P 

K 

A 

B 

E 

220-330; 

decline 

between 

1983 and 

2000, 

20097,9] 

Sarigan 5.0 

(1.9) 

Habitat loss 

• agriculture 

• ungulates 

Hunting 

Habitat loss 

• invasive nonnative plants 

Hunting 

K 

P 

A 

B 

140; stable, 

20077,8

Guguan 4.0 

(1.5) 

Volcanic activity Hunting P B 350; status 

unknown, 

20009 

Alamagan 11.0 

(4.3) 

Habitat loss 

• agriculture 

• ungulates 

Hunting 

Habitat loss 

• ungulates 

Hunting 

Volcanic activity 

K 

K 

P 

A 

B 

E 

200; numbers 

and status 

uncertain9, 10

Pagan 47.7 

(18.4) 

Habitat loss 

• agriculture 

• ungulates 

Hunting 

Volcanic activity 

Habitat loss 

• ungulates 

• invasive nonnative plants 

Hunting 

Military training 

activities 

Volcanic activity 

K 

K 

P 

P 

A 

B 

E 

E 

500-800; 

decline 

between 1983 

and 2000, 

recent 

numbers 

uncertain, 

20077,9 
Draft Revised Recovery Plan for the Mariana Fruit Bat 

22

Island Area 

km2

(mi2

) 

Historical Factors Known and 

Potential Current 

Threats 

Known 

(K) or 

Potential 

(P) 

Threat 

Listing 

Factor* 

Fruit bat 

numbers and 

status 

(current 

estimates**, 

year) 

Agrihan 47.4 

(18.3) 

Unknown Habitat loss 

• ungulates 

Hunting 

Volcanic activity 

K 

K 

P 

A 

B 

E 

1,000; status 

unknown, 

20009

 

Asuncion 7.4 

(2.9) 

Volcanic activity Volcanic activity 

Hunting 

P 

P 

E 600; status 

unknown, 

200111 

Maug 2.0 

(0.8) 

Unknown Unknown 

Hunting P 

 <25, status 

unknown, 

198310

* Listing Factors (as described in Section 4(a)(1) of the Endangered Species Act: 

A – Destruction or curtailment of habitat or range 

B – Overutilization for commerce, recreation, education, or scientific purposes 

C – Disease or predation 

D – Inadequacy of existing laws 

E – Other natural or human-related factors 

** Note: most estimates are rough and represent a range of methods and survey effort and 

frequency. 

*** Based on anecdotal information from local hunters, it is believed that hunting of fruit bats 

once occurred throughout the Marianas. Today, hunting of fruit bats is illegal under Territorial, 

Commonwealth, and Federal law. 

Sources: 

1

 A. Brooke, pers. comm.. 2007 

2 C. Kessler, pers. comm. 2007

3

 G. Wiles, pers. comm. 2007 

4 S. Vogt, pers. comm.. 2007 

5 Laura Williams, CNMI-DFW, pers. comm. 2004 

6 Tim Sutterfield, U.S. Navy, in litt. 1997 7 C. Kessler, pers. comm. 2009 (estimate for Pagan based on brief 2007 survey) 

8 Wiles and Johnson 2004 

9 Cruz et al. 2000f (Agrihan); 2000e (Pagan); 2000b (Alamagan), 2000a (Guguan) 10 Wiles et al. 1989 11 Johnson 2001 
Draft Revised Recovery Plan for the Mariana Fruit Bat 

23

prized delicacy. In a Guam survey of Chamorros, the culture indigenous to the 

Mariana Islands, 53 percent of the respondents indicated that they enjoyed eating 

fruit bats (Sheeline 1991). The Mariana fruit bat clearly is an important symbol in 

the Mariana Islands, as 82 percent of the respondents to the same survey believed 

that fruit bats had cultural value. However, 85 percent of the respondents also 

expressed a belief that people should stop hunting and eating fruit bats if such 

activity would lead to the species’ extinction (Sheeline 1991). 

Traditional hunting methods, such as nets, traps, thorny branches on poles, 

and stone projectiles, ensured limited success in capturing fruit bats (Lemke 

1992a). Today, bats are taken more efficiently and in greater numbers with 

shotguns and rifles fired at roosting and feeding sites or along flyways. 

Gregarious species such as the fanihi are particularly vulnerable to hunting at their 

roost sites. One shotgun blast may kill several bats or knock them to the ground, 

and a successful raid can glean up to 50 bats (Wiles 1987b; Lemke 1992a). 

Hunting at colonies can also result in direct mortality and abandonment of infant 

and juvenile bats (Lemke 1992a). 

Demand for fruit bats for human consumption in the Marianas is clearly 

demonstrated by the large commercial trade in bats that existed from the 1960s to 

1990s (Wiles and Payne 1986; Stinson et al. 1992; Wiles 1992; Wiles et al.

1997). It is estimated that several hundred thousand fruit bats were imported to 

Guam and the CNMI from other sources in the Pacific region during this period. 

This number included about 16,500 Mariana fruit bats shipped to Guam from 

Rota, Saipan, Tinian, and Pagan from 1975 to 1982 (excluding 1977), although 

this figure is likely inflated to some extent. This trade ended in 1994, leaving a 

void in the availability of bats for Chamorro consumers and potentially increasing 

illegal hunting for bats within the island chain. In recent years, a single locally 

killed fruit bat can sell for $40 to $75 in the CNMI (Worthington and Taisacan 

1996; C. Kessler, in litt. 2003; A. Brooke, pers. comm. 2007, Dana Lujan, U.S. 

Department of Agriculture, pers. comm. 2007). Despite this, many of the bats 

killed are taken for personal consumption (often for important social and cultural 

occasions) rather than for commercial purposes. 

 
Draft Revised Recovery Plan for the Mariana Fruit Bat 

24

Southern Islands

Hunting was one of the important causal factors in the fruit bat declines on 

Guam, Saipan, Tinian, and Aguiguan through the 1970s (Perez 1972; Wheeler 

1980; Wiles 1987b), although bans on the hunting of bats were enacted on Guam 

in 1973 and in the CNMI in 1977 (Public Law 5-21, September 1977). Today, 

although few bats remain on Guam, they are probably still hunted 

opportunistically on private property when they transit the island, and by deer 

hunters or poachers on Andersen Air Force Base and other military lands on 

northern Guam near the last remaining colonial roost (Guam National Wildlife 

Refuge staff, pers. comm. 2005). Numerous documented reports indicate that 

illegal hunting continues to be a major threat to the Mariana fruit bat in the 

CNMI, particularly on Rota (see below). In 1987, between three and eight bats 

were reported to be taken by hunters in a small colony on Saipan (Glass and 

Taisacan 1988). In 1997, there was a report of nearly 90 bats shot and killed on 

Tinian from a colony that roosted on the island briefly (T. Sutterfield, pers. 

comm. 1998). 

Rota

Numerous documented reports indicate that hunting continues to be a major 

threat to the Mariana fruit bat on Rota (Glass and Taisacan 1988; Lemke 1992b; 

Marshall et al. 1995; Worthington and Taisacan 1996; Stan Taisacan, CNMIDFW, pers. comm. 1997; Rainey 1998; Nathan Johnson, CNMI-DFW, pers. 

comm. 2000; C. Kessler, pers. comm. 2004; Arlene Pangelinan, U.S. Fish and 

Wildlife Service, pers. comm. 2004; Esselstyn et al. 2006, J. Boland, pers. 

comm., 2009). This long history of observations by biologists on Rota indicates 

that hunting is ongoing in spite of the law. Although hunting pressure is difficult 

to quantify, an interview of 29 hunters suggested an annual mean hunting rate of 

9.4 bats/hunter during normal years (e.g., those without major storms) (Esselstyn 

et al. 2006). 

Most recently, CNMI-DFW biologists have made the following observations 

(J. Boland, pers. comm., 2009): between April 2008 and April 2009 four 

maternity colonies and an unknown number of commuting and foraging 

individuals were disturbed by poachers. Disturbances have resulted in complete 

abandonment of three roost sites occupied by maternity colonies and partial 

abandonment of the fourth. Disturbances to bats while they commute between 
Draft Revised Recovery Plan for the Mariana Fruit Bat 

25

roosts and foraging areas have resulted in bats changing preferred flight paths. 

Alternate flight paths may put bats at greater risk of death and injury due to an 

increase in energy expenditure required to travel longer distances over open water 

with greater exposure to strong winds and rain. Poaching incidents since April 

2008 have resulted in the loss of an estimated 200 bats. 

Post-typhoon Hunting on Rota 

Defoliation and other habitat damage caused by severe storms can increase the 

vulnerability of fruit bats to illegal hunting (see the “Natural Disturbance” section, 

below). Following supertyphoon Roy in 1988, the near-total defoliation of the 

island’s forests (Fancy and Snetsinger 1996) and other damage caused by the 

storm forced bats on Rota to forage during the day in areas close to human 

habitation (Stinson et al. 1992). As a result, extensive opportunistic hunting 

occurred, contributing to a reduction of total bat numbers on Rota by more than 

half (Stinson et al. 1992). Bat numbers recovered to more than 2,000 before 

supertyphoon Pongsona in December 2002, but again declined by more than half 

following this storm, most probably with hunting as a significant contributing 

factor. This decline was documented by monthly surveys conducted by the same 

individuals using the same techniques (evening colony departures, direct colony 

counts, and searches for solitary bats). These surveys yielded estimates of fewer 

than 750 animals for most of the 15 months following the supertyphoon 

(Esselstyn et al. 2006). Similar sharp increases in hunting of fruit bats following 

severe storms have been documented elsewhere in the Pacific as well as in the 

Mariana Islands (Craig et al. 1994; McConkey et al. 2004). 

Continued illegal hunting on Rota diminishes the fruit bat population’s ability 

to recover to pre-storm abundance (Worthington and Taisacan 1996). Hunter 

interviews indicated that hunting pressure on fruit bats increased by roughly 30 

percent in the year following Pongsona (Esselstyn et al. 2006). In June and July 

2004, residents of Rota reported that one or more illegal hunting incidents killed 

at least 40 fruit bats, resulting in the abandonment of the largest colony on the 

island, and another smaller colony was abandoned at this time as well (C. Kessler, 

pers. comm. 2004). On August 22 and 23, 2004, 21 months after supertyphoon 

Pongsona, Rota again sustained severe damage from supertyphoon Chaba, with as 

much as 60 to 75 percent of the island defoliated (A. Pangelinan, pers. comm. 

2004). These conditions may have facilitated another pulse of post-typhoon 
Draft Revised Recovery Plan for the Mariana Fruit Bat 

26

hunting. Hypothetically, multiple episodes of increased illegal hunting would 

exacerbate the adverse effects of severe storms on bat numbers and impede the 

natural rebound of the population (see Factor E, below). 

CNMI Northern Islands

Hunting of fruit bats on the northern islands is occasionally reported, but 

monitoring on these islands and of vessels returning from the north is limited. It 

is generally assumed that illegal hunting of fanihi takes place on all islands with 

human populations and on islands visited in transit to and from these (Wiles et al. 

1989; L. Williams, pers. comm. 2007; G. Wiles pers. comm. 2007). In 1996, 

hunting in the northern islands was reported to be an increasingly significant 

problem (Worthington and Taisacan 1996). On Anatahan, which lies only 151 

kilometers (94 miles) from heavily-populated Saipan, remains of recently cooked 

fruit bats were found in the main campsite area in 1995, and a team of biologists 

on the island observed residents of Anatahan cooking and eating fruit bats 

(Marshall et al. 1995; Worthington et al. 2001). 

In 1998, 14 Mariana fruit bats shot in the northern islands were confiscated 

from a CNMI vessel returning to Saipan (T. Eckhardt, in litt. 1998), and hunting 

of Mariana fruit bats was reported on Sarigan (Zoology Unlimited LLC 1998). 

On Pagan, 7 recently expended .410 (very small bore) shotgun shells were found 

in 1999, 4 more were found in 2000, and a .410 shell and fresh remains of cooked 

fruit bat were found during a helicopter refueling stop in 2001 (Cruz et al. 2000e; 

Johnson 2001). Most recently, a team of CNMI officials visiting Pagan in 2007 

reported being served cooked fanihi by the island’s residents (L. Williams, pers. 

comm. 2007). 

2. Habitat Loss and Degradation [Listing Factor A] 

The ecological integrity of the remaining native forest habitat in the Mariana 

Islands is threatened by development, agriculture, and the long-term ecological 

impacts of ungulates, invasive plants, and, on Guam, the brown treesnake. 

Although some non-native forest types (e.g., forest dominated by coconut or other 

agricultural plants) do provide food resources for fruit bats, and bats roost in some 

forests that provide little or no forage (e.g., Casuarina forest), loss and 

degradation of native forest is a key threat to Mariana fruit bats because essential 
Draft Revised Recovery Plan for the Mariana Fruit Bat 

27

food and roosting resources are lost. For example, essential, limiting nutrients 

such as calcium are more abundant in native plants than in agricultural plants 

eaten by fruit bats in American Samoa (Nelson et al. 2000), and in the 

Philippines, fruit bats preferentially forage in native forest although they use both 

native and non-native/disturbed forests (Mildenstein et al. 2005). 

Over the past several centuries, the southern Marianas have lost most of their 

original native forest, primarily to agriculture, growing human populations, 

economic development, and military activities (Bowers 1950; Fosberg 1960). 

Nonetheless, the overall extent of native forest and potential habitat for fanihi is 

greatest in the southern islands (Table 4), although these forests may be degraded 

(see below). Only a few of the northern islands are relatively unaltered by human 

activities; Guguan, Asuncion, and possibly Maug have for the most part escaped 

the effects of millennia of continuous human settlement, World War II, and postwar activities that caused the initial, extensive habitat alteration and fragmentation 

of forest habitat (see Table 4). Human impacts are evident on the other islands, in 

the form of swordgrass savannahs maintained by burning, old copra (coconut) 

plantations, the presence of feral ungulates, and intermittent human habitation. 

The absolute amount of forest habitat in the northern islands is much less than in 

the southern islands, in part because the islands are much smaller. In addition, the 

relative proportions of northern islands considered to be habitat for fanihi also are 

smaller than in the southern islands (see Table 4). 

On many islands in the archipelago, ungulates have caused severe damage to 

forests by browsing on plants, causing erosion (Marshall et al. 1995; Kessler 

1997; U.S. Fish and Wildlife Service 1998a, b), retarding plant growth and 

regeneration (Lemke 1992b; Wiles 2005), and facilitating the establishment of 

invasive plants (Marshall et al. 1995; Kessler 1997; U.S. Fish and Wildlife 

Service 1998a, b; Wiles et al. 1999), which can impede forest regeneration by 

displacing or smothering native species (Kessler 2000b). This is a particular 

problem on highly altered islands, where what little mature native forest remains 

continues to be threatened by the degradation and fragmentation associated with 

introduced ungulates. For example, although Guam is dominated by native forest 

(see Table 4), Wiles (2005) recently documented a rapid decline in the native 

breadfruit, an important bat food, even in the remaining native forest on the 
Draft Revised Recovery Plan for the Mariana Fruit Bat 

28

Table 4. Estimated extent of fanihi habitat in the Marianas. “Native” 

includes mature and secondary forest, but these forests may be degraded by 

ungulates. The variety and extent of forest types provide habitat of variable 

quality; see text for discussion. Data compiled by USFWS from published 

and unpublished sources. 

Island Area 

km2

 (mi2) 

Total forest 

cover 

ha 

Fanihi habitat 

ha 

Fanihi 

habitat 

(percentage 

of all forest 

cover) 

Fanihi 

habitat 

(percentage 

of total land 

area) 

Guam 549.0(212.0) 25,833 25,711 

Native: 25,711 

100 48 

Rota 85.2(32.9) 6,977 6,663 

Native: 6,202 

Coconut: 446 

Agroforest: 15 

95 70 

Aguiguan 7.0(2.7) 302 302 

Native: 302 

100 43 

Tinian 101.8(39.3) 8,283 6,481 

Native: 6,361 

Coconut: 118 

Agroforest: 2 

78 64 

Saipan 122.9(47.5) 9,105 5,355 

Native: 4,080 

Coconut: 1,236 

Agroforest: 39 

59 44 

Farallon de 

Medinilla 

2.0(0.8) 0 0 0 0 

Anatahan 32.3(12.5) Not available Not available 

Sarigan 5.0(1.9) 162 162 

Native: 29 

Coconut: 133 

100 33 
Draft Revised Recovery Plan for the Mariana Fruit Bat 

29

Island Area 

km2

 (mi2) 

Total forest 

cover 

ha 

Fanihi habitat 

ha 

Fanihi 

habitat 

(percentage 

of all forest 

cover) 

Fanihi 

habitat 

(percentage 

of total land 

area) 

Guguan 4.0(1.5) 140 140 

Native: 140 

100 35 

Alamagan 11.0(4.3) 350 350 

Native: 230 

Coconut: 120 

100 32 

Pagan 47.7(18.4) 1,720 900 

Mixed native and 

coconut: 900 

52 19 

Agrihan 47.4(18.3) 2,300 2,300 

Native: 1,250 

Coconut: 800 

Agroforest: 250 

100 58 

Asuncion 7.4(2.9) 305 305 

Native: 305 

100 41 

Maug 2.0(0.8) Not available Not available Not 

available* 

Not available 

* Owing to Maug’s remoteness and extreme topography, all of its forest cover is native.

northern end of the island. This decline was attributed to a combination of 

mortality of mature trees in the wake of several severe typhoons and a lack of 

recruitment caused by deer and pig foraging on fallen fruit (with seeds) and 

seedlings. 

Non-native pigs, goats, and cattle continue to degrade native forests on some 

of the northern islands as well. The introduction of these ungulates to some of the 

northern islands as recently as 40 years ago has resulted in rapid degradation and 

loss of native forest cover, notably on Anatahan and Pagan, two of the largest 
Draft Revised Recovery Plan for the Mariana Fruit Bat 

30

islands that have supported relatively large numbers of fruit bats (Kessler 1997, 

2000a; Worthington et al. 2001). 

These vegetation and landscape changes can deprive fruit bats of the forests 

on which they depend for food and roosting. The diminished quality and extent 

of forest thus leads to an associated reduction in the number of fruit bats that the 

remaining habitat is able to support. In the most extreme case, tree loss associated 

with past and current live-fire training activities by the military on Farallon de 

Medinilla has all but eliminated roosting and foraging opportunities for bats 

transiting between the Southern and Northern islands. 

3. Predation by Brown Treesnakes [Listing Factor C] 

The brown treesnake is believed to adversely affect bat recruitment on Guam. 

Numerous sightings of this predator have occurred on Saipan, and treesnake 

interdiction is a critical concern throughout the CNMI. Predation by brown 

treesnakes has caused the decline or extirpation of most landbird species on Guam 

(Savidge 1987, Wiles et al. 2003) and is probably responsible for decreases in 

recruitment of young bats observed in the island’s low remaining numbers of bats 

(Wiles 1987a; Wiles et al. 1995). Although only two or three cases of snake 

predation have been reported (e.g., Wiles 1983), the snake is considered capable 

of preying on non-volant young bats (U.S. Fish and Wildlife Service 1990). Data 

collected at the island’s main bat colony from 1982 to 2006 have indicated a 

consistent pattern wherein small pups are routinely recorded with their mothers, 

but medium-sized young are much rarer and large young are virtually absent 

(Wiles 1987b; Wiles et al. 1995; G. Wiles, unpubl. data; D. Janeke, pers. comm. 

2006). Most observations were made in areas where native forest bird 

populations had collapsed, indicating the establishment of high densities of brown 

treesnakes. However, for several months in 1983, after the colony moved to a 

new location where native birds persisted, sizable numbers of larger pups were 

detected (G. Wiles, pers. comm. 2007). 

The brown treesnake was accidentally introduced to Guam between 1945 and 

1952, probably in ship cargo (Rodda et al. 1992), and was present throughout the 

island by 1986 (Savidge 1987). Because of the snake’s distribution and biology 

and Guam’s location and role as a major transportation hub in the Pacific, the 
Draft Revised Recovery Plan for the Mariana Fruit Bat 

31

probability is high that human conveyance will disperse brown treesnakes from 

Guam to other Pacific islands (Fritts 1988). 

 

Reports of brown treesnakes in the CNMI, especially on Saipan, have 

increased since 1982 (Brown Treesnake Control Plan 1996). As of October 2007, 

76 credible treesnake sightings, including 11 captures, had occurred on the island 

(Nathan Hawley, U.S. Fish and Wildlife Service, pers. comm. 2007). The 

frequency of sightings during this period suggests that brown treesnakes are 

present on Saipan island (Colvin et al. 2005; Draft Brown Treesnake Control Plan 

2009) thus leading to increased predation risks on native fauna. However, 

documentation of treesnakes breeding on Saipan still is lacking (Rodda and 

Savidge 2007; Draft Brown Treesnake Control Plan 2009). There have also been 

two documented captures of treesnakes on Rota and five sightings on Tinian. 

4. Natural Disturbance, Development, and Military Training Activities 

[Listing Factor E] 

Natural Disturbance 

The fanihi has coexisted for millennia with typhoons, volcanic eruptions, and 

other natural phenomena. These disturbances, when sufficiently severe, probably 

have drastically reduced bat numbers at times on all islands in the Marianas. 

With sufficient time and suitable remaining habitat, bat numbers presumably 

rebounded through natural recruitment or immigration from neighboring islands. 

Today, natural disturbances are likely to lead to more severe and lasting 

impacts on the compromised fanihi population throughout the archipelago 

because these impacts are exacerbated by anthropogenic pressures. Individual 

islands harbor smaller numbers of bats than they did historically, and numbers on 

some islands are so small that they are highly vulnerable to extirpation. Natural 

or anthropogenic disturbance and resulting changes to a population’s intrinsic 

demographic parameters, such as sex ratio, age structure, and other traits, are 

more likely to affect the population growth rate and persistence when numbers are 

low. The decreasing numbers and increasingly fragmented distribution of fanihi 

throughout the islands through time make recolonization events less likely after 

disturbance from severe environmental events. The total extent of high quality, 

native forest for bats has been reduced on most islands, which limits opportunities 

for bats to shift the location of their roost sites and foraging activity in response to 
Draft Revised Recovery Plan for the Mariana Fruit Bat 

32

disturbance (and by extension limits the resilience of the ecosystem as a whole). 

Taken together, these circumstances reduce the ability of bats to recover rapidly 

in the wake of severe natural disturbances. 

Storms 

Severe storms, in particular, pose a threat to fanihi, primarily because of the 

dramatic increase in hunting that sometimes occurs in the wake of such storms, 

augmenting the impact of that natural disturbance (Esselstyn et al. 2006). 

Evidence suggests that severe typhoons usually do not directly kill fruit bats at 

their roost sites (Lemke 1992b; Esselstyn et al. 2006; Wiles and Brooke in press; 

but see Wiles 1987b), but the synergistic effect on the fanihi of illegal hunting and 

severe storms is well documented on Rota (see “Hunting” section, above) and in 

other locations (e.g, Craig et al. 1994; McConkey et al. 2004). Severe storms can 

alter fruit bat foraging and roosting behavior by decimating food resources, 

removing protective foliage cover, temporarily modifying forest structure, and 

changing vegetation composition, especially by facilitating encroachment of 

nonnative species (Lemke 1992b). Loss of food resources can drive bats to forage 

closer to areas of human activity in daylight hours, thereby increasing their 

vulnerability to illegal hunting (Craig et al. 1994; Pierson et al. 1996). Finally, 

tropical cyclones are likely to become more severe and more frequent in coming 

decades as a result of human impacts to the global climate (e.g., Emanuel 2005; 

Webster et al. 2005). Environmental changes on this rapid timescale may exceed 

the ability of natural systems to adapt. 

Volcanism 

The Mariana Archipelago was formed by volcanic activity along the Mariana 

Trench, a subduction zone where one tectonic plate of the Earth’s crust is moving 

beneath another. The northern islands of the CNMI are still volcanically active; 

eruptions have altered the landscapes and habitats of many of these islands in 

recorded history. The potential thus exists for volcanic eruptions to destroy 

habitat for fruit bats in the northern islands. Pagan last erupted in 1981, when ash 

and a lava flow covered much of the northern half of the island. Anatahan 

erupted multiple times between 2003 and 2005, and much of the island was 

denuded. As described above, fruit bat numbers on Anatahan declined 

precipitously following the 2003 eruption, and after a brief increase during 2004, 
Draft Revised Recovery Plan for the Mariana Fruit Bat 

33

have remained at fewer than 200 individuals since 2005 (C. Kessler, pers. comm. 

2007). 

Few people have visited Anatahan since the eruptions began, and illegal 

hunting there is thus unlikely to have confounded the response of the island’s bats 

to this natural disturbance. However, owing to anthropogenic disturbance, the 

islands on either side of Anatahan, Sarigan and Farallon de Medinilla, harbor few 

and no bats, respectively (Table 3). Therefore, more time and opportunity will be 

necessary for significant number of bats from other islands to immigrate to 

Anatahan, even when the island’s vegetation has recovered sufficiently to support 

greater numbers of bats. An increase in bat numbers on Anatahan may be more 

dependent on intrinsic growth than on immigration for some time to come. 

Development and Military Training Activities 

Urbanization and other forms of development remain a threat to the Mariana 

fruit bat. This threat may manifest as fragmentation or degradation of forest 

habitat, direct disturbance of bats, and/or increased likelihood of new 

introductions of the brown treesnake or other predators to islands. On Guam, 

development takes the form of urbanization associated with an increasing 

population and tourism industry and the expansion and refurbishment of military 

infrastructure. On Rota and Tinian, development includes the clearing of lands 

set aside as agricultural homesteads (CNMI Senate Bill 13-32, C.S. 1, November 

2002; CNMI Senate Bill 14-44, S.S. 1, July 2004), military infrastructure and new 

businesses such as the casino on Tinian. On Saipan, increasing urbanization, road 

building, and the tangantangan charcoal industry are ongoing issues of concern. 

The Department of Defense has several military installations and training 

programs in the Mariana Islands. The Department of Defense live fire and 

bombing exercises on Farallon de Medinilla have effectively precluded that island 

as a foraging or roosting site for fruit bats. However, survey crews in 1996 and 

2008 each observed a single bat on the island, indicating that Farallon de 

Medinilla may still function as a stopover site for bats in transit (A. Brooke, pers. 

comm., 2009). Recent and new activities proposed by the U.S. Air Force at 

Andersen Air Force Base on northern Guam have been determined likely to 

adversely affect fruit bats under section 7 of the Endangered Species Act, 
Draft Revised Recovery Plan for the Mariana Fruit Bat 

34

prompting formal consultation with the U.S. Fish and Wildlife Service to estimate 

the risk of take and develop measures to avoid and minimize that take. 

As of this writing, the Department of Defense is developing Environmental 

Impact Statements for new training, development, and other activities on Guam 

and in the CNMI associated with the redeployment of a U.S. Marine Corps 

Expeditionary Force from Okinawa to the Marianas. We do not currently have 

sufficient information to summarize in this draft revised recovery plan the 

potential threat to fanihi posed by these actions and will evaluate these proposed 

activities under section 7 of the Endangered Species Act once the data have been 

provided. 

G. Conservation Efforts 

Prompted by severe declines in fanihi numbers, the CNMI legislature passed a 

moratorium on the taking of bats on all islands in 1977 (Public Law 5-21, 

September 1977); however, hunting of bats was subsequently permitted on 

Anatahan. The bat has since been listed as threatened or endangered by the 

CNMI government on Rota, Saipan, Tinian, and Aguiguan (CNMI 1991), but the 

CNMI’s designation of threatened or endangered species did not include 

prohibition on take (K. Garlick, U.S. Fish and Wildlife Service, in litt. 1997) or 

any other protection (A. Palacios, in litt. 1990; Worthington and Taisacan 1996). 

Current CNMI hunting regulations (Part 4, Section 10.7.i (Commonwealth 

Register Vol. 23, August 16, 2001, p. 18266)) prohibit the hunting, killing, or 

possession of threatened, endangered, and protected species. 

 

The Mariana fruit bat is listed as an endangered species by the Government of 

Guam and take is prohibited under this designation (Wiles 1982). The species 

was originally listed under the Federal Endangered Species Act as an endangered 

subspecies endemic to Guam in 1984 (49 FR 33881). We designated critical 

habitat for the Mariana fruit bat on Guam in a final rule published in the Federal 

Register on October 28, 2004 (USFWS 2004). On Guam, the bat is legally 

protected from hunting by its status under U.S. and Guam laws. 

In January 1990, declines in bat populations throughout Micronesia resulted in 

the reclassification of P. mariannus from Appendix II to Appendix I of the 
Draft Revised Recovery Plan for the Mariana Fruit Bat 

35

Convention on International Trade in Endangered Species (CITES). This 

reclassification was an effort to control shipments of other subspecies from Palau, 

Yap, and Kosrae into the Mariana Islands and to encourage exporting countries to 

conserve their bat populations. Enforcement of CITES regulations, outreach by 

the U.S. Fish and Wildlife Service and the governments of Guam and the CNMI, 

and Palau’s change in political status to one of free association with the United 

States resulted in the termination of all legal Pteropus imports into the Marianas 

in 1994. CITES protection has been highly successful and very few illegal 

shipments of bats now enter Guam and the CNMI (Wiles and Brooke in press). 

Since the 1970s, surveys of fanihi have been conducted on various islands and 

at variable intervals by the Guam Division of Aquatic and Wildlife Resources 

(DAWR) and CNMI DFW. Personnel at these agencies and at the Guam National 

Wildlife Refuge have conducted studies to obtain information about the ecology, 

life history, movements, and take by hunters of fanihi (e.g., Perez 1972; Wheeler 

and Aguon 1978; Wheeler 1980; Wiles 1982, 1983, 1987a, 1987b; Glass and 

Taisacan 1988; Wiles et al. 1989, 1995; Wiles and Glass 1990; Lemke 1992; Rice 

and Taisacan 1992; Stinson et al. 1992; Worthington and Taisacan 1995, 1996; 

Worthington et al. 2001, Wiles and Johnson 2004; Esselstyn et al. 2006). The 

CNMI DFW has a staff biologist position dedicated to study and monitoring of 

fanihi. A University of Guam student recently concluded a study of fanihi 

movements and habitat use on Guam (Janeke 2006). Studies of fruit bats by 

biologists at the agencies listed above, the U.S. Navy, and other institutions are 

ongoing. Genetic analysis of differentiation between island groups is currently 

being conducted by Gary McCracken at the University of Tennessee. Wiles 

(2005) reported on Guam’s declining population of native breadfruit, a favored 

food of fanihi. 

Other projects in the Mariana Islands may not specifically address bat 

conservation, but indirectly benefit the species. Examples of such projects 

include: the designation of conservation status for Guguan, Asuncion, and Maug, 

the three northernmost islands of the CNMI; locally legislated conservation lands 

in the CNMI Sabana Conservation Area and on Guam; the establishment of Guam 

National Wildlife Refuge at Ritidian Point and the overlay refuges on most 

military lands on Guam; ungulate eradications on Sarigan and Anatahan; ongoing, 

multi-agency research and management to control brown treesnakes on Guam; 
Draft Revised Recovery Plan for the Mariana Fruit Bat 

36

and research and recovery implementation for other listed species in the Marianas, 

for example the aga or Mariana crow (Corvus kubaryi), the nosa Luta or Rota 

bridled white-eye (Zosterops rotensis), and the ga`ga` karisu or nightingale reedwarbler (Acrocephalus luscina). 

 

Because illegal hunting of fanihi continues, we conclude that to date, 

enforcement of laws that prohibit hunting has been mostly ineffectual. Survey 

efforts and research listed above have contributed information about the threat 

posed by hunting that can aid in recovery, but this information has not yet been 

implemented effectively on the ground in the form of outreach, education, and/or 

punitive consequences for those who break the law. Because habitat degradation 

and loss is a principal threat to the fanihi, projects that protect or restore native 

forest are likely to have high conservation benefits. At this time, few section 7 

consultations have been conducted that have included measures to conserve 

fanihi, and no section 10 incidental take permits have been issued that directly 

address fanihi recovery. However, section 10 habitat conservation planning 

efforts that are currently underway on Rota for the Mariana crow may result in 

mitigation that will benefit other native birds and fanihi as well, by protecting 

native forest habitat on Rota. 
Draft Revised Recovery Plan for the Mariana Fruit Bat 

37

II. RECOVERY 

A. Recovery Goal and Objective 

The goal of this recovery plan is to bring about the recovery of the fanihi and 

delist the species in the Mariana Islands. The major objective of this recovery 

plan is to outline a strategy and describe actions that will result in increasing 

numbers of fanihi and the establishment of a viable metapopulation in the 

Mariana Islands. To do this, one aim is to strengthen individual island 

subpopulations across the archipelago as well as increase potential for interaction 

among island subpopulations within the archipelago. Such interactions among 

multiple islands provide insurance against a catastrophe such as a severe typhoon 

or volcanic eruption that may decimate bat numbers on one or more islands. A 

viable, archipelago-wide population is one that is resilient to natural disturbances 

and has a probability of extinction that is ten percent or less over 100 years. 

Meeting this overarching objective will require a combination of extensive 

community outreach and law enforcement to curb illegal hunting and disturbance 

of bats at their roosts, habitat restoration projects, and ongoing research and 

monitoring. Background for and description of these actions are provided in parts 

I and III, respectively, of this plan. 

B. Recovery Strategy 

The strategy for achieving the distribution and long-term population stability 

described above will involve several broad categories of actions: unlawful hunting 

of fanihi must be effectively controlled throughout the archipelago; habitat on 

islands where bats occur must be protected and enhanced, and habitat where bats 

do not currently or regularly occur must be protected or restored to facilitate 

movement of bats among islands and address existing or incipient disjunctions in 

the fanihi’s distribution; and brown treesnakes must be controlled on Guam and 

Saipan, and most importantly, must be prevented from dispersing to other islands. 

Re-establishing and maintaining a widely distributed, robust metapopulation 

of fanihi in the Mariana Islands is critical to the long-term persistence of the 

species. Distribution of bats in both the Northern and Southern islands is 

essential, including relatively large numbers of bats on some islands (e.g., Rota, 
Draft Revised Recovery Plan for the Mariana Fruit Bat 

38

Pagan, Agrihan, and/or Anatahan) and smaller numbers on small islands (e.g., 

Aguiguan, Guguan, and Asuncion) or islands with limited habitat (e.g., Sarigan). 

We anticipate that establishment of bats in currently unoccupied habitat will occur 

by natural colonization from adjacent sites or other islands, once numbers in those 

locations are released from hunting pressure and can grow. Even if numbers on 

individual islands fluctuate, for example, as a result of a volcanic eruption or 

severe typhoon, the distribution of healthy numbers of bats on other islands 

provides insurance against a single environmental catastrophe which otherwise 

may have significant population-level effects on the fanihi and hinder recovery. 

Illegal hunting is a major threat to fanihi, and implementing recovery actions 

(e.g., outreach, education, and enforcement measures) to address this threat will 

be challenging, because the bat has great cultural value as a food item among the 

indigenous people and some immigrant groups in the Marianas. In the near term, 

consistent enforcement of Federal, Territorial, and Commonwealth law that 

prohibits hunting these animals is essential to ensure their continued existence. 

Immediately and in the long term, however, our efforts must focus on raising 

awareness of the fact that until the archipelago-wide fanihi population recovers, a 

sustainable hunt will not be possible. The public comments we received on the 

proposed rule to list the fanihi as threatened throughout its range suggest that the 

conservation constituency for this animal within the Mariana Islands is small. 

Thus, a key aspect of the recovery strategy for fanihi is involving hunters and the 

general public in on-the-ground research and conservation efforts to build that 

constituency. This involvement will be essential as a means of both tapping into 

an important source of natural history information and garnering local community 

support for and commitment to fanihi recovery and conservation for the long 

term. 

Implementing these actions requires building long-term support for and 

participation in the recovery effort; enhancing existing survey methodologies; 

conducting research and monitoring to address critical gaps in our knowledge and 

provide new information for effective conservation and recovery; and application 

of research and monitoring through adaptive management. The recovery strategy 

will be implemented as a collaborative effort among technical experts, agencies, 

the governments of the CNMI and Guam, and other participants and stakeholders. 
Draft Revised Recovery Plan for the Mariana Fruit Bat 

39

Recovery of the fanihi will be tracked via monitoring and annual reporting 

through federal and local wildlife agencies. Research recommended in this 

recovery plan will provide the information needed to ensure the recovery strategy 

is as effective as possible. As we acquire new information through monitoring, 

research, and other sources, we will revise this recovery plan and adapt the 

strategy for its implementation to ensure the greatest efficiency and effectiveness 

of recovery efforts for the fanihi. The specific recovery actions described in the 

narrative outline address the threats to fanihi with a focus on islands, and areas 

within islands, as necessary. The fanihi is widely distributed in the Mariana 

archipelago, and threats and recovery potential vary among islands. The 

expensive and complex logistics of working on the remote northern islands may 

pose significant challenges to recovery implementation. Long-term recovery 

objectives will likely require prioritization among islands. 

C. Recovery Criteria 

Delisting of a threatened species is achieved through a formal rulemaking 

process. The recovery criteria set forth in a recovery plan are intended to serve as 

objective, measurable guidelines to assist us in determining when a listed species 

has recovered to the point that the protections afforded by the Endangered Species 

Act are no longer necessary. However, the actual delisting process is not 

dependent solely on achieving the recovery criteria. The formal rulemaking 

process is based upon a five-factor analysis (per section 4(a)(1) of the Endangered 

Species Act), in conjunction with an analysis of the recovery criteria, that results 

in a determination that the threats have been sufficiently controlled or eliminated 

such that delisting is warranted.1

To achieve the goal of delisting the fanihi, we have defined recovery criteria 

that will demonstrate we have ensured the conservation and survival of the 

species, and that the threats to the species have been reduced to the point that the 

protections afforded by the Endangered Species Act are no longer necessary. 

These criteria are listed below. However, currently we lack information 

necessary for refining these criteria and the methods, such as population models, 

for measuring progress toward them. We need new estimates of the numbers of 

bats occurring on most islands and improved information on population 

 

1

 See section I.F above for a summary of the five factors as they apply to fanihi. 
Draft Revised Recovery Plan for the Mariana Fruit Bat 

40

dynamics and demographics (for example, fanihi-specific data on productivity, 

survivorship, age structure, sex ratios), specific habitat requirements, interisland 

movements, genetic structure of the fanihi population, and mortality rates 

associated with disturbances such as typhoons and hunting. We need this 

information to develop a scientifically credible estimate of the target population 

size, growth rate, distribution, and extent and characteristics of the habitat 

necessary to ensure the long-term viability of fanihi. Further research on the life 

history and ecology of the species thus is needed to set measurable thresholds for 

recovery and accurately monitor and project the fanihi’s response to management 

of threats. Results of such research may require us to revisit and revise these 

criteria. 

Recovery criteria below address the major threats to the species, including 

general criteria for population distribution and post-delisting monitoring, as well 

as additional criteria organized by the five Listing Factors to be used in a delisting 

analysis. Although storms and volcanic eruptions are a threat under Listing 

Factor E, no foreseeable management effort is likely to reduce the incidence of 

these threats. However, a well-distributed and stable or increasing population that 

meets Criterion 1 below is expected to be resilient enough to accommodate some 

local catastrophic events without endangering the overall survival of the species. 

The fanihi may be considered for delisting when the following criteria have 

been met: 

1. Population and Distribution 

The total size of the fanihi population has increased, based on data from 

scientifically defensible monitoring protocols, and stable or increasing 

subpopulations of sufficient size to avoid genetic and demographic risks 

associated with very small populations are distributed among the Mariana 

Islands so that the probability of the fanihi’s persistence over 100 years is high 

(at least 90 percent). 

To meet this criterion, we propose that stable or increasing fanihi 

subpopulations should be distributed as follows: three of the five southern 

islands (Saipan, Tinian, Aguiguan, Rota, and Guam), and six of the eight 

islands north of Saipan where fanihi have persisted historically (Anatahan, 
Draft Revised Recovery Plan for the Mariana Fruit Bat 

41

Sarigan, Guguan, Alamagan, Pagan, Agrihan, Asuncion, and Maug). Of the 

six northern islands with stable or increasing fanihi populations, two of these 

must include Pagan, Anatahan, or Agrihan, the three largest of the northern 

islands. This distribution may be modified as appropriate based on future 

monitoring data or population viability modeling (Recovery Action 5.2). 

2. Post-delisting monitoring 

A post-delisting monitoring program for the species has been developed to 

reliably detect population trends and is ready for implementation. 

3. Listing Factor A: Habitat Loss and Degradation 

Specific actions to restore habitat (including ungulate control and control of 

invasive non-native plants as appropriate for specific islands – see Table 3) 

have been identified and management plans developed as necessary for 

recovery under Criterion 1, and these actions and plans are being successfully 

implemented so that habitat loss and degradation no longer endanger the 

survival of the fanihi. 

4. Listing Factor B: Hunting 

Specific actions to reduce illegal hunting have been identified and 

management plans developed as necessary for recovery under Criterion 1, and 

these actions and plans are being successfully implemented so that illegal 

hunting no longer endangers the survival of the fanihi. 

5. Listing Factor C: Brown Treesnakes 

Long term measures are being successfully implemented to control the 

incipient brown treesnake population on Saipan and to prevent the 

introduction of the brown treesnake from Guam and Saipan to other islands in 

the CNMI. 

6. Listing Factor E: Development and Military Training Activities 

Impacts of urban development and military training on the fanihi are 

successfully being avoided, minimized, or mitigated so that they do not 

endanger the survival of the fanihi. 
Draft Revised Recovery Plan for the Mariana Fruit Bat 

42

III. RECOVERY ACTIONS 

A. Outline of Recovery Actions 

1. Develop education, outreach and enforcement programs to control illegal 

hunting of fanihi 

1.1. Develop education programs 

1.1.1. Elementary and secondary schools education 

1.1.2. Provide public access to fanihi, with interpretation 

1.1.3. Education for conservation officers and other public servants 

1.2. Develop outreach and incentive programs 

1.2.1. Involve hunters and local residents in research and recovery 

1.2.2. Work with local government officials, managers of protected 

areas, and private landowners to establish protected roost site 

sanctuaries for the bats where they can be monitored by local 

participants. 

1.2.3. Foster the creation of grass-roots conservation clubs and nonprofit organizations (similar to Audubon chapters), and provide 

these with community habitat restoration projects 

1.2.4. Establish an “ambassador” program 

1.3. Minimize illegal hunting of fanihi through law enforcement actions 

1.3.1. Provide law enforcement support to CNMI and Guam 

Conservation Officers for effective patrolling to prevent hunting 

at known roost sites and flyways on public and private lands, 

especially on Rota. 

1.3.2. Develop a multi-agency strategy, including roles and 

responsibilities of all agencies involved, to curb fanihi hunting 

though proactive law enforcement. 

1.3.3. Encourage local and Federal judicial authorities to investigate 

and prosecute illegal fanihi hunting aggressively. 

1.3.4. Improve inspection of all aircraft and vessels at ports of entry in 

the CNMI and on Guam to prevent illegal traffic of fanihi and 

other Pteropus species. 

1.3.5. Improve monitoring of permitted deer and pig hunters on 

Andersen Air Force Base 

2. Protect and restore habitat 
Draft Revised Recovery Plan for the Mariana Fruit Bat 

43

2.1. Determine Management Areas on islands where necessary, and prioritize 

sites for recovery actions 

2.2. Control ungulates 

2.2.1. Eradicate or significantly reduce populations of ungulates in 

Management Areas on Guam, Rota, Tinian, Aguiguan, and 

Saipan 

2.2.2. Eradicate or significantly reduce populations of ungulates and 

maintain reduction on Alamagan, Pagan, and Agrihan such that 

native forest can recover and fanihi numbers can be maintained 

or improved. 

2.2.3. Complete the ungulate eradication on Anatahan. 

2.2.4. Prevent the introduction of ungulates to Guguan, Asuncion, 

Maug, and prevent the reintroduction of ungulates to Sarigan 

2.2.5. Discourage the release of ungulates on public lands on all 

islands. 

2.3. Restore native vegetation 

2.3.1. Control invasive non-native plant species that limit native forest 

persistence and sustainability 

2.3.2. Propagate and out-plant native trees and shrubs 

3. Control the threat of brown treesnake predation on fanihi 

3.1. Determine impact of brown treesnakes to fanihi on Guam and 

determine best methods for protecting bats from treesnakes 

3.2. Prevent the spread of brown treesnakes and support brown treesnake 

control and interdiction efforts undertaken by all agencies 

4. Develop Federal cooperative conservation projects and amend existing plans 

and agreements with willing landowners to implement recovery on Federal 

and non-Federal lands 

5. Conduct monitoring and research to increase the knowledge base for 

management and updating the recovery plan 

5.1. Improve monitoring methods and monitor populations 

5.2. Conduct research on fanihi biology and ecology 

5.2.1. Conduct life history studies 

5.2.2. Conduct ecological studies 

5.2.3. Determine the genetic diversity within the fanihi population and 

across the range of Pteropus mariannus 

5.3. Conduct research on human-fanihi interaction 
Draft Revised Recovery Plan for the Mariana Fruit Bat 

44

5.4. Conduct or support relevant research on habitat restoration and 

conservation 

5.4.1. Support research to quantify impacts of alien ungulates and 

plants on native forest resources critical to fanihi. 

5.4.2. Support research to develop improved control and eradication 

methods for these alien species. 

6. Monitor implementation of the recovery plan and practice adaptive 

management in which recovery tasks are revised by the U.S. Fish and Wildlife 

Service in coordination with the Recovery Team and other partners as 

pertinent new information becomes available. 

6.1. Periodically review the recovery plan and revise or update it as 

appropriate 

6.2. Maintain an active recovery team, as needed 

6.2.1. Establish short-term objectives for the recovery program 

6.2.2. Coordinate recovery actions with other recovery and ecosystem 

management efforts 
Draft Revised Recovery Plan for the Mariana Fruit Bat 

45

B. Recovery Action Narrative 

1. Develop outreach, education, and enforcement programs to control illegal 

hunting of fanihi 

Building grassroots public support for fanihi recovery is vital to overcoming 

administrative and political barriers to recovery. Because such barriers 

develop as a result of the sociocultural environment, enhancement of public 

perception of the value of a healthy fanihi population will be a critical 

component of successful recovery efforts. To this end, this recovery plan 

includes a strong focus on education, outreach, and the close involvement of 

local landowners, communities, and decision-makers in the planning and 

implementation of recovery projects. 

1.1. Develop education programs 

1.1.1. Elementary and secondary schools education 

Improving children’s knowledge of natural systems and 

understanding of the role fanihi plays in the ecosystems of their 

islands is one of the most important and proactive long-term 

recovery actions for this species. This can be accomplished 

through a wide variety of specific projects, such as educating 

teachers in fanihi biology and ecology and developing natural 

science curricula that include fanihi; equipping local biologists to 

visit schools with fanihi awareness lessons; and “citizen science” 

school projects that involve school children in hands-on research 

that contribute to fanihi conservation (e.g., collecting fanihi 

droppings under a roost and have students assist with 

germinating the seeds and planting these in areas to be restored). 

1.1.2. Provide public access to fanihi (zoos, interpretation) 

Opportunity to view fanihi and have their behavior and ecology 

interpreted is a key step in increasing community acquaintance 

with fanihi as a living animal that is not only a delicious treat but 

also an essential part of the natural heritage of the Mariana 

Islands. Potential locations for developing fanihi viewing and 
Draft Revised Recovery Plan for the Mariana Fruit Bat 

46

interpretive materials and programs for the public include the 

Rota zoo and other facilities in the CNMI and on Guam. 

1.1.3. Education for conservation officers and other public servants 

To gain local support for fanihi conservation and enforcement of 

local and Federal laws, conservation officers and government 

officials at the island and Territorial and Commonwealth levels 

must become better educated about the ecology of the animal, the 

impacts that hunting has on the fanihi population, and the 

impossibility of sustainable hunting without a recovered 

population. This education could be accomplished through 

annual or quarterly classes or workshops organized and taught by 

multi-agency staff drawn from local and Federal natural 

resources management agencies. Certificates of completion 

could be offered to provide incentive for attendance. 

1.2. Develop outreach and incentive programs 

Public outreach and broad-based community planning can help promote 

public support and understanding of recovery actions for the fanihi. 

Fostering activities that engage, and ideally employ, the community in 

population monitoring and long-term management of fanihi will be 

crucial to conservation that has long term effects on both the fanihi and 

the perceptions and attitudes of the community. It will be important to 

identify the sectors of society in which outreach efforts will be most 

effective, and this may be accomplished by conducting marketing-style 

surveys to identify target audiences. In addition, monitoring the success 

of outreach projects is essential (for example, through surveys before 

and following to determine whether views and behaviors have changed). 

For all of the actions described below, even if an “offshore” firm is 

contracted to design and implement the work, individuals in the 

community must be sought who can model and champion local 

participation in conservation and stewardship of fanihi. 

1.2.1. Involve hunters and local residents in research and recovery 

Local hunters and other residents have specific local knowledge 

of fanihi and can provide assistance with location of roosts, 
Draft Revised Recovery Plan for the Mariana Fruit Bat 

47

seasonal feeding locations, and flight paths; fanihi diet research; 

and discussion of how best to control hunting. This community 

should be closely involved and employed if possible in 

information gathering and recovery implementation. The 

development of a community-based, “citizen science”, 

monitoring program is one way to do this. It will be essential to 

engage such a program in monitoring activities during periods 

when fanihi are most vulnerable (e.g., before fiestas and after 

large storms. 

1.2.2. Work with local government officials, managers of protected 

areas, and private landowners to establish protected roost 

site sanctuaries for the bats where they can be monitored by 

local participants 

This effort can be used to obtain information about roosting bats 

such as reproductive status, resighting of banded bats, etc., as 

well as generate a sense of “ownership” and stewardship of 

fanihi and their roost sites in the community. 

1.2.3. Foster the creation of grass-roots conservation clubs and 

non-profit organizations, and provide these with community 

habitat restoration projects 

Involving local people in projects such as controlling invasive 

plants, propagating and outplanting native species, and 

participating in other restoration work and in study and 

monitoring of fanihi (e.g., similar to the work done by Audubon 

Society chapters) creates a lasting investment by local 

communities and commitment to the ongoing conservation of 

specific places they have worked hard to restore. Currently 

outreach efforts of this kind are minimal in the Marianas. 

1.2.4. Establish an “ambassador” program 

A team of community members, scientists and agency personnel 

should meet with groups of citizens on Guam and in the CNMI 

to discuss fanihi recovery. Such meetings could be organized 
Draft Revised Recovery Plan for the Mariana Fruit Bat 

48

and hosted by clubs/organizations described in 1.2.3., above. 

These ambassadors should be prepared to discuss the causes of 

endangerment, findings to date, planned actions for achieving 

recovery, the ramifications of the Draft Revised Recovery Plan 

for the fanihi, Federal and local laws concerning endangered 

species, and the special needs and concerns of citizens. 

1.3. Minimize illegal hunting of fanihi through law enforcement actions 

 

1.3.1. Provide law enforcement support to CNMI and Guam 

Conservation Officers 

In the close-knit communities of the Pacific, enforcing laws can 

be a social and cultural challenge. In the Marianas, conservation 

officers would benefit from collaboration with and support from 

Federal law enforcement officers, such as USFWS special 

agents, to improve effective patrolling of known hunting areas, 

apprehend illegal hunters, and conduct other actions to prevent 

hunting at roost sites and flyways on public and private lands, 

especially on Rota. An exchange program could be developed to 

strengthen ties between Federal, Territorial, and Commonwealth 

officers, facilitate additional training for Territorial and 

Commonwealth officers, and build capacity for conservation law 

enforcement in the Marianas. 

1.3.2. Develop a multi-agency strategy, including roles and 

responsibilities of all agencies involved, to curb fanihi 

hunting though proactive law enforcement 

The fanihi is protected under Guam, CNMI, and U.S. Federal 

law. Greater cooperation and coordination are needed among 

entities responsible for enforcing these laws to ensure this is 

done consistently, proactively, and in a manner that promotes 

rather than discourages voluntary compliance by the public. 

Coordination and outreach through the Stakeholder Subgroup of 

the Recovery Team and other avenues will be pursued to inform 

the public of this recovery plan and to include public input into 

recovery implementation. 
Draft Revised Recovery Plan for the Mariana Fruit Bat 

49

1.3.3. Encourage local and Federal judicial authorities to 

investigate and prosecute illegal fanihi hunting aggressively 

Ongoing and new education and outreach efforts to curb hunting 

of fanihi will take time to reach the target communities in the 

CNMI and on Guam. In the interim, local and Federal judicial 

authorities should be made aware of the serious threat that illegal 

hunting poses to the fanihi, and, together with law enforcement 

agencies, should be provided the resources to effectively 

investigate and successfully prosecute cases of unlawful hunting. 

In tandem with education, effective law enforcement is a key 

component of conservation. 

 

1.3.4. Improve inspection of all aircraft and vessels at ports of 

entry in the CNMI and on Guam to prevent illegal traffic of 

fanihi and other Pteropus species 

Better and more frequent inspections are needed of boats and 

aircraft arriving on Saipan from the Northern Islands and Rota, 

arriving on Guam from Rota, and arriving anywhere in the 

Marianas from other sources of or transit points for illegal 

shipments of fanihi or other fruit bats. Sniffer dogs and 

increased inspection and enforcement staff are needed to carry 

out these inspections. 

1.3.5. Improve monitoring of permitted deer and pig hunters on 

Andersen Air Force Base 

Anecdotal reports exist of ungulate hunters on AAFB 

opportunistically taking fanihi that are transiting or foraging in 

the tracts of forest on the base. Ingress and egress of hunters 

should be monitored; sniffer dogs and conservation officers 

could be placed at stations to check bags of ungulate hunters on 

AAFB for illegally taken fanihi. 

2. Protect and restore habitat 
Draft Revised Recovery Plan for the Mariana Fruit Bat 

50

2.1. Determine Management Areas on islands where necessary, and 

prioritize sites for recovery actions 

Opportunities for fruit bat recovery will vary among islands in the 

archipelago; conservation actions may not be pressing or necessary on 

every island, and recovery does not depend upon an even geographic 

distribution of recovery efforts. Rather, we anticipate that recovery 

efforts will be focused on islands, and in specific Management Areas 

where necessary (i.e., on the islands with large human populations), that 

are determined by the U.S. Fish and Wildlife Service in consultation 

with our partners in the Mariana Islands and the recovery team to offer 

the best opportunities to meet the criteria for recovery of the fanihi. 

These Management Areas within islands might be areas that harbor 

active or historically active roost sites or fanihi habitat determined to be 

essential to recovery. We recognize that some jurisdictions and 

landowners on the islands may not wish to participate in the recovery 

effort. Within these conditions, priorities for habitat protection and 

restoration should be (a) areas/islands currently harboring fanihi, (b) 

areas/islands critical to recovery whether or not they harbor fanihi now, 

and (c) other islands/areas. 

2.2. Control ungulates 

2.2.1. Eradicate or significantly reduce and confine ungulates in 

Management Areas on Guam, Rota, Tinian, Aguiguan, and 

Saipan 

In the context of large and growing human populations in the 

Southern Islands of the Marianas, native forests must be retained 

and restored to a condition of natural regeneration in areas of 

sufficient size and connectivity to ensure that habitat containing 

key resources for fanihi and other native wildlife is safeguarded 

for the long term. In addition to data gathering and other handson outreach projects described above, forest restoration and 

maintenance are activities that should involve and employ the 

local community. 
Draft Revised Recovery Plan for the Mariana Fruit Bat 

51

2.2.2. Eradicate or significantly reduce and confine ungulates and 

maintain reductions in the CNMI Northern Islands 

The isolated Northern Islands with their occasional, small human 

populations are the stronghold of the fanihi population, and the 

habitat for native wildlife on these islands must be protected and, 

where necessary, restored. Ungulates must be controlled on 

these islands so that the native forest is retained and restored to a 

condition of natural regeneration to ensure that habitat containing 

key resources for fanihi and other native wildlife is safeguarded 

for the long term. Alamagan, Pagan, and Agrihan should be 

cleared of ungulates or, if human settlers are present, livestock 

should be maintained in small numbers specifically to meet the 

needs of people living on the island, and should be removed 

when settlements are abandoned. 

2.2.3. Complete the ungulate eradication on Anatahan 

When Anatahan’s volcano erupted in 2003, eradication of goats 

and pigs from the island was nearing completion. This island is 

one of the three largest in the Northern Islands and in the past has 

harbored a large number of bats. Completing this eradication 

now, while the island is still relatively devegetated, should be the 

first priority for fanihi recovery in the Northern Islands.

2.2.4. Prevent the introduction of ungulates to Guguan, Asuncion, 

Maug, and prevent the reintroduction of ungulates to 

Sarigan 

These four islands are designated “Conservation Islands” by the 

CNMI government and should remain free of ungulates and other 

new non-native species. 

2.2.5. Discourage the release of ungulates on public lands on all 

islands 

Additional populations of ungulates on any islands will only 

hamper habitat restoration and increase the degradation of 

existing forest habitat. Public outreach and education may help 
Draft Revised Recovery Plan for the Mariana Fruit Bat 

52

to dissuade new introductions of ungulates by individuals or 

groups. 

2.3. Protect and restore native vegetation 

Once released from hunting pressure, fanihi can potentially use habitat 

ranging over all of an inhabited island such as Rota. Thus, a general 

policy should be pursued of “no net loss” of native forest and restoration 

of forests on islands and in areas critical to fanihi recovery. 

2.3.1. Control invasive plant species that limit native forest 

persistence and sustainability 

The control of species such as Leucaena leucocephala (tangantangan) and Casuarina equisetifolia that form monotypic stands 

and provide few resources for native wildlife should be 

undertaken strategically in areas prioritized to protect and expand 

existing native forests. 

2.3.2. Propagate and out-plant native trees and shrubs 

Outplanting will increase the rate of native forest recovery (e.g., 

in areas where ungulates or invasive plants are removed). When 

undertaken around fragments of native forest, outplanting 

increases the resilience of these patches to incursion by alien 

plant species and reduces the threat of these species becoming 

established over large areas. However, outplanting trees and 

plants on their native islands is a broadly beneficial conservation 

action around villages and around and within agroforests, as 

well. In particular, plants known to be important resources for 

fanihi, such as Artocarpus mariannus (seeded breadfruit),

Pandanus tectorius (kafu), Cycas micronesica, Terminalia 

catappa (umbrella tree), and native Ficus spp., should be 

propagated and planted where appropriate. 

3. Control the threat of brown treesnake predation on fanihi 

 
Draft Revised Recovery Plan for the Mariana Fruit Bat 

53

3.1. Determine impact of brown treesnakes to fanihi on Guam and 

determine and implement best methods for protecting bats from 

treesnakes 

Scientists and land managers currently conducting treesnake research 

and control on Guam should provide a special focus on protecting fanihi 

on Guam insofar as predation by treesnakes plays a role in exacerbating 

the decline of bats on the island. Examining the stomach contents or 

conducting stable isotope analysis of tissues of snakes trapped in the 

vicinity of the fanihi roost in northern Guam may yield further evidence 

of snakes preying on bats. Control of brown treesnakes should be 

undertaken in the vicinity of the roost on Guam only if there is 

conclusive evidence that (a) such control will demonstrably benefit the 

bats, (b) such work can be undertaken without running the risk of the 

roost being abandoned, and (c) the results of such work will provide 

information that could be applicable to protecting fruit bats from 

predation by treesnakes on Saipan, or on other islands. 

3.2. Prevent the spread of brown treesnakes and support brown 

treesnake control and interdiction efforts undertaken by all agencies

Barriers, traps, and other interdiction procedures must be maintained or 

established at all ports of entry to the CNMI to minimize the risk of 

treesnake introduction from Guam and Saipan. 

4. Develop Federal cooperative conservation projects and amend existing 

plans and agreements with willing landowners to implement recovery on 

Federal and non-federal lands 

An important potential source of protection for fanihi and their habitat is 

management codified in conservation agreements with local governments and 

willing landowners on Federal and non-Federal lands. Management will 

include maintaining or improving forest habitat, reducing or eliminating direct 

harvest of fanihi through education and law enforcement actions, eliminating 

non-native species or reducing their populations through rigorous planned 

management, and other actions. 

Existing land use and natural resource management plans for islands or areas 

within islands, including Integrated Natural Resource Management Plans for 
Draft Revised Recovery Plan for the Mariana Fruit Bat 

54

military bases and other documents, should be reviewed by the U.S. Fish and 

Wildlife Service and the appropriate entities to ensure that recovery needs for 

fanihi are addressed and triggers for implementation of recovery actions and 

monitoring of results are included. All actions proposed on U.S. military 

lands on Guam and in the CNMI must receive thorough review by biologists 

familiar with fanihi and its habitat. 

Through surveys, monitoring, and regulatory review and consultation under 

the National Environmental Policy Act and Endangered Species Act, we can 

assess the potential effects on the fanihi of activities planned by the U.S. 

Navy, U.S. Air Force, and other defense agencies, and determine the best 

methods of avoiding, minimizing, and mitigating adverse impacts to this and 

other listed species. For example, clearing of forested land in proximity to a 

fanihi roost is an action that should be proposed to the U.S. Fish and Wildlife 

Service and reviewed as appropriate before being implemented. 

5. Conduct monitoring and research to increase the knowledge base for 

management and updating the recovery plan 

Monitoring is necessary to track the status of fanihi throughout the 

archipelago, to assess threats to bats and their habitat, and to evaluate the 

implementation and effectiveness of conservation actions and this recovery 

plan. Development of scientifically acceptable monitoring protocols is needed 

to accomplish these tasks. Significant gaps remain in our scientific 

knowledge of the fanihi. It is incumbent on us to see that those gaps are filled 

as they pertain to achieving a recovered fanihi population; we must ensure that 

the objectives we set and the tools used to meet them are the right ones to 

reach our definition of “recovered” for this species. Therefore some research 

will be integral to recovery for this animal. The results of recovery-oriented 

research will over time aid in refining the fanihi recovery program. 

Specifically, research is needed to address questions about life history and 

ecology that are crucial to accurately projecting population persistence, 

monitoring response to management, and adapting and refining our approach 

to management to reflect new information. Critically, the results of this 

research will aid in determination of thresholds for bat numbers and other 

conservation criteria on each island that could indicate when the fanihi 

population has recovered sufficiently for some legal hunting to resume. The 
Draft Revised Recovery Plan for the Mariana Fruit Bat 

55

fanihi population is clearly too precarious at present to support a hunting 

season. However, if illegal hunting and other threats are reduced so that the 

species’ status can improve substantially and our knowledge of population 

dynamics and threats allows us to identify a limited, sustainable hunting 

regime consistent with recovery of the species, such a hunt might be allowed 

under the Endangered Species Act either by delisting the species or through a 

special rule under section 4(d) of the Act. 

5.1. Improve monitoring methods and monitor populations 

A great deal of effort has been invested in fanihi surveys and monitoring 

in the past, but the goals for monitoring were not always identified, 

methods used were not standardized, and there has been little 

coordinated monitoring among the islands. Thus, we currently have 

imprecise estimates of population size and trends and limited ability to 

make comparisons among islands. 

Enhancement of existing monitoring methods is needed to develop 

scientifically rigorous population monitoring programs for the Southern 

and Northern Islands. Such enhancement must include: improved 

measures of the variability in survey data (e.g., observer error; detection 

of bats at roosts, where not all bats may be visible) and in numbers 

through time (e.g., are there seasonal increases associated with 

recruitment of juveniles and/or seasonal declines associated with eventspecific hunting?). Study and experimentation are needed to determine 

the most efficient monitoring schedule and best suite of survey methods 

for monitoring fanihi bats on individual islands and throughout the 

archipelago. Monitoring programs in the Southern Islands may be 

intensive and involve frequent surveys; in the Northern Islands, 

infrequent access and logistical constraints may require development of 

rapid but rigorous monitoring methods. To the extent that logistical 

considerations permit, monitoring methods must be standardized or 

statistically comparable between islands and between surveys. Finally, 

training programs are needed to ensure consistency and full 

understanding of monitoring methods and objectives. This applies 

equally to expert and “community” observers participating in bat 

monitoring. Monitoring data will be compiled into reports to assess 
Draft Revised Recovery Plan for the Mariana Fruit Bat 

56

recovery plan implementation and whether the recovery criteria are 

being addressed. Where appropriate, summaries of monitoring data 

should be made available to the public as part of the outreach program. 

A project that includes intensive monitoring at intervals through the year 

on an island without (e.g., Sarigan) and with (e.g., Rota) hunting or other 

human disturbance would provide much needed information about 

sources and magnitude of variability in survey data. This information is 

needed to refine survey methods so that data collected may be used to 

detect trends on a timescale sufficient to address threats and measure 

progress toward recovery. 

5.2. Conduct research on fanihi biology and ecology 

We have conducted preliminary experiments with population viability 

analysis (PVA) to examine the sensitivity of the fanihi population to 

various threats and to potential recovery actions. These models will help 

us further refine recovery criteria, set population thresholds, and project 

the species’ response to management. Available data describing fanihi 

vital rates (and other traits of the species) to refine input parameters for 

population projection are minimal and need improvement. Two areas 

where more research is needed to develop better parameter estimates are 

migration patterns among islands, and the response of bat populations 

(mortality, migration, reproduction rates) after natural disturbance events 

of different severity. 

5.2.1. Conduct life history studies 

Length of gestation, age of sexual maturity, reproductive success, 

annual survivorship of various age classes, and recruitment into 

the breeding population are unknown for the fanihi; these life 

history parameters influence the population-level effects of 

natural and anthropogenic disturbance events. These parameters 

also influence the response of the population to recovery actions. 

Data describing these traits as well as reproductive strategies, 

generation time, and reproductive rates and success are needed 

for modeling intrinsic capacity for population growth and long 

term persistence with any degree of confidence. This research 
Draft Revised Recovery Plan for the Mariana Fruit Bat 

57

could be undertaken using direct observation of bats at roosts, 

mark-resight studies, and potentially more invasive study of 

closely related proxy species, as has been done for the critically 

endangered Guam Micronesian kingfisher or sihek (Halcyon 

cinnamomina cinnamomina), which currently survives only in 

captivity. 

5.2.2. Conduct ecological studies 

Conservation-directed ecological study of fanihi includes 

determination of dietary and other habitat selection requirements, 

foraging range, seasonal variation in foraging behavior, the 

quality and areal extent of habitat for bats on each island (and 

investigation of how this relates to carrying capacity for 

individual islands), and direct and indirect impacts of typhoons 

on bat mortality. This information will inform and refine habitat 

conservation and restoration efforts. This research could be 

undertaken using a combination of many approaches, such as 

radio-tracking; direct foraging observation; interviews with 

hunters and fruit farmers; collection and analysis of bat ejecta 

and feces beneath foraging and roosting trees; and defining, 

mapping, and quantification of bat habitat and typhoon impacts 

using ground surveys, remotely-sensed imagery, and Geographic 

Positioning and Information Systems (GPS and GIS). 

5.2.3. Determine the genetic diversity within the fanihi population 

and across the range of Pteropus mariannus 

Genetic information is needed to infer the historical connectivity 

between islands, determine whether negative inbreeding effects 

may be a concern, and determine the relatedness of fanihi with 

subspecies of Pteropus mariannus in other archipelagos. Some 

genetic work is needed to ascertain whether additional endemic 

subspecies occur in the Mariana archipelago and should be listed 

as separate entities. Molecular analysis could be undertaken 

using genetic material extracted from tissue samples or, 

potentially, from fecal matter and ejecta. 
Draft Revised Recovery Plan for the Mariana Fruit Bat 

58

With further research, DNA extracted from fecal material and 

ejecta also holds potential for conducting non-invasive markrecapture studies, and assessing maternity and paternity rates and 

connectivity among roost sites. By collecting these samples 

beneath roosts, the potential injury and other trauma that may be 

associated with trapping bats is avoided. 

5.3. Conduct research on human-fanihi interaction 

Hunters and other residents are a critical source of information about 

local knowledge of fanihi ecology and biology as well as perceptions of 

the limits of bat populations (as an exploitable resource). Understanding 

the interests and perspectives of hunters is also crucial to identifying the 

best ways to address hunting regulation. Finally, hunters and local 

residents are likely to provide insights into other strategies for promoting 

fanihi conservation, such as how fanihi may play a role in local pride-ofplace or as a flagship conservation species. Hunters and other local 

stakeholders are important links to the local community and their input 

should help to identify strategies for engaging local communities and 

encouraging participatory conservation management. 

Interviews of hunters and other local residents will be conducted to help 

determine mortality rates and spatial and temporal patterns in hunting 

pressure. This information will help to refine our understanding of 

population-level impacts of hunting and to target implementation of 

education, outreach, and enforcement efforts where and when they will 

be most effective. 

5.4. Conduct or support relevant research on habitat restoration and 

conservation 

5.4.1. Support research to quantify impacts of alien ungulates and 

plants on native forest resources critical to fanihi 

Foraging by ungulates is known generally to reduce or preclude 

recruitment of plant species, reduce native species diversity, and 

ultimately lead to devegetation and erosion. However, ungulate 

eradication can result in “release” of alien plant species that can 
Draft Revised Recovery Plan for the Mariana Fruit Bat 

59

impede the recovery of native vegetation. The dynamics of alien 

species invasions of native plant communities in the Marianas 

are poorly known. Study of the impacts of alien invaders on 

plant species that provide resources for fanihi (e.g., Artocarpus 

mariannensis, Erythrina variegata (Indian coral tree), and Ficus

spp.) will help us develop strategies for control of alien species 

such as goats and the vine Operculina ventricosa (St. Thomas 

lidpod) and prioritize sites for treatment. 

5.4.2. Support research to develop improved control and 

eradication methods for alien species. 

Efficient and cost-effective methods are needed for control and 

eradication of alien species, especially invasive plants. A 

particular need exists for methods that can be used effectively in 

the remote northern islands, where access is difficult and 

intermittent at best. 

6. Monitor implementation of the recovery plan and practice adaptive 

management in which recovery tasks are revised by the U.S. Fish and 

Wildlife Service in coordination with the Recovery Team and other 

partners as pertinent new information becomes available. 

The list of actions above is necessarily broad and not all-inclusive. As 

additional information becomes available, new risks and opportunities may 

come to light, and additional tasks will be identified and priorities shifted as 

appropriate. 

6.1. Periodically review the recovery plan and revise or update it as 

appropriate 

The restoration of an endangered species is an uncertain science that 

requires continual critique and reevaluation of approach. A regularly 

updated recovery plan will assure all participants that recovery is guided 

by the best available science and will be in keeping with our current 

guidance on recovery planning.

6.2. Maintain an active recovery team, as needed 
Draft Revised Recovery Plan for the Mariana Fruit Bat 

60

The recovery team can serve as a source of recommendations and 

guidance regarding fanihi recovery. The team is a forum in which issues 

surrounding recovery may be discussed and effective, coordinated 

recovery strategies developed. Although significant experience and 

expertise are represented on the team now, we will also need access to 

specialists outside the team for input on specific points and questions. 

Several principles should guide the team's work. The team should 

maintain an awareness of all activities that have a major impact on fanihi 

recovery. The team should encourage peer review and publication of all 

scientific findings used in fanihi management. Management 

recommendations unsuitable for publication should nonetheless be 

subjected to independent peer review. The team should make substantial 

and continuing efforts to identify stakeholders in fanihi recovery and 

draw them into the recovery program to make meaningful contributions. 

6.2.1. Establish short-term objectives for the recovery program 

With assistance from team members, we will develop 2- to 5-

year implementation plans, providing the rationale for each 

objective. These implementation plans should: (a) describe 

specific objectives; (b) explain how achieving the objective will 

contribute to recovery; (c) provide evidence that achieving the 

objective is feasible; (d) describe the funding and other resources 

needed; and (e) provide evidence that the resources to be 

committed are best used for the proposed activity rather than for 

some other aspect of fanihi recovery. Each objective should 

include measures that can be used to monitor progress. 

6.2.2. Coordinate recovery actions with other recovery and 

ecosystem management efforts 

Many of the members of the recovery team are ideally situated to 

help us coordinate recovery implementation for the fanihi with 

other conservation initiatives on Guam and in the CNMI. 

Particular attention should be given to coordination on efforts to 

prepare habitat conservation plans with the government of the 

Commonwealth of the Northern Mariana Islands and with groups 
Draft Revised Recovery Plan for the Mariana Fruit Bat 

61

focused on the recovery of other listed species in the Marianas 

(e.g., Mariana crow recovery team, Guam Micronesian 

Kingfisher recovery committee). 
Draft Revised Recovery Plan for the Mariana Fruit Bat 

62

IV. RECOVERY IMPLEMENTATION SCHEDULE FOR 

2010-2014 

The Implementation Schedule that follows lists and prioritizes the actions 

and estimated costs for the recovery of the fanihi. It is a guide for meeting the 

recovery goals outlined in this plan. Recovery actions in the Implementation 

Schedule have been prioritized in a ranking system with each action being 

assigned a “priority number” from 1 (highest priority) to 3 (lowest priority; see 

definitions below). The numbers in the Action Number column correspond to the 

descriptions of recovery actions in the Narrative Outline of Recovery Actions. 

Parties with authority, responsibility, or expressed interest to implement a 

specific recovery action are also identified in the Implementation Schedule. 

When more than one party has been identified the proposed lead party is indicated 

by an asterisk (*). In cases where a lead party has not been identified, each party 

listed is individually responsible for implementing the recovery action. The 

listing of a party in the Implementation Schedule does not require, nor imply, that 

the identified party will implement the action(s) or secure funding for 

implementing the action(s). However, parties willing to participate may benefit 

by being able to show in their own budgets that their funding request is for a 

recovery action identified in an approved recovery plan and is therefore 

considered a necessary action for the overall coordinated effort to recover the 

fanihi. Also, section 7(a)(1) of the Endangered Species Act (16 USC 1531 et 

seq.) directs all Federal agencies to utilize their authorities in furtherance of the 

purposes of the Endangered Species Act by carrying out programs for the 

conservation of threatened and endangered species. 

Definition of action priorities: 

• Priority 1: An action that must be taken to prevent extinction or prevent 

the species from declining irreversibly in the foreseeable future. 

• Priority 2: An action that must be taken to prevent a significant decline in 

species population or habitat quality, or some other significant negative 

impact short of extinction. 

• Priority 3: All other actions necessary to meet the recovery objectives. 
Draft Revised Recovery Plan for the Mariana Fruit Bat 

63

Definition of action durations: 

• Continuous: An action that will be implemented on a routine basis once 

begun for the period of time estimated to recovery (in this case, 10 years). 

• Ongoing: An action that is currently being implemented and will continue 

until the time estimated to recovery. For the purposes of cost estimation, 

we used our best estimate of the time that may be required to complete the 

action. 

• Unknown: Either action duration or associated costs are not known at this 

time. For the purposes of cost estimation, we used our best estimate of the 

time that may be required to complete the action. 

Threat categories: 

We consider the role of five potential factors affecting the species in order 

to list, delist, or reclassify a taxon. These factors are: 

(A) the present or threatened destruction, modification or curtailment of its 

habitat or range; 

(B) overutilization for commercial, recreational, scientific, or educational 

purposes; 

(C) disease or predation, 

(D) the inadequacy of existing regulatory mechanisms; 

(E) other natural or manmade factors affecting its continued existence. 

Recovery actions are designed to address the threats in the Listing Factor 

column in order to meet the recovery criteria (see Recovery Criteria section). The 

majority of the recovery actions in this plan address habitat loss (factor A), 

overutilization (factor B), predation (factor C), and other natural factors affecting 

its continued existence (factor E). Existing regulatory mechanisms (factor D) 

appear adequate, as the fanihi is listed as threatened by the Federal government 

and consequently receives protection under the provisions of the Endangered 

Species Act. 
Draft Revised Recovery Plan for the Mariana Fruit Bat 

64

Responsible Parties for Action Implementation: 

We have statutory responsibility for implementing this recovery plan. 

Only Federal agencies are mandated to take part in the effort under section 7(a)(1) 

of the Endangered Species Act (16 USC 1531 et seq.). However, species 

recovery will require the involvement of the full range of Federal, territorial, 

private, and local interests. The expertise and contributions of additional agencies 

and interested parties will be needed to implement recovery actions and to 

accomplish education and outreach objectives. For each recovery action 

described in the Implementation Schedule, the column titled “Responsible 

Parties” lists the primary Federal and local agencies we have identified as having 

the authority and responsibility for implementing recovery actions and other 

groups, partners, and partnerships who are actively involved in recovery. 

Key to Acronyms used in the Implementation Schedule: 

• CNMI: Government of the CNMI 

• DAWR: Guam Division of Aquatic and Wildlife Resources 

• DFW: CNMI Division of Fish and Wildlife 

• DLNR: CNMI Department of Land and Natural Resources 

• DoD: U.S. Department of Defense 

• GoG: Government of Guam 

• NGO: Non-governmental organizations, including community groups 

• U/P: University or privately contracted researchers 

• USFWS-ES: United States Fish and Wildlife Service, Ecological Services 

• USFWS-LE: United States Fish and Wildlife Service, Law Enforcement 

• USFWS-R: United States Fish and Wildlife Service, Refuges Division 

• USGS-BRD: United States Geological Survey, Biological Research 

Discipline 

• WS: United States Department of Agriculture, Wildlife Services 
Draft Revised Recovery Plan for the Mariana Fruit Bat 

65

Cost estimates: 

The costs of implementing the identified recovery actions are estimated over two 

timeframes: the first five years covered by this recovery plan (5-Year Costs 

column) and the total costs of recovery over a 10-year period (10-Year Costs 

column).
66

Cost Estimate (in $10

Priority 

Number 

Action 

Number 

Listing 

Factor 

Action 

Description 

Action 

Duration 

Responsible 

Parties 

10- 

Year 

Costs 

FY 

10 

FY 

11 

FY 

12 

1 1.2.2 B Work with local 

government, land managers and owners to establish roost site sanctuaries 

Ongoing DLNR, 

USFWS-ES 

3.8 .3 .5 .5 

1 1.3.1 B Provide law enforcement 

support to CNMI and Guam Conservation Officers for fanihi protection 

Continuous USFWSLE*, ES; 

DAWR, DLNR 

150 15 15 15 

1 1.3.2 B Develop a multi-agency 

strategy to curb illegal hunting through law enforcement 

Continuous DAWR, 

DLNR, USFWS-LE 

6 .5 1 1 

1 1.3.3 B Encourage local and 

Federal judicial authorities to investigate and prosecute fanihi hunting aggressively 

Continuous USFWS-ES, 

LE; DAWR, DFW* 

5 .5 .5 .5 

2 1.2.1 B Involve hunters and local 

residents in research and recovery 

Continuous DFW*, 

DAWR, USFWS-ES 

19 1 2 2 
67

Cost Estimate (in $10

Priority 

Number 

Action 

Number 

Listing 

Factor 

Action 

Description 

Action 

Duration 

Responsible 

Parties 

10- 

Year 

Costs 

FY 

10 

FY 

11 

FY 

12 

2 1.3.4 B Improve inspection at 

ports of entry to prevent illegal traffic in fanihi 

Continuous USFWSLE*, DoD, 

GoG, CNMI 

50 3 7 5 

2 1.3.5 B Improve monitoring of 

permitted ungulate hunters on AAFB 

Continuous DoD 20 2 2 2 

2 2.2.1 A Eradicate or reduce and 

confine ungulates in Management Areas on Southern Islands 

Ongoing DOA*, 

DLNR*, DoD*, USFWS-R 

30 3 3 3 

2 2.2.2 A Eradicate or reduce and 

confine ungulates in the CNMI Northern Islands 

10 years DLNR*, 

DoD 

50 5 5 5 

2 2.2.3 A Complete ungulate 

eradication on Anatahan 

3 years DoD*, 

USFWS-ES 

12 4 4 4 
68

Cost Estimate (in $10

Priority 

Number 

Action 

Number 

Listing 

Factor 

Action 

Description 

Action 

Duration 

Responsible 

Parties 

10- 

Year 

Costs 

FY 

10 

FY 

11 

FY 

12 

2 2.2.4 A Prevent the introduction 

of ungulates to CNMI Conservation Islands (Sarigan, Guguan, Asuncion, Maug) 

Ongoing CNMI, 

DLNR 

2 .2 .2 .2 

2 2.2.5 A Discourage the release of 

ungulates on public lands on all islands 

Continuous CNMI, 

DLNR 

2 .2 .2 .2 

2 3.1 C Determine best methods 

for protecting bats from BTS on Guam 

2 years USGS-BRD, 

USDA-WS, DAWR 

6 3 3 - 

2 3.2 C Prevent the spread of BTS 

from Guam and Saipan 

Ongoing USDA-WS 1,600 160 160 160 

2 4 E Develop cooperative 

conservation projects; amend existing plans and agreements 

Continuous 

USFWSES*, DoD 

230 90 30 30 

2 5.1 A, B,C, 

E 

Improve monitoring methods and monitor fanihi population 

Continuous DFW*, 

DAWR*, USFWS-ES, USFWS-R 

34 5.5 5.5 5.5 
69

Cost Estimate (in $10

Priority 

Number 

Action 

Number 

Listing 

Factor 

Action 

Description 

Action 

Duration 

Responsible 

Parties 

10- 

Year 

Costs 

FY 

10 

FY 

11 

FY 

12 

2 5.2.1 A, B, C, 

E 

Conduct life history studies 

10 years DFW*, 

DAWR, USGS-BRD, USFWS-ES, U/P 

52 4 8 8 

2 5.2.2. A, B, C, 

E 

Conduct ecological studies 

5 years DFW*, 

DAWR, USGS-BRD, USFWS-ES, U/P 

32 4 8 8 

3 1.1.1. A, B Education in elementary 

and secondary schools about fanihi biology, cultural value, and threats 

Continuous DFW*, 

DAWR 

16 4 4 1 

3 1.1.2 B Provide public access to 

fanihi in zoos or other facilities 

Ongoing DFW, 

DAWR 

10 1 1 1 

3 1.1.3 B Education for 

conservation officers and other public servants 

Continuous DFW, 

DAWR 

6.5 2 .5 .5 
70

Cost Estimate (in $10

Priority 

Number 

Action 

Number 

Listing 

Factor 

Action 

Description 

Action 

Duration 

Responsible 

Parties 

10- 

Year 

Costs 

FY 

10 

FY 

11 

FY 

12 

3 1.2.3 A, B, E Foster the creation of 

grass-roots conservation clubs, provide these with community habitat restoration projects 

Continuous DLNR, 

DOA 

2.7 0.5 0.2 0.3 

3 1.2.4 A, B, E Establish an 

“ambassador” program 

Continuous DLNR, 

DOA 

6 1.5 0.5 0.5 

3 2.3.1 A Strategic control of 

invasive plant species (e.g., Leucaena, 

Casuarina) 

Continuous DLNR, 

DOA 

26 5 5 2 

3 2.3.2 A 

Propagate and outplant native trees and shrubs 

Continuous DOA, 

DLNR, NGO 

7 2 1 0.5 

3 5.2.3 A, B, C, 

E 

Determine genetic diversity within fanihi population and across range of P. mariannus

3 years U/P 

6 3 2.5 .5 

3 5.3 A, B 

Conduct research on human-fanihi interaction 

3 years DFW*, 

DAWR, U/P 2.5 1 1 0.5 
71

Cost Estimate (in $10

Priority 

Number 

Action 

Number 

Listing 

Factor 

Action 

Description 

Action 

Duration 

Responsible 

Parties 

10- 

Year 

Costs 

FY 

10 

FY 

11 

FY 

12 

3 5.4.1 A 

Support research on ungulate impacts on native forest 

3 years DLNR, 

DOA, DoD, USFWS-ES, R 

6 2 2 2 

3 5.4.2 A 

Support research on ungulate control and eradication methods 

2 DOA, DoD, 

DLNR, USFWS-ES 

4 2 2 - 

3 6.1 A, B, C, 

E 

Periodically review and revise or update the recovery plan 

1 year USFWS-ES 

3 - - - 

3 6.2.1 A, B, C, 

E 

Establish short-term (2-5 year) objectives for recovery implementation 

Continuous USFWS-ES 

5 1 - 1 

3 6.2.2 A, B, C, 

E 

Coordinate recovery actions with other recovery and management efforts 

Continuous USFWS-ES 

3 .3 .3 .3 

TOTALS: 2,304.3 
Draft Revised Recovery Plan for the Mariana Fruit Bat 

72

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