samoensis Species Report May 2020

U.S. Fish and Wildlife Service Pacific Islands Fish and Wildlife Office Honolulu, HI Cover Photo Credits Mao, Gymnomyza samoensis photographs courtesy R. Stirnemann.

Suggested Citation USFWS. 2020. Species Status Assessment for Mao (Gymnomyza samoensis). May 2020 (Version 1.1). U.S. Fish and Wildlife Service, Pacific Islands Fish and Wildlife Office, Honolulu, HI. 46 pp.

Primary Authors Version 1.1 of this document was prepared by Fred Amidon and James Kwon of the Pacific Islands Fish and Wildlife Office, Honolulu, Hawaii. Preparation and review was conducted by Gregory Koob, Megan Laut, and Stephen E. Miller of the Pacific Islands Fish and Wildlife Office.

ii

Executive Summary The Mao or Ma‘oma‘o (Gymnomyza samoensis) is a large endangered (Family Meliphagidae) endemic to the Samoa archipelago. Historically, the species occurred on the islands of Savai‘i and ‘Upolu in the independent nation of Samoa, and on the island of Tutuila in American Samoa, an unincorporated territory of the United States. Historical population estimates are unknown but the species was considered widespread in forested on Savai‘i and ‘Upolu. Currently the species is only known to occur on the islands of Savai‘i and ‘Upolu and the total population is estimated to be around 500 individuals. The range restriction and population declines are believed to be associated with the loss and degradation of Mao and high rates of nest failure and fledgling depredations. Mao habitat loss and degradation is related to human development (.e.g, agriculture and ), increase in the distribution and number of invasive plant species, increase in number of feral ungulates, and impacts from naturally occurring tropical cyclones (hurricanes). Increased nest failures and fledgling depredations are related to nest predation by introduced rats, depredation of juveniles by feral cats, and nest loss due to naturally occurring tropical cyclones (hurricanes). This Species Report evaluates the viability of the Mao, defined as the likelihood that it will persist over time. Viability is assessed by the resiliency, representation, and redundancy of the Mao populations. Each of these terms, and the Mao’s resultant viability, under current conditions are described below. Currently, there are no known Mao in American Samoa. Resiliency is the capacity of a population to withstand stochastic events. There are no historical population estimates, but the species was considered widespread in forested habitats on Savai‘i and ‘Upolu. Currently, the Savai‘i, ‘Upolu, and Tutuila populations have low to moderate, low, and no resiliency, respectively. The Tutuila population is believed extirpated and both the Savai‘i and ‘Upolu populations have experienced population restrictions and declines. These trends are expected to continue if the threats to the species continue at current levels or worsen. Representation is having one or more populations of a species occupying the full range of habitat types used by the species. Currently, the species is considered to have low to moderate representation due to range restrictions on Savai‘i and ‘Upolu and the extirpation of the Tutuila population. These range restrictions are expected to continue if the threats to the species continue at current levels or worsen. Redundancy is minimizing the risk of extinction of the species by establishing multiple populations across a landscape so some populations will always survive catastrophic events. Currently, the species is considered to have moderate redundancy as the species still occurs in two of the three original populations. Redundancy is expected to decrease in the future if threats to the species continue at current levels or worsen, potentially resulting in the extirpation of one or both of the two remaining populations.

iii

Table of Contents Cover Photo Credits ...... ii Suggested Citation ...... ii Primary Authors ...... ii Executive Summary ...... iii List of Figures ...... v List of Tables ...... vi Introduction ...... 1 Regulatory History ...... 1 Methodology ...... 1 Part 1. Life History and Historical Status ...... 4 Geography ...... 4 Species Description ...... 12 Life History...... 14 Historical Range, Distribution, and Population Status ...... 18 Part 2. Current Conditions and Species Status...... 20 Current Range and Distribution ...... 20 Factors Affecting Viability ...... 22 Resiliency, Representation, and Redundancy of the Species ...... 30 Resiliency of Current Populations ...... 30 Current Representation and Redundancy of the Species ...... 30 Literature Cited ...... 32

iv

List of Figures Figure 1. The three conservation biology principles of resiliency, redundancy, and representation, or the “3Rs”...... 2 Figure 2. Islands of the Samoan archipelago...... 5 Figure 3. Monthly mean precipitation for selected areas of the Samoan archipelago. All data from PCCSP (2019)...... 6 Figure 4. Monthly mean temperatures for selected areas of the Samoan archipelago. All data from PCCSP (2019)...... 7 Figure 5. Cyclone (hurricane) tracks for the central South Pacific from 1898 to 2019. All data from Knapp et al. (2010) and NOAA IBTrACS (2019)...... 7 Figure 6. Locations of roads in the Samoan archipelago...... 9 Figure 7. Vegetation land cover on American Samoa (from data by Meyer et al., 2017)...... 10 Figure 8. Vegetation land cover for Samoa (from data by LINZ, 2015a-i)...... 11 Figure 9. Photograph of an adult Mao by R. Stirnemann...... 13 Figure 10. Beak, wing, tarsus, and tail measurements from Mao in Western Samoa (Stirnemann et al., 2015a, p. 381)...... 13 Figure 11. Breeding cycle of the Mao (from data in Stirnemann et al., 2015b, pp.5-7) ...... 15 Figure 12. Generalized life stages of the Mao...... 17 Figure 13. The estimated historical range of the Mao...... 19 Figure 14. Sites where Mao where observed in Samoa from 1987 to 2000. (Land cover from MNRE, 2006, p. 3)...... 20 Figure 15. Locations in 2005-2006 where Mao were detected (●) and not detected (+) (MNRE 2006, p. 4; Tipamaa 2007, in litt., cited in Birdlife International 2012). Land cover from MNRE, 2006, p. 4...... 21 Figure 16. Estimated forest cover in Samoa from 1954 to 1999 (MNRE 2014, in litt.; FAO 2005, in litt.) Maps from Figures 4-6 of MNRE (2006, p. 6-7)...... 23

v

List of Tables Table 1. Census estimates of the American Samoa population on the islands of Tutuila, Ta’ū, Ofu, Olosega, and Swains...... 8 Table 2. Census estimates of the Samoa population by island and for the entire country...... 8 Table 3. Major land cover types for the Samoan archipelago. Area estimates in acres (hectares)...... 11 Table 4. Estimated acreages of vegetation types for the main islands of American Samoa. Area in acres (hectares)...... 12 Table 5. List of Gymnomyza species and their conservation status. All data obtained from Higgins et al. (2008, p. 670-671)...... 14 Table 6. Diet and foraging locations recorded for Gymnomyza species. All data obtained from Higgins et al. (2008, p. 670-671)...... 14 Table 7. Nest descriptions and nest location information for Gymnomyza species...... 16 Table 8. Estimated lifespans, sizes, and weights of in Australia. Lifespan estimates are based on banded wild ...... 17

vi

Introduction This document synthesizes the available information on the Mao and provides an assessment of the its current viability. The Mao or Ma‘oma‘o (Gymnomyza samoensis) is a large honeyeater (Family Meliphagidae) endemic to the Samoa archipelago (Figure 1). Historically, the species occurred on the islands of Savai‘i and ‘Upolu in the independent nation of Samoa and the island of Tutuila in American Samoa, an unincorporated territory of the United States (Watling 2001, p. 175). Currently the species is only known to occur on the islands of Savai‘i and ‘Upolu.

Regulatory History The U.S. Fish and Wildlife Service issued a final listing rule in 2016, in which the Service determine endangered status under the Act of 1973, as amended, for two endemic American Samoan land snails (Eua zebrina and Ostodes strigatus), the American Samoa distinct population segment of the Friendly Ground-dove, the Pacific Sheath-tailed Bat, (South Pacific subspecies) (Emballonura semicaudata semicaudata), and the Mao (Gymnomyza samoensis). The Mao was listed as endangered under the Endangered Species Act in 2016, and is also classified as endangered by the IUCN (USFWS 2016a, Birdlife International 2016). The effect of this regulation added these species to the List of Endangered and Threatened Wildlife. There are no additional regulatory documents for Mao, apart from a 2014 Candidate Notice of Review (CNOR) and a proposed rule for listing in 2016. (Links to the CNOR,and the proposed and final listing rules are at https://www.fws.gov/endangered/, then do a species search for Gymnomyza samoensis.

Methodology This Species Report provides an assessment of the Mao’s current viability, based on the best information available at this time, including peer-reviewed literature, gray literature (government, academic, business, and industry reports), and expert elicitation. Viability is the ability or likelihood of the species to maintain populations over time, i.e., likelihood of avoiding extinction. The viability of federally listed species is currently being assessed using the three conservation biology principles of resiliency, redundancy, and representation, or the “3Rs” (Figure 1; USFWS 2016b, p 12-16). The viability of Mao was evaluated by describing what the needs to be resilient, redundant, and represented, followed by comparing this assessment of what is needed for viability to the status of the bird in its current condition or its condition based on the most recent information. Our analysis follows the SSA Framework (Figure 1) developed by the USFWS (USFWS 2016b), in which viability is defined as the likelihood that a species will persist over time and is a product of resiliency, representation, and redundancy, otherwise known as “the three Rs” (USFWS 2016b, p. 12-13).

1

Figure 1. The three conservation biology principles of resiliency, redundancy, and representation, or the “3Rs”.

The definitions of the “3Rs” are defined below, and are used to infer the status of the species. • Resiliency is the capacity of a population or a species to withstand the more extreme limits of normal year-to-year variation in environmental conditions such as temperature and rainfall extremes, and unpredictable but seasonally frequent perturbations such as fire, flooding, and storms (i.e., environmental stochasticity). Quantitative information on the resiliency of a population or species is often unavailable. However, in the most general sense, a population or species that can be found within a known area over an extended period of time (e.g., seasons or years) is likely to be resilient to current

2

environmental stochasticity. If quantitative information is available, a resilient population or species will show enough reproduction and recruitment to maintain or increase the numbers of individuals in the population or species, and possibly expand the range of occupancy. Thus, resiliency is positively related to population size and growth rate, and may also influence the connectivity among populations. • Redundancy is having more than one resilient population distributed across the landscape, thereby minimizing the risk of extinction of the species. To be effective at achieving redundancy, the distribution of redundant populations across the geographic range should exceed the area of impact of a catastrophic event that would otherwise overwhelm the resilient capacity of the populations of a species. In the report, catastrophic events are distinguished from environmental stochasticity in that they are relatively unpredictable and infrequent events that exceed the more extreme limits of normal year-to-year variation in environmental conditions (i.e., environmental stochasticity), and thus expose populations or species to an elevated extinction risk within the area of impact of the catastrophic event. Redundancy is conferred upon a species when the geographic range of the species exceeds the area of impact of any anticipated catastrophic event. In general, a wider range of habitat types, a greater geographic distribution, and connectivity across the geographic range will increase the redundancy of a species and its ability to survive a catastrophic event. • Representation is having more than one population of a species occupying the full range of habitat types used by the species. Alternatively, representation can be viewed as maintaining the breadth of genetic diversity within and among populations, in order to allow the species to adapt to changing environmental conditions over time. The diversity of habitat types, or the breadth of the genetic diversity of a species, is strongly influenced by the current and historic biogeographical range of the species. Conserving this range should take into account historic latitudinal and longitudinal ranges, elevation gradients, climatic gradients, soil types, habitat types, seasonal condition, etc. Connectivity among populations and habitats is also an important consideration in evaluating representation.

The viability of a species is derived from the combined effects of the 3Rs. A species is considered viable when there are a sufficient number of self-sustaining populations (resiliency) distributed over a large enough area across the range of the species (redundancy) and occupying a range of habitats to maintain environmental and genetic diversity (redundancy) to allow the species to persist indefinitely when faced with annual environmental stochasticity and infrequent catastrophic events. Common ecological features are part of each of the 3Rs. This is especially true of connectivity among habitats across the range of the species. Connectivity sustains dispersal of individuals, which in turn greatly affects genetic diversity within and among populations. Connectivity also sustains access to the full range of habitats normally used by the species, and is essential for re-establishing occupancy of habitats following severe environmental stochasticity or catastrophic events (see Figure 1 for examples of overlap among the 3Rs). Another way the three principles are inter-related is through the foundation of population

3

resiliency. Resiliency is assessed at the population level, while redundancy and representation are assessed at the species level. Resiliency populations are the necessary foundation needed to attain sustained or increasing Representation and Redundancy within the species. The assessment of viability is not binary, in which a species is either viable or not, but rather on a continual scale of degrees of viability, from low to high. The health, number and distribution of populations were analyzed to determine the 3Rs and viability. In broad terms, the more resilient, represented, and redundant a species is, the more viable the species is. The current understanding of factors, including threats and conservation actions, will influence how the 3Rs and viability are interpreted for Mao.

Part 1. Life History and Historical Status Geography The Samoan archipelago is a chain of 13 islands and two atolls in the Pacific Ocean south of the equator (Figure 2), and is divided into the unincorporated United States Territory American Samoa to the southeast and the independent nation of Samoa to the northwest (Craig 2009, p. 5). These islands extend northwest to southeast over 298 miles (mi) (480 kilometers (km)) between 13o and 15o south latitude, and 168o to 172o west longitude (Goldin 2002, p. 4). They date to the early Pleistocene and were formed as hot-spot shield volcanoes, with the older islands located on the northwestern end of the chain (Thornberry-Ehrlich 2008, pp. 16, 28). American Samoa consists of five high islands and two atolls: Tutuila (the largest island; 54 square (sq.) mi (140 sq. km)); Aunu‘u (1 sq. mi (2 sq. km)) off the southeast end of Tutuila; Ofu and Olosega (3.5 sq. mi (9 sq. km)) separated by a narrow channel now spanned by a bridge; Ta‘ū (15 sq. mi (39 sq. km)); Rose Atoll (1.5 sq. mi (4 sq. km)), a National Wildlife Refuge) with two small uninhabited islands, Rose and Sand; and Swains Island (0.6 sq. mi (1.5 sq. km)), which is politically part of American Samoa, but geologically and biologically part of the Tokelau archipelago , which is a territory of New Zealand (Goldin 2002, pp. 5–6). These islands and atolls range in elevation from 3,170 feet (ft) (966 meters (m)) at Mt. Lata on Ta‘ū to 4 to 6 ft (1 to 2 m) above sea level at Rose Atoll (Goldin 2002, pp. 5-6). Samoa is located less than 100 mi (160 km) northwest of Tutuila Island, American Samoa, and consists of two large inhabited islands, ‘Upolu (424 sq. mi (1,100 sq. km)) and Savai‘i (703 sq. mi (1,820 sq. km)), and 8 small offshore islets, several of which are inhabited. Samoa lies between 13o to 14o south latitude and 170o to 173o west longitude and has a total land area of approximately 1,133 sq. mi (2,934 sq. km)) (Watling 2001, p. 26). The highest point in Samoa is Mt. Silisili on Savai‘i at 6,093 ft (1,857 m).

4

Figure 2. Islands of the Samoan archipelago.

Physical Environment The Samoan archipelago lies within the tropics, where it is hot, humid, and rainy year-round. The wet season is from October to May, with a slightly cooler and drier season from June through September (Figures 3 and 4). Temperatures average about 81.5o Fahrenheit (F) (27o Celsius (C)) at lower elevations. Rainfall averages 125 inches (in) (318 centimeters (cm)) annually at lower elevations, but can vary greatly depending upon topography, reaching 300 in (750 cm) or greater annually in the mountain areas. Tropical cyclones (called hurricanes north of the equator and east of 180o latitude, and typhoons north of the equator and west of 180o latitude) are a regular feature of natural disturbance in the Samoan Archipelago (Figure 5), and occur at intervals of between 1 to 13 years (Goldin 2002, p. 7). Land-Use In 2010/2011, the population of American Samoa and Samoa was 55,519 and 187,820, respectively (U.S. Census Bureau 2010; Samoa Bureau of Statistics (SBS) 2011, p. 14). Ninety- eight percent of the American Samoa population occurs on the island of Tutuila (Table 1). Census data collected since 1920 for American Samoa shows initial increases in populations on the Manu‘a Islands (Ta‘ū, Ofu and Olosega) and then declines by 2010 (Table 1). Tutuila shows a steadily increasing population from 1920 to 1980 and then some fluctuation in subsequent years. In Samoa, 76% of the population occurs on ‘Upolu (which includes Manono and Apolima islands, Table 2). Samoa shows a steadily increasing population from 1951 to 2011, with the population on Savai‘i remaining stable from 1981 to 2016 while ‘Upolu shows a steady increase during that period (Table 2).

5

Figure 3. Monthly mean precipitation for selected areas of the Samoan archipelago. All data from PCCSP (2019).

6

Figure 4. Monthly mean temperatures for selected areas of the Samoan archipelago. All data from PCCSP (2019).

Figure 5. Cyclone (hurricane) tracks for the central South Pacific from 1898 to 2019. All data from Knapp et al. (2010) and NOAA IBTrACS (2019).

7

Table 1. Census estimates of the American Samoa population on the islands of Tutuila, Ta’ū, Ofu, Olosega, and Swains.

American Island Year Samoa Tutuila Ta‘ū Ofu Olosega Swains Island 19201 8,056 6,185 1,155 361 355 - 19301,2 10,055 7,809 1,243 466 438 99 19401,2 12,908 10,164 1,588 500 509 147 19501,2 18,937 15,954 1,698 576 545 164 19602 20,051 17,250 1,661 605 429 106 19703 27,159 24,973 1,320 412 380 74 19803 47,283 45,524 1,138 345 249 27 19904 46,773 45,043 1,136 353 225 16 20005 57,291 55,876 873 289 216 37 20105 55,519 54,359 790 176 177 17 1Unkown (1956) , 2 Levin and Wright (1974), 3 U.S. Census Bureau (1982), 4 U.S. Census Bureau (1992), 5 U.S. Census Bureau (2010)

Table 2. Census estimates of the Samoa population by island and for the entire country.

Island Year1 Samoa ‘Upolu Savai‘i 1951 84,909 - - 1961 114,427 - - 1971 146,647 - - 1981 156,349 113,199 43,150 1991 161,298 116,248 45,050 2001 176,710 133,886 42,824 2011 187,820 143,418 44,402 1All data from Samoa Bureau of Statistics (SBS) (2011 p. 14, 17).

Due to the steep topography of the islands, human habitation is primarily located along the coastlines (Figure 6). On American Samoa, small-scale agriculture occurs inland from villages in former lowland rainforest areas on slopes that sometimes exceed 45o (Atkinson and Medeiros 2006, p. 4). Approximately 11% (5,456 acres (ac), 2,208 hectares (ha)) of American Samoa is classified as developed (i.e., urban areas, roads, infrastructure, and 6% (2,824 ac, 1,143 ha) of American Samoa is classified as agriculture (Table 3). The majority of the development and agriculture in American Samoa occurs on Tutuila where the largest human population resides (Table 1 and Table 3).

In Samoa approximately 3 percent (21,869 ac, 8,850 ha) is classified as developed and 25 percent (174,449 acres, 70,597 hectares) is agriculture. The majority of the development and agriculture occurs on ‘Upolu, where the majority of the human population resides (Table 2 and Table 3).

8

Figure 6. Locations of roads in the Samoan archipelago.

Biological Environment Mueller-Dombois and Fosberg (1998, p. 361) describes seven general categories of vegetation in the Samoan archipelago: • Littoral vegetation generally refers to vegetation that is directly exposed to salt spray and occurs along the coasts. It includes a variety of forest, scrub, and herbaceous communities (see Whistler (1992) and Mueller-Dombois and Fosberg (1998) for detailed descriptions). • Lowland rainforests occur below approximately 1,600-2,000 ft (500-600 m) elevation and include a variety of subtypes based on the dominate canopy species. • Montane rainforests occur above 1,600-2,000 ft (500-600 m) elevation and are generally cooler and rainier than lowland . • Cloud forest and cloud scrub is limited to Savai‘i, ‘Upolu, Ta‘ū and Olosega and are wetter than montane forests due to the persistence of clouds. • Wetland vegetation includes mangroves, swamp forest, coastal marsh, and montane marsh and bog areas. Montane marsh and bog areas are limited to Savai‘i and ‘Upolu. • Vegetation on recent volcanic surfaces includes scrubby woody vegetation, ferns, and grasses that may represent pioneer rainforest. • Modified vegetation includes all areas that were formerly dominated by native vegetation but were altered due to human activity (e.g., urban areas, agriculture areas, secondary forest, etc.).

Approximately 70% of American Samoa is forested, with the majority classified as lowland tropical rainforest (Figure 7; Tables 3 and 4). Montane rainforest makes up a smaller percent of

9 the forest areas and is primarily restricted to Ta‘u with a small amount on Tutuila (Figure 7, Table 4). Approximately 58% of Samoa is forested (Figure 8, Table 3). However, not all of this is native forest. It is estimated that only 360 ac (146 ha) of native lowland rainforests (below 2,000 ft or 600 m) remain on Savai‘i and ‘Upolu as a result of forest clearing (MNRE 2013, p. 47). Savai‘i still has extensive upland forests that are for the most part undisturbed and composed of native species (MNRE 2013, p. 40). However, upland forest on ‘Upolu is largely dominated by introduced species today (MNRE 2013, p. 12).

Figure 7. Vegetation land cover on American Samoa (from data by Meyer et al., 2017).

10

Figure 8. Vegetation land cover for Samoa (from data by LINZ, 2015a-i).

Table 3. Major land cover types for the Samoan archipelago. Area estimates in acres (hectares).

Samoa American Samoa Land cover1 Ofu/ Savai‘i ‘Upolu Minor Islands Tutuila Ta‘ū Olosega Developed 6,934 (2,806) 14,794 (5,987) 141 (57) 5,179 (2,096) 101 (41) 175 (71) Agriculture 77,905 (31,527) 96,037 (38,865) 507 (205) 2,664 (1,078) 42 (17) 119 (48) 281,593 124,830 Forest 722 (292) 23,262 (9,414) 2,797 (1,132) 8,300 (3,359) (113,957) (50,517) Scrub 29,734 (12,033) 17,957 (7,267) 171 (69) 1,564 (633) 52 (21) 1,873 (758) Grassland/ 16,131 (6,528) 21,730 (8,794) 37 (15) 440 (178) 35 (14) 625 (253) Herbaceous Mangrove 67 (27) 801 (324) 15 (6) 79 (32) 0 (0) 0 (0) Wetland 306 (124) 726 (294) 67 (27) 72 (29) 7 (3) 17 (7) Open Water 91 (37) 692 (280) 10 (4) 62 (25) 0 (0) 0 (0) Barren 7,811 (3,161) 247 (100) 49 (20) 432 (175) 94 (38) 128 (52) 1 All data from Meyer et al. (2017) and MNRE (2014).

11

Table 4. Estimated acreages of vegetation types for the main islands of American Samoa. Area in acres (hectares).

Type1 Subtype Tutuila Aunu‘u Ofu/Olesega Ta‘ū Herbaceous 26 (11) 22 (9) 1 (<1) 572 (231) Strand Littoral Littoral Forest 608 (246) 35 (14) 295 (119) 338 (137) Vegetation Littoral Scrub 117 (47) 4 (2) 9 (4) 17 (7) Pandanus Scrub 18 (7) 0 (0) 0(0) 0 (0) Coastal Marsh 7 (3) 34 (14) 7 (3) 7 (3) Wetland Mangrove Scrub 79 (32) 14 (6) 0 (0) 0 (0) Vegetation & Forest Swamp Forest 64 (26) 29 (12) 0 (0) 10 (4) Lowland - 20,507 (8,299) 0 (0) 480 (194) 761 (308) Rainforest Montane - 95 (38) 0 (0) 92 (37) 2,809 (1,137) Rainforest Cloud Forest & - 0 (0) 0 (0) 0 (0) 546 (221) Scrub Vegetation on Recent Volcanic - 223 (90) 0 (0) 0 (0) 0 (0) Surfaces Fern lands 3 (1) 2 (1) 13 (5) 34 (14) Managed Land 10,296 (4,167) 139 (56) 406 (164) 513 (208) Modified Secondary 0 (0) 53 (22) 1,696 (686) 5,421 (2,194) Vegetation Forest Secondary 1,214 (491) 14 (6) 37 (15) 78 (31) Scrub Open Water - 62 (25) 11 (4) 0 (0) 0 (0) Terrestrial - 431 (175) 18 (7) 93 (38) 129 (52) Non-vegetated 1 Data from Meyer et al. (2017). Species Description Appearance The Mao is a large honeyeater with dark varying from blackish on the head and breast to olive- green on the wings, tail, and body (Stirnemann et al. 2015a, p. 1). It has an olive-green stripe under the eye (Figure 9). The bill is long, curved, and black in adults. Males have blue- grey and brown eyes, and females have brown eyes only (Stirnemann et al. 2015b, p. 383). Males are significantly larger than females with respect to wing, bill, tarsus, and tail length, although there is considerable overlap in size (Stirnemann et al. 2015b, pp. 380–381). Juveniles have a shorter bill than adults, and eye color changes 2 months post-fledging (Stirnemann et al. 2015b, p. 383). The Mao is a very vocal species and makes a variety of loud distinctive calls with bouts of calling lasting up to a minute (Watling 2001, p. 174). Calls differ between sexes (Stirnemann et al. 2015b, p. 382).

12

Figure 9. Photograph of an adult Mao by R. Stirnemann.

) mm ) mm Length ( Length (

) ) mm mm Length ( Length (

Figure 10. Beak, wing, tarsus, and tail measurements from Mao in Western Samoa (Stirnemann et al., 2015a, p. 381).

13

Measurements and Mass The Mao is a large honeyeater approximately 11 to 12 in (28 to 31 cm) long (Higgins et al. 2008, p. 670). Morphometric data from the Samoa populations indicates that males are generally larger than females, with some overlap (Figure 10).

Systematics The Mao is one of three species in the genus Gymnomyza in the honeyeater family, Meliphagidae (Table 5). The species was first described in 1841 by Hombron and Jacquinot from a specimen collected in ‘Upolu, Samoa (Higgins et al. 2008, p. 670)

Table 5. List of Gymnomyza species and their conservation status. All data obtained from Higgins et al. (2008, p. 670-671).

Species Common Name Status Distribution Gymnomyza virdis Not Globally Threatened Fiji Gymnomyza samoensis Mao Endangered Samoa Gymnomyza aubryana Crow Honeyater Endangered New Caledonia

Life History Foraging Biology Diet The Mao forage on invertebrates, fruits and nectar and may also prey on small reptiles (Table 6; Higgins et al. 2008, p. 670-671; Butler and Stirnemann 2013, p. 30). Potential nectar sources include native and non-native species like Erythrina spp., Heliconia spp., etc. (Butler and Stirnemann 2013, p. 30).

Table 6. Diet and foraging locations recorded for Gymnomyza species. All data obtained from Higgins et al. (2008, p. 670-671).

Species Diet Foraging Location Nectar, Fruit, Invertebrates, Canopy, Subcanopy, Gymnomyza virdis Reptiles Occasionally on Ground Gymnomyza samoensis Nectar, Arthropods, Fruit Canopy and Subcanopy Gymnomyza aubryana Arthropods and Nectar Canopy, Mid-storey

14

Foraging Method and Location Mao forage by gleaning invertebrates from leaves, probe dead material and moss for invertebrates, and probe flowers for nectar (Higgins et al. 2008, p. 671; Butler and Stirnemann 2013, p. 30). They forage primarily in the canopy and subcanopy in forested areas, like other members of their genus (Table 6). Females will also forage for invertebrates under dead leaves on the forest floor to feed their fledglings (Butler and Stirnemann 2013, p. 30). Fledglings solicit food from the female by begging continually on the he forest floor (Butler and Stirnemann 2013, p. 28). Breeding Biology Figure 11 provides an overview of the breeding cycle of the Mao. The Mao’s breeding cycle is typical for with most breeding activities centered on the nest site. Once a nest is constructed the female lays and incubates the egg for 19-21 days (Stirnemann et al. 2015b, p.4- 7). The egg hatches after incubation and the emerging chick or hatchling is fed and cared for by the female for 22-24 days until it leaves the nest (i.e., fledging). The immature or juvenile birds is dependent on its parents for 8-10 weeks post-fledging. Specific details on the breeding characteristics of the species are outline below.

Figure 11. Breeding cycle of the Mao (from data in Stirnemann et al., 2015b, pp.5-7)

Breeding Season Based on observations of 10 nests, Mao breeding can occur over 9 to 10 months, although peak egg-laying appears to occur from late May to October, when rainfall levels are decreased (Stirnemann et al. 2015b; p.4-5). The Mao will re-nest if the first nest fails, but not if the first nesting attempt produces a chick (Stirnemann et al. 2015b; pp. 4–7). Nest Characteristics The Mao’s nest consists of young branches of various trees and contains little lining (Butler and Stirnemann 2013, p. 25). Nests are oval, cup-shaped, approximately 5.5 in (14 cm, n = 10) by 3.1 in (8 cm, n = 10), and are constructed in the junction of branches (Stirnemann et al. 2015b; p. 5). Heights of 10 nests ranged from 8 to 20 m above the forest floor, and percent nest cover ranged from 0 to 80% (Stirnemann et al. 2015b, p. 6). Nest descriptions and nest heights were similar among the species in the same genus (Table 7). Stirnemman et al. (2015b, p. 6) reported nests in the following native and introduced tree species: Planchonella garberi (Alaa, n = 1), Loranthus

15

samoensis (Native mistletoe, n = 1), Dysoxylum huntii (Maota, n = 1), Glochidion ramiflorum (Masame, n = 1), Fagraea berteroana (Pua lulu, n = 1), Hevea brasiliensis (Rubber tree, n = 2), and Dendrocnide harveyi (Salato, n = 2). The duration of the nest building process is unknown.

Table 7. Nest descriptions and nest location information for Gymnomyza species.

Species Nest Description Nest Location Gymnomyza virdisa Cup made of rootlets ~ 20m above ground, outer foliage Gymnomyza samoensisb Cup 14.2 ± 1.9 m (range 8-20 m, n = 10) Gymnomyza aubryanaa Cup ~ 20m above ground, branch fork aHiggins et al. 2008, p670-671; bStirnemann et al. 2015b, p. 5.

Egg Stage One off-white egg is produced per clutch (Stirnemann et al. 2015b; p. 5; n = 10). Higgins et al. (2008, p. 671) reports a clutch size of 2-3 eggs, however no information is reported on the number of nests observed or the original citation for this observation. The incubation period, based on two nest observations, is 19-21 days (Stirnemann et al. 2015b, p. 5) and only females incubate eggs. Males feed females during incubation, infrequently. Hatchling Stage Based on observations of six nests, nestling fledge 22-24 days after hatching (Stirnemann et al. 2015b, p. 6). The female is almost exclusively responsible for feeding the chick. Fledgling Stage Juveniles are dependent on adults for approximately 8 to 10 weeks post-fledging (Stirnemann et al. 2015b; pp. 4–7). The female is almost exclusively responsible for feeding the fledgling. Survivorship and Lifespan Figure 12 shows the main life stages of the Mao. The survival rates of the species at each of these life stages are important determinants of population viability. The sections below provide details on known survival rates for each of the stages and the overall lifespan of the species.

16

Figure 12. Generalized life stages of the Mao.

Breeding Success Nest survival was estimated to be 2.8% (±4.1% SE, n = 12) for a 42-day nesting period (egg and nestling stage; Stirnemann et al. 2015c, p.74). However, based on observations of 19 females over two breeding seasons, overall reproductive productivity was 0.34 juveniles per female per breeding season (Stirnemann et al. 2015c, p. 74). The inconsistency between low nest survival and overall reproductive productivity is believed to be related to repeated nesting attempts by Mao, with failed nests, over the breeding season (Stirnemann et al. 2015c, p.76). Juvenile and Adult Survival Strinemman et al. (2015c, p. 74) estimated 0.95 ± 0.07 annual survival and 100% annual survival for 19 monitored adult females and 12 monitored adult males. Juvenile survival was estimated to be 100% during the 8-10 week post-fledging parental care period. Estimates of juvenile survival to adulthood are not available. Lifespan Estimates of lifespan have not been determined for the Mao. Estimates for other species in the Meliphagidae family range from 6.8 to 15.3 years (Table 8).

Table 8. Estimated lifespans, sizes, and weights of honeyeaters in Australia. Lifespan estimates are based on banded wild birds.

Lifespan Size5 Weight5 Species Common Name (yrs) (cm) (g) melanops Yellow-tufted Honeyeater 15.34 16.5-21 14.2-31 Phylidonyris novaehollandiae3 New Holland Honeyeater 14.54 16-20 10-15 Lichenostomus fuscus Fuscous Honeyeater 14.14 13.5-17 12-24

17

Meliphaga lewinii Lewins Honeyeater 134 19-21.5 27.3-49.5 Lichenostomus chrysops Yellow-faced Honeyeater 12.14 15-17.5 12-20.5 Phylidonyris niger White-cheeked Honeyeater 10.41 16-20 15.5-25 Anthochaera phrygia Regent Honeyeater 9.114 20.4-24 33-45 Melithreptus brevirostris Brown-headed Honeyeater 72 11.5-14 12-17 Melithreptus lunatus White-naped Honeyeater 6.83q A 11.5-25 1Editor 2014, p. 668; 2Lane and Lonnon 1967, p. 33; 3Leishman 2012, p. 14; 4Geering 2002; 5Higgins et al., 2008, pp. 589, 597, 604, 607, 622, 667, 668, 673, 675.

Habitat Mao was thought to be primarily restricted to mature, well- developed, moist, mossy forests at upper elevations (Watling 2001, p. 175; Engbring and Ramsey 1989, p. 68), but has recently been observed at elevations ranging from 932 to 5,075 ft (284 to 1,547 m) and in including lowland rainforest, disturbed secondary forest, and montane rainforest (MNRE 2006, pp. 9–10). The birds use the mid- to upper-canopy levels of the forest and will also forage along forest edges and brushy forest openings (Engbring and Ramsey 1989, p. 68). The Mao has also been recorded visiting trees near the coast (Watling 2001, p. 175). Butler and Stirnemann (2013, p. 30) provide the following information about the Mao’s habitat use. The birds occur only in forested areas with a canopy layer, including modified habitat such as plantations where large trees also are present. They do not occur in logged areas with no large trees or canopy. Mao are primarily found in the high canopy layer, but also spend considerable time foraging on the trunks of trees and feeding on nectar sources near the ground (such as ginger (family Zingiberaceae)) and in low bushes (such as Heliconia spp.). The Mao selects territories with high tree species diversity and with appropriate nectar sources and a large tree from which the male sings. Trees near a commonly used singing tree are selected for nesting. Mao maintain a territory during breeding with both sexes defending the territory (Stirnemann et al. 2015b, p. 4-5). Territory sizes for the Mao are currently unknown. However, the (Gymnomyza aubryana) has an estimated territory size of 0.5 to 0.75 km2 (Higgins et al. 2008, p. 671). Historical Range, Distribution, and Population Status Mao is endemic to the islands of Savai’i and ‘Upolu, Samoa, and Tutuila Island, American Samoa (Figure 13; Engbring and Ramsey 1989, p. 68; Watling 2001, p. 174). It was once found throughout Savai‘i and ‘Upolu in Samoa, likely in forests ranging from the coast to mountain tops (MNRE 2006, p. 2). Bellingham and Davis (1988, p.124-125) recorded the species in lowland and montane forests on both islands in 1984 but found them more abundant in montane forests on Savai‘i and unlogged forest on ‘Upolu. Reed (1980, p. 158) also recorded the species as more abundant in high elevation forests in 1979. No population estimates were provided.

18

Figure 14 summarizes the records of Mao from 1987 to 2000. These records show the species largely restricted to native forest on both islands. Little is known about the historical distribution and population status of the species on Tutuila. It was observed during U.S. Exploring Expedition in 1839 (Amerson et al. 1982a, p. 72) but apparently no specimens were collected (Banks 1984, p. 166). Three specimens were collected on Tutuila by the Whitney South Seas Expedition in February 1924 (two males with enlarged testes and a potential young female; Banks 1984, p. 166). The species was not recorded during surveys by Amerson et al. in 1975 and 1976 (1982a, p. 72). However, P. Bruner recorded an observation of the species on Tutuila in 1977 (Pratt et al. 1987, p. 280). Extensive surveys of Tutuila in 1986 and from 1992 to 1996 failed to detect the species (Engbring and Ramsey 1989, p 68; Freifeld et al 2004, p. 104).

Figure 13. The estimated historical range of the Mao.

19

Figure 14. Sites where Mao where observed in Samoa from 1987 to 2000. (Land cover from MNRE, 2006, p. 3).

Part 2. Current Conditions and Species Status Current Range and Distribution Mao is currently found only on the islands of Savai‘i and ‘Upolu in Samoa (MNRE 2006, p. 2). Birdlife International (2016) reported that the population size is around 500 individuals, however, extensive surveys have not been conducted. The following sections provide information on population trends in each of the historical populations.

Savai‘i and ‘Upolu, Samoa Historically the Mao was reported as common in undisturbed upland forests (foothill, montane, and cloud forests above 1,970 ft (600 m)) of ‘Upolu and Savai‘i (Bellingham and Davis 1988, p. 124). A decline in distribution was observed in the 1990s following a period in which several powerful cyclones (hurricanes) hit Samoa: Tusi (1987), Ofa (1990), and Val (1991) (Lovegrove et al. 1992,p. 26; MNRE 2006, pp. 2, 4). Otherwise, no detailed surveys of the Mao were conducted before 2005, and little information exists regarding changes in abundance and distribution (MNRE 2006, p. 2). Surveys conducted in 2005– 2006 found Mao at seven sites on ‘Upolu and Savai‘i in upland forested habitat (Figure 15), yielded a rough estimate of 500 individuals and indicated that numbers are declining (MNRE 2006, p. 4; Tipamaa 2007, in litt.,

20 cited in Birdlife International 2012). The Rapid Assessment of Upland Savai‘i, Samoa, conducted in 2012, detected small numbers of the Mao at two sites on the island (Atherton and Jefferies 2012, p. 14). None of the recent surveys (Atherton and Jeffries 2012, p. 110; MNRE 2013, p. 87) or studies (Stirnemann 2015a) has yielded an updated population estimate. However, researchers observed that the species is rarer than previously thought and recommended that comprehensive surveys be conducted to generate a new population estimate (Stirnemann 2015b, in litt).

Tutuila, American Samoa The Mao is likely extirpated from Tutuila Island in American Samoa (Freifeld 1999, p. 1,208). Surveys conducted on Tutuila Island in 1986 and from 1992 to 1996 did not detect the Mao (Engbring and Ramsey 1989; p. 68; Freifeld et al 2004, p. 104). Given that the species is noisy and conspicuous, it is believed unlikely that a population on Tutuila was missed during those surveys (Engbring and Ramsey 1989; p. 68). More recent surveys conducted by DMWR, in forested habitats likely to support Mao, also failed to detect their presence, further indicating the likelihood that the species no longer occurs on Tutuila (MacDonald 2015, in litt.).

Figure 15. Locations in 2005-2006 where Mao were detected (●) and not detected (+) (MNRE 2006, p. 4; Tipamaa 2007, in litt., cited in Birdlife International 2012). Land cover from MNRE, 2006, p. 4.

21

Factors Affecting Viability This section describes the various factors that may be affecting the viability of the current Mao populations and may have led to the observed population and range declines from historical levels. Habitat Loss and Degradation The following sections provide information on habitat loss and degradation due to development, introduced ungulates, and introduced plants. This section also provides information on efforts to restore and improve potential Mao habitat. Development Before the arrival of Polynesians, approximately 3,000 years ago, the Samoan archipelago was likely covered by forest, except for recent volcanic flows, marshes, and steep and windswept regions of the islands (Amerson et al. 1982b, p. 20; Mueller-Dombois and Fosberg 1998, p. 360). Thousands of year of subsistence agriculture and, more recent, commercial agriculture has resulted in the alteration and great reduction in area of forests at lower elevations in the Samoan archipelago (Whistler 1994, p. 40; Mueller-Dombois and Fosberg 1998, p. 361; Whistler 2002, pp. 130–131). On the island of Tutuila, American Samoa, agriculture and urban development covers approximately 24 percent of the island, and up to 60 percent of the island contains slopes of less than 30 percent where additional land clearing is feasible (ASCC 2010, p. 13; DWMR 2006, p. 25). Farmers are increasingly encroaching into some of the steep forested areas as a result of suitable flat lands already being occupied with urban development and agriculture (ASCC 2010, p. 13). Consequently, agricultural plots have spread from low elevations up to middle and some high elevations on Tutuila. In Samoa, the amount of forested area declined from 74 to 40 percent of total land area between 1954 and 1990 (Figure 16; MNRE 2014, in litt.) and continued to decline by roughly 2 percent or 7,400 ac (3,000 ha) per year from 1990 to 2000 (Figure 16, FAO 2005, in litt.). The clearing of land for commercial agriculture has been the leading cause of deforestation in Samoa (Whistler 2002, p. 131). The transition from subsistence agriculture to cash crops for export (e.g., taro, bananas, cacao), coupled with rapid population growth and new technologies, led to increased forest clearing in Samoa (Paulson 1994, pp. 326–332; Whistler 2002, pp. 130–131). Shifting or slash- and-burn cultivation is also a concern in upland forest as farmers use roads from heavily logged lowland forests to gain access to formerly inaccessible land (MNRE 2006, p. 5). For example, there is much concern about potential forest loss because of road that has been bulldozed into the cloud forest (above 3,280 ft (1,000 m)) on Savai‘i, apparently illegally (Atherton and Jeffries 2012, p. 16). Logging has also been a cause of forest loss in Samoa. Between 1978 and 1990, 20 percent of all forest losses in Samoa were attributable to logging, with 97 percent of the logging having occurred on Savai’i (Government of Samoa 1998 in Whistler 2002, p. 132). These logging activities have slowed down in recent years due to the reduction of forest accessible to logging,

22 but is an ongoing problem on Savai’i despite years of effort to phase it out (MNRE 2006, p. 5; Atherton and Jeffries 2012, p. 17). Today, only 360 ac (146 ha) of native lowland rainforests (below 2,000 ft or 600 m) remain on Savai‘i and ‘Upolu as a result of logging, agricultural clearing, residential clearing (including relocation due to tsunami), and natural causes such as rising sea level and cyclones (hurricanes) (MNRE 2013, p. 47). On ‘Upolu, direct or indirect human influence has caused extensive damage to native forest habitat above 2,000 ft (600 m) elevation (MNRE 2013, p. 13). Although upland ‘Upolu is forested, almost all of the upland forests are largely dominated by introduced species today (MNRE 2013, p. 12). Savai‘i still has extensive upland forests that are for the most part undisturbed and composed of native species (Figure 16; MNRE 2013, p. 40). As a result, forest clearance as a result of development is considered an ongoing threat to the Mao (MNRE 2006, p. 5).

1954 1987 1999 Figure 16. Estimated forest cover in Samoa from 1954 to 1999 (MNRE 2014, in litt.; FAO 2005, in litt.) Maps from Figures 4-6 of MNRE (2006, p. 6-7).

Invasive Plants Nonnative plants are known to have invaded native ecosystems in American Samoa and Samoa, with documented adverse impacts to native forests (Space and Flynn 2000, pp. 5, 12; Space and Flynn 2002, pp. 4–5; Whistler 2002, p. 122; Atkinson and Medeiros 2006, pp. 17–18; Craig 2009, pp. 94, 98; ASCC 2010, p. 22; NPSA 2012, in litt.; Atherton and Jeffries 2012, p. 103; Butler and Stirnemann 2013, p. 30; MNRE 2013, p. 29). The native flora of the Samoan archipelago (plant species that were present before humans arrived) consisted of approximately 550 taxa, 30 percent of which were endemic (species that occur only in the American Samoa and Samoa) (Whistler 2002, p. 8). An additional 250 plant species have been introduced intentionally or accidentally and have become naturalized, with 20 or more of these considered invasive or potentially invasive in American Samoa (Whistler 2002, p. 8; Space and Flynn 2000, pp. 23–24). Of these approximately 20 or more nonnative pest plant species, at least 10 have altered or have the potential to alter the habitat of the Mao (Atkinson and Medeiros 2006, p. 18; Craig 2009, pp. 94, 97–98; ASCC 2010, p. 15). Nonnative plants have been documented to degrade native ecosystems in the Pacific island through the following means: (1) modifying the availability of light through alterations of the

23 canopy structure; (2) altering soil–water regimes; (3) modifying nutrient cycling; (4) converting native- dominated plant communities to nonnative plant communities through with native species; and (5) increasing the frequency of landslides and erosion (Smith 1985, pp. 217– 218; Cuddihy and Stone, 1990, p. 74; Matson 1990, p. 245; D’Antonio and Vitousek 1992, p. 73; Vitousek et al. 1997, pp. 6–9; Atkinson and Medeiros 2006, p. 16). Nonnative plant species can also exploit the disturbance caused by other natural and human caused factors, such as cyclones (hurricanes), agriculture and development, and feral ungulates, and thus, in combination reinforce or exacerbate their negative impacts to native habitats. The following list provides a brief description of the nonnative plants that may have the greatest negative impacts to the native forest habitat for the Mao in American Samoa (Space and Flynn 2000, pp. 23–24; Craig 2009, pp. 94, 96–98; ASCC 2010, p. 15): • Adenanthera pavonina (red bean tree, coral bean tree, lopa), native to India and Malaysia, is a medium-sized tree up to 50 ft (15 m) high that invades intact forests as well as disturbed sites, and can quickly form large stands (GISD 2006). In American Samoa, it is invasive in secondary forests, but also has the ability to become more widely established on Tutuila and the Manua Islands (Space and Flynn 2000, p. 4). It is considered to have negative impacts on the native forests in American Samoa because the trees produce large quantities of seed, grow on a variety of soils, and can overtop many native trees and eventually form monotypic stands (Space and Flynn 2002, p. 5). • Castilla elastica (Mexican rubber tree, pulu mamoe), native to tropical America, is a medium-sized tree 15 to 30 ft (5 to 10 m) high that can invade intact forest where it reproduces prolifically and can crowd out native species (NPSA 2012, in litt.). It has displaced significant areas of lowland forest in Samoa, and is now considered to be an important threat to native forests in American Samoa (Atkinson and Medeiros 2006, p. 18). • Cinnamomum verum (cinnamon, tinamoni), native to south Asia, is a fast-growing, medium-sized tree up to 30 ft (9 m) high with aromatic bark and leaves. It forms dense root mats that inhibit establishment of other plants, and can shade out other tree species and thus create monotypic stands. On Tutuila, it is actively spreading in the ridge forests of Mt. Matafao, Matuu, and Maloata (Space and Flynn 2000, p. 4; NPSA 2012, in litt.). • The shrub Clidemia hirta (Koster's curse), native to the New World from Mexico to Argentina, grows to be 6.6 ft (2 m) in height, forms a dense understory, shades out native plants, and prevents their regeneration (Wagner et al. 1985, p. 41; Smith 1989, p. 64). On Ta‘ū, it has become a serious problem in the unique summit scrub community (Whistler 1992, p. 22). • Falcataria moluccana (albizia, tamaligi), native to Moluccas, New Guinea, New Britain, and the Solomon Islands, is a tree that can reach 131 ft (40 m) in height and has a wide- spreading canopy. It grows rapidly and outcompetes slow-growing native trees by reducing light availability, and its abundant, high-nutrient litter alters soil chemistry (GISD 2008). Its shallow root system may lead to soil instability and landslides (Atkinson and Medeiros 2006, p. 17).

24

• Funtumia elastica (African rubber tree, pulu vao), is a medium-sized tree up to 100 ft (30 m) tall native to tropical Africa (USDA 2006). This tree is invasive because of its “parachute seeds” that can disperse long distances and germinate in sunny or shady conditions (Whistler 2002, p. 122). Funtumia has become a dominant subcanopy and understory tree in the western half of ‘Upolu where it can form monotypic forests (Pearsall and Whistler 1991, p. 30). It is also established and becoming dominant on eastern Savai‘i (Whistler 2002, p. 122). This species has the potential to become a major problem in American Samoa due to its proximity and the volume of traffic with Samoa (Space and Flynn 2000, p. 12). • Leucaena leucocephala (wild tamarind, lusina, fua pepe), a shrub native to the neotropics, is a nitrogen-fixer and an aggressive competitor that often forms the dominant element of the vegetation (Geesink et al. 1999, pp. 679–680). It crowds out native species and resprouts vigorously after cutting, and seeds can remain viable for 10 to 20 years (Craig 2009, p. 98). • Merremia peltata (Merremia, fue lautetele), is an indigenous, sprawling, or high-climbing vine that can invade areas following disturbances such as land-clearing and cyclones (hurricanes). This fast-growing vine can smother plantation and forest trees (Craig 2009, p. 98). • Mikania micrantha (mile-a-minute vine, fue saina), native to tropical America, is a scrambling or climbing herbaceous vine, that retards forest regeneration with its smothering growth (Whistler 1994, p. 42). This sun-loving, shade-intolerant vine is a major pest of plantations and forests on all major American Samoa islands (Space and Flynn 2000, p. 5; Craig 2009, p. 94). • Psidium cattleianum (strawberry guava, kuava) is a tall shrub or small tree that forms dense stands in which few other plants can grow, displacing native vegetation through competition. The fruit is eaten by feral pigs and birds that disperse the seeds throughout the forest (Smith 1985, p. 200; Wagner et al. 1985, p. 24). It is thought to have been cultivated in American Samoa for more than 40 years and has become naturalized in lowland rainforest on western Tutuila. • Spathodea campanulata (African tulip, faapasi), native to tropical Africa, is a large tree up to 80 ft (24 m) or more in height with showy red-orange tulip-like flowers and pods containing hundreds of wind-dispersed seeds (PIER 2013). It is particularly invasive in low-to mid-elevation forests, and can spread in open agricultural land, waste areas, and intact native forest, forming dense stands that shade out other vegetation (GISD 2010). Mao forage and occasionally nest in modified habitat, such as plantation areas where nonnative trees provide nectar and nesting habitat (e.g., Falcataria moluccana). However, these habitats lack the high tree-species diversity, which Mao are believed to prefer, and may also place the species at a greater risk of predation by nonnative predators that may inhabit these forests (Butler and Stirnemann 2013, p. 30).

25

Introduced Ungulates Feral pigs (Sus scrofa) and feral cattle (Bos taurus) are both factors in forest degradation in Samoa. Feral pigs are known to cause deleterious impacts to processes and functions throughout their worldwide distribution (Aplet et al. 1991, p. 56; Anderson and Stone 1993, p. 201; Campbell and Long 2009, p. 2,319). Feral pigs are extremely destructive and have both direct and indirect impacts on native plant communities. Pigs are a major vector for the establishment and spread of invasive, nonnative plant species by dispersing plant seeds on their hooves and fur, and in their feces (Diong 1982, pp. 169–170, 196–197), which also serve to fertilize disturbed soil (Siemann et al. 2009, p. 547). In addition, pig rooting and wallowing contributes to erosion by clearing vegetation and creating large areas of disturbed soil, especially on slopes (Smith 1985, pp. 190, 192, 196, 200, 204, 230–231; Stone 1985, pp. 254– 255, 262– 264; Tomich 1986, pp. 120– 126; Cuddihy and Stone 1990, pp. 64– 65; Aplet et al. 1991, p. 56; Loope et al. 1991, pp. 18–19; Gagne and Cuddihy 1999, p. 52; Nogueira-Filho et al. 2009, p. 3,681; CNMI-SWARS 2010, p. 15; Dunkell et al. 2011, pp. 175–177; Kessler 2011, pp. 320, 323). Erosion resulting from rooting and trampling by pigs impacts native plant communities by contributing to watershed degradation and alteration of plant nutrient status, and increasing the likelihood of landslides (Vitousek et al. 2009, pp. 3,074–3,086; Chan-Halbrendt et al. 2010, p. 251; Kessler 2011, pp. 320–324). In the Hawaiian Islands, pigs have been described as the most pervasive and disruptive nonnative influence on the unique native forests, and are widely recognized as one of the greatest current threats to Hawai‘i’s forest ecosystems (Aplet et al. 1991, p. 56; Anderson and Stone 1993, p. 195). In American Samoa, feral pigs negatively affect forested habitats. Feral pigs have been present in American Samoa since humans first settled the islands (American Samoa Historic Preservation Office 2015, in litt.). In the past, hunting pressure kept their numbers down, however, increasing urbanization and increasing availability of material goods has resulted in the decline in the practice of pig hunting to almost nothing (Whistler 1992, p. 21; 1994, p. 41). Feral pigs are moderately common to abundant in many forested areas, where they spread invasive plants, damage understory vegetation, and destroy riparian areas by their feeding and wallowing behavior (DMWR 2006, p. 23; ASCC 2010, p. 15). Feral pigs are a serious problem in the National Park of American Samoa (NPSA) because of the damage they cause to native vegetation through their rooting and wallowing (Whistler 1992, p. 21; 1994, p. 41; Hoshide 1996, p. 2; Cowie and Cook 1999, p. 48; Togia pers. comm. in Loope et al. 2013, p. 321). Such damage to understory vegetation is likely to reduce foraging opportunities for the Mao. Pig densities have been reduced in some areas by snaring and hunting, but remain high in other areas (ASCC 2010, p. 15). In Samoa, feral pigs are present throughout lowland and upland areas on Savai‘i, and are considered to have a negative impact on the ecological integrity of upland forests of Savai‘i, an important conservation area for the Mao and other rare species (Atherton and Jeffries 2012, p. 17). During recent surveys, feral pig activity was common at most sites in upland forests on Savai‘i, and was even detected at the upper range of the Mao at an elevation of 4,920 ft (1,500 m) (Atherton and Jefferies 2012, pp. 103, 146).

26

Finally, significant numbers of feral cattle (Bos taurus) were present in an upland site on Savai‘i where their trampling had kept open grassy areas within forested flats, and where Mao had previously been observed (Atherton and Jeffries 2012, pp. 103– 105). Trampling in forested areas damages understory vegetation and is likely to reduce foraging opportunities for Mao as well as provide vectors for invasion by nonnative plants. In addition, feral cattle consume tree seedlings and browse saplings, and combined with undergrowth disturbance, prevent forest regeneration, subsequently opening the forest to invasion by nonnative species (Cuddihy 1984, p. 16).

Conservation Efforts to Reduce Habitat Loss and Degradation American Samoa – The National Park of American Samoa (NPSA) was established to preserve and protect the tropical forest and archaeological and cultural resources, to maintain the habitat of flying foxes, to preserve the ecological balance of the Samoan tropical forest, and, consistent with the preservation of these resources, to provide for the enjoyment of the unique resources of the Samoan tropical forest by visitors from around the world (Public Law 1988; Public Law 2006). Under a 50-year lease agreement between local villages, the American Samoa Government, and the Federal Government, approximately 8,000 ac (3,240 ha) of forested habitat on the islands of Tutuila, Ta‘ū, and Ofu are protected and managed (ASG-NPSA Lease Agreement 1993). Several programs and partnerships to address the threat of nonnative plant species have been established and are ongoing in American Samoa. Since 2000, the NPSA has implemented an invasive plant management program that has focused on monitoring and removal of nonnative plant threats. The nonnative plant species prioritized for removal include the following: Adenanthera pavonina or lopa, Castilla elastica or pulu mamoe, Falcataria moluccana or tamaligi, Leucaena leucocephala or lusina, and Psidium cattleianum or strawberry guava (Togia 2015, in litt.). In particular, efforts have been focused on the removal of the tamiligi from within the boundaries of the NPSA as well as in adjacent areas (Hughes et al. 2012). The thrip Liothrips urichi is an insect that was introduced to American Samoa in the 1970s as a biocontrol for the weed Clidemia hirta (Tauiliili and Vargo 1993, p. 59). This thrip has been successful at controlling Clidemia on Tutuila. Though Clidemia is still common and widespread throughout Tutuila, thrips inhibit its growth and vigor, preventing it from achieving ecological dominance (Cook 2001, p. 143). In 2004, the American Samoa Team (ASIST) was established as an interagency team of nine local government and Federal agencies. The mission of ASIST is to reduce the rate of invasion and impact of invasive species in American Samoa with the goals of promoting education and awareness on invasive species and preventing, controlling, and eradicating invasive species. In 2010, the U.S. Forest Service conducted an invasive plant management workshop for Territorial and Federal agencies, and local partners (Nagle 2010). More recently, the NPSA produced a field guide of 15 invasive plants that the park and its partners target for early detection and response (NPSA 2012, in litt.).

27

In 1996, the NPSA initiated a feral pig control program that includes fencing and removal of pigs using snares in the Tutuila Island and Ta‘ū Island Units. Two fences have been constructed and several hundred pigs have been removed since 2007 (Togia 2015, in litt.). The program is ongoing and includes monitoring feral pig activity twice per year and additional removal actions as needed (Togia 2015, in litt.). Samoa – The Mt. Vaea Ecological Restoration Project surveyed and mapped the presence of native bird and plant species and invasive plant species within lowland forest habitat of the 454- ac (183-ha) Mt. Vaea Scenic Reserve on ‘Upolu, Samoa (Bonin 2008, pp. 2–5). The project was envisioned as the first demonstration project of invasive species management and forest restoration in Samoa. Phase I of the project resulted in the development of a restoration plan recommending removal of five priority invasive plant species and planting of native tree species (Bonin 2008, pp. viii, 24). Phase 2 of the project resulted in identifying techniques for treatment of two problematic rubber tree species (Castilla elastica or pulu mamoe and Funtumia elastica or pulu vao) and replanting areas with native tree species (Bonin 2010, pp. 20–21). Predation The following sections provide information about predation by rats (Rattus spp.) and feral cats (Felis catus) and existing efforts to address these stressors. Rats Nest predation by rats has negative impacts on many island birds, including the Mao (Atkinson 1977, p. 129; 1985, pp. 55–70; Butler and Stirnemann 2013, p. 29; O’Donnell et al. 2015, pp. 24–26). Three species of rats occur in American Samoa and are likely to be present on the islands of Ofu and Olosega: the Polynesian rat, Norway rat, and black rat (Atkinson 1977, p. 38; DMWR 2006, p. 22; Caruso 2015, in litt.). Rats have been identified as the main cause of decline in the closely related Gymnomyza aubryana in New Caledonia (MNRE 2006, p. 8). Butler and Stirnemann (2013, p. 29) captured footage of one nest depredation event on Mao by a black rat (Rattus rattus), which took an egg. The rat gained access to the egg by jumping on the incubating female’s back from the branch above, driving the female off the nest. Combined with the disappearance of two females during the breeding season, this footage suggests that adult females are potentially vulnerable to predation on the nest at night, while they are incubating (Butler and Stirnemann 2013, p. 31), a phenomenon documented or suspected in other island bird species, which lack innate behavioral defenses against nonnative mammalian predators (see for example Robertson et al. 1994, p. 1,084; Armstrong et al. 2006, p. 1,034; VanderWerf 2009, p. 741). This potential bias toward predation of females has the potential to create a skewed sex ratio in Mao populations (Robertson et al. 1994, pp. 1,083–1,084). The location of Mao nests affects their vulnerability to predation by rats. Nests in close proximity to plantation habitats, where rats are most abundant, are particularly susceptible and experience low reproductive success (Butler and Stirnemann 2013, p. 31). Nests within 160 ft (50 m) of a plantation are 40 percent more likely to be depredated than nests in forested areas farther from plantations (Butler and Stirnemann 2013, p. 31). Cats 28

Predation by cats has been directly responsible for the extinction of numerous birds on oceanic islands (Medina et al. 2011, p. 6). Native mammalian carnivores are absent from oceanic islands because of their low dispersal ability, but once introduced by humans, they can become significant predators on native such as seabirds and land birds that are not adapted to predation by terrestrial carnivores (Nogales et al. 2013, p. 804; Scott et al. 1986, p. 363; Ainley et al. 1997, p. 24; Hess and Banko 2006, in litt.). The considerable amount of time spent on the ground (up to 7 days) and poor flight ability of Mao chicks post- fledging increases the risk of predation by feral cats (Butler and Stirnemann 2013, p. 28). Evidence of feral cat presence exists in montane forests and along an elevational gradient on Savai‘i (Atherton and Jeffries 2012, pp. 76, 103). Predation by feral cats has been suggested as a contributing factor in the Mao’s extirpation from Tutuila (Stirnemann 2015b, in litt.). However, feral cats have not commonly been observed in native forest areas on Tutuila, though they have been observed in remote and forested areas on Ta‘ū (Badia 2014, in litt.; Arcilla 2015, in litt.). Conservation Efforts to Reduce Predation Rats – A project to restore habitat for the Mao and other priority species by removing the threat of predation by the Polynesian rat (R. exulans) was attempted on the uninhabited islands of Nu‘utele (267 ac (108 ha)) and Nu‘ulua (62 ac (25 ha)) off the eastern end of ‘Upolu, Samoa (Tye 2012, in litt). The demonstration project aimed to eradicate the Polynesian rat from both islands through aerial delivery of poison baits. Post-project monitoring detected rats on Nu‘utele, suggesting that rats survived the initial eradication effort or were able to recolonize the island (Tye 2012, in litt.). Cats – There are no known projects to reduce feral cat predation of Mao. Disease Avian malaria and pox have been important factors in the decline of Hawai‘i’s avifauna (Warner 1968, Van Riper et al. 1986). Field and laboratory investigations suggest that avian malaria may be indigenous and non-pathogenic in American Samoa and, therefore, is unlikely to affect bird populations (Jarvi et al. 2003, p. 636; Seamon 2004, in litt.). The best available information does not indicate there are other diseases affecting the Mao populations in Samoa (MNRE 2006, p. 8). Competition No information is available on competition between other species and the Mao. Natural Events

Tropical Cyclones (Hurricanes) Tropical Cyclones (hurricanes) are a common natural disturbance in the tropical Pacific and have occurred in the Samoan archipelago with varying frequency and intensity (Figure 5). Although severe storms are a natural disturbance with which the Mao has coexisted for millennia, such storms can exacerbate the threats to its remaining small, isolated populations by, at least temporarily, damaging or redistributing habitat and food resources for the birds and causing

29 direct mortality of individuals (Wiley and Wunderle 1993, pp. 340–341; Wunderle and Wiley 1996, p. 261). Two storms in the 1990s, Cyclones Ofa (1990) and Val (1991), severely damaged much of the remaining forested habitat in Samoa, reducing forest canopy cover by 73 percent (MNRE 2006, pp. 5, 7). In addition, Cyclone Evan struck Samoa in 2012 causing severe and widespread forest damage, including defoliation and downed trees in 80 to 90 percent of the Reserves and National Parks on ‘Upolu (Butler and Stirnemann 2013, p. 41). Secondary forests were also severely damaged by the storm, and most trees in the known Mao locations were stripped of their leaves, fruits, and flowers (Butler and Stirnemann 2013, p. 41).

Resiliency, Representation, and Redundancy of the Species Resiliency of Current Populations Savai‘i, Samoa The Mao was once found throughout Savai‘i likely in forests ranging from the coast to mountain tops (MNRE 2006, p. 2). Historical population estimates are not available but the species was considered common but patchily distributed in high elevation forests in 2005 (Pratt and Mittermeirer 2016, p. 226). Surveys conducted in 2012 detected small numbers at two sites (Atherton and Jefferies 2012, p. 14), indicating the population may have declined. In addition, reductions in the distribution of the population were noted in the 1990s and 2000s (Lovegrove et al. 1992, p. 26; MNRE 2006, pp. 2, 4). Based on the available information, we estimate the resiliency of the Savai‘i Mao population to be moderate due to an apparent decline in the population size and reduction in range. ‘Upolu, Samoa The Mao was once found throughout ‘Upolu likely in forests ranging from the coast to mountain tops (MNRE 2006, p. 2). Historical population estimates are not available but the species was considered common in preferred habitat in 1984 (Bellingham and Davis 1988, p. 124). Subsequent surveys indicate the population may have declined (Lovegrove et al. 1992, p. 26). In addition, reductions in the distribution of the population were noted in the 1990s and 2000s (Lovegrove et al. 1992, p. 26; MNRE 2006, pp. 2, 4). Based on the available information, we estimate the resiliency of the ‘Upolu Mao population to be low to moderate due to an apparent decline in the population size and reduction in range. Tutuila, American Samoa The Mao is believed to be extirpated from Tutuila (Freifeld 1999, p. 1,208). Therefore, this population has no resiliency.

Current Representation and Redundancy of the Species Species Representation

30

Historically the Mao was found on the islands of Savai‘i and ‘Upolu, Samoa, and Tutuila Island, American Samoa (Engbring and Ramsey 1989, p. 68; Watling 2001, p. 174). The Mao has not been observed on Tutuila since 1977 and is likely absent from that island (Freifeld 1999, p. 1,208; MacDonald 2015, in litt.). In addition, the Mao was once found throughout Savai‘i and ‘Upolu (Samoa) likely in forests ranging from the coast to mountain tops (MNRE 2006, p. 2). Distribution declines were noted in the 1990s and 2000s (Lovegrove et al. 1992, p. 26; MNRE 2006, pp. 2, 4) and the species appears restricted to forested areas at higher elevations (Figure 15). Based on the available information, we estimate the representation of the Mao to be low to moderate due to loss of the Tutuila population and restricted altitudinal ranges on the remaining islands. Species Redundancy Historically the Mao was found on the islands of Savai‘i and ‘Upolu, Samoa, and Tutuila Island, American Samoa (Engbring and Ramsey 1989, p. 68; Watling 2001, p. 174). The Mao has not been observed on Tutuila since 1977 and is likely absent from that island (Freifeld 1999, p. 1,208; MacDonald 2015, in litt.). While there are multiple locations of Mao on Savai‘i and ‘Upolu, each of these islands represents a single population. Historically, there were three populations of Mao. Currently there are only two populations, and those two populations are reduced in their area of occupancy and in number of individuals. Based on the available information, we estimate the redundancy of the Mao to be moderate due to loss of the Tutuila population.

31

Literature Cited Ainley, D.G., R. Podolsky, DeForest, L., and G. Spencer. 1997. New insights into the status of the Hawaiian petrel on Kauai. Colonial Waterbirds 20: 24-30. American Samoa Historic Preservation Office. 2015, in litt. Cultural history of American Samoa, http://www.ashpo.org/index.php/history.html?tmpl=component&print=1&page=, accessed 24 APR 2015. Amerson, A.B., Jr., W.A. Whistler, and T.D. Schwaner. 1982a. Wildlife and wildlife habitat of American Samoa. II. Accounts of flora and fauna. U.S. Fish and Wildlife Service, Washington, D.C. 151 pp. Amerson, A.B., Jr., W.A. Whistler, and T.D. Schwaner. 1982b. Wildlife and wildlife habitat of American Samoa. I. Environment and Ecology. U.S. Fish and Wildlife Service, Washington, D.C. 119 pp Anderson, S.J. and C.P. Stone. 1993. Snaring to control feral pigs Sus scrofa in a remote Hawaiian rain forest. Biological Conservation 63: 195–201. Aplet, G.H., S.J. Anderson, and C.P. Stone. 1991. Association between feral pig disturbance and the composition of some alien plant assemblages in Hawaii Volcanoes National Park. Vegetatio 95: 55–62. Arcilla, N. 2015, in litt. Email from former DMWR wildlife biologist, Nicole (Dauphine) Arcilla to FWS biologist J. Kwon providing information on presence of domestic cats on Ofu and Olesega Islands, 08 MAY 2015. Armstrong, D.P., Raeburn, E.P., Lewis, R.M., and D. Ravine. 2006. Modeling vital rates of a reintroduced New Zealand robin population as a function of predator control. Journal of Wildlife Management 70: 1,028-1,036. [ASCC] American Samoa Community College. 2010. American Samoa forest assessment and resource strategy 2011-2015. Prepared by Forestry Program, Division of Community and Natural Resources. 63 pp. [ASG-NPSA] American Samoa Government (ASG) and National Park of American Samoa (NPSA). 1993. Lease agreement between landowners of American Samoa and the American Samoa Government, and the Department of Interior. Atherton, J. and B. Jefferies. 2012. Rapid Biodiversity Assessment of Upland Savaii Samoa. Apia, Samoa. 176 pp. Atkinson, I.A.E. 1977. A reassessment of factors, particularly Rattus rattus L., that influenced the decline of endemic forest birds in the Hawaiian Islands. Pacific Science 31:109-133. Atkinson, C.T., and A.C. Medeiros. 2006. Trip report: Pilot study of factors linking watershed function and coastal ecosystem health in American Samoa, Pacific Island Ecosystems Research Center, Honolulu, HI. Open File Report 2006-1383. Badia, R. 2014, in litt. Crake project reports May – September 2013. Banks, R.C. 1984. Bird specimens from American Samoa. Pacific Science 38:150-169.

32

Bellingham, M. and A. Davis. 1988. Forest bird communities in western Samoa. Notornis 35: 117-128. BirdLife International. 2016. Gymnomyza samoensis. The IUCN Red List of Threatened Species 2016: e.T22704317A93962858. http://dx.doi.org/10.2305/IUCN.UK.2016- 3.RLTS.T22704317A93962858.en. Downloaded on 16 May 2019. Bonin, M. J. 2008. Mt. Vaea ecological restoration project, phase I. Prepared for Conservation International Pacific Islands Program and the Ministry of Natural Resources and Environment (MNRE). 73 pp. Bonin, M. J. 2010. Mt. Vaea ecological restoration project, phase II. Prepared for the MNRE and JICA Samoa. 32 pp. Butler, D. and R. Stirnemann. 2013. Leading the recovery of two of Samoa’s most threatened bird species the tooth-billed pigeon (Manumea) and the mao (Mao) through ecological research to identify current threats. Biodiversity Conservation Lessons Learned Technical Series 25. 50 pp. Campbell, T.A. and D.B. Long. 2009. Feral swine damage and damage management in forested ecosystems. Forest Ecology and Management 257:2319−2326. Caruso, C. 2015, in litt. Email from National Park of American Samoa Manua District ranger Carlo Caruso to FWS biologist J. Kwon providing information on rats on Manua Islands, 22 APR 2015. Chan-Halbrendt, C., T. Lin, F. Yang, and G. Sisior. 2010. Hawaiian residents’ preferences for Miconia control program attributes using conjoint choice experiment and latent class analysis. Environmental Management 45: 250–260. [CNMI-SWARS] Commonwealth of the Northern Mariana Islands (CNMI) and (SWARS) Council. 2010. Commonwealth of the Northern Mariana Islands (CNMI) statewide assessment and resource strategy 2010-2015+. 78 pp. Cook, R. P. 2001. Specificity of Liothrips urichi (Thysanoptera: Phlaeothripidae) for Clidemia hirta in American Samoa. Proc. Hawaii Entomol. Soc. 35: 143–144. Cowie, R.H. and R.P. Cook. 1999. The distribution and abundance of land snails in the National Park of American Samoa, with particular focus on Partulidae. Cooperative National Park Resources Studies Unit, Hawaii, Technical Report 123, i-iii, 1-143. Craig, P. 2009. Natural history guide to American Samoa. National Park Service. 131 pp. Cuddihy, L.W. 1984. Effects of cattle grazing on the mountain parkland ecosystem, Mauna Loa, Hawaii. Cooperative National Park Resources Studies Unit, University of Hawaii, Manoa, Honolulu. 135 pp. Cuddihy, L.W. and C.P. Stone. 1990. Alteration of native Hawaiian vegetation: effects of humans, their activities and introductions. Cooperative National Park Resources Studies Unit, University of Hawaii, Manoa, Honolulu. 138 pp. D’Antonio, C.M. and P.M. Vitousek. 1992. Biological invasions by exotic grasses, the grass/fire cycle, and global change. Annual Review of Ecology and Systematics 23: 63–87.

33

Diong, C.H. 1982. Population biology and management of the feral pig (Sus scrofa L.) in Kipahulu Valley, Maui. Thesis for doctorate of philosophy in the Department of Zoology, University of Hawaii at Manoa, Honolulu. 408 pp. [DMWR] Department of Marine and Wildlife Resources, American Samoa Government. 2006. A comprehensive strategy for wildlife conservation in American Samoa. September 2005, Revised November 2006. 116 pp. Dunkell, D.O., G.L. Bruland, C.I. Evensen, and C.M. Litton. 2011. Runoff, sediment transport, and effects of feral pigs (Sus scrofa) exclusion in a forested Hawaiian watershed. Pacific Science 65: 175–194. Editor. 2014. Recovery round-up. Corella 36(4):104 Engbring, J., and F.L. Ramsey. 1989. A 1986 survey of the forest birds of American Samoa. U.S. Fish and Wildlife Service, Department of Interior, Honolulu, Hawaii. [FAO] Food and Agricultural Organization. 2005, in litt. Forests and forestry in small island developing states. http://www.fao.org/forestry/sids/4142/en/, accessed 30 JUN 2015. Freifeld, H.B. 1999. Habitat relationship of forest birds on Tutuila Island, American Samoa. Journal of Biogeography. 26:1191-1213. Freifeld, H.B., C. Solek, and A. Tualaulelei. 2004. Temporal variation in forest bird survey data from Tutuila Island, American Samoa. Pacific Science 58:99-117. Gagne, W.C. and L.W. Cuddihy. 1999. Vegetation. In Manual of the Flowering Plants of Hawaii, Revised Edition, Vol. 1, W.L. Wagner, D.R. Herbst, and S.H. Sohmer (eds.), University of Hawaii Press and Bishop Museum Press, Honolulu. Pp. 45–114. Geering, D. 2002. Longevity of Aust birds. Accessed 30 May 2019 at http://bioacoustics.cse.unsw.edu.au/birding-aus/2002-04/msg00467.html Geesink, R., W.L. Wagner, and D.R. Herbst. 1999. 47. Fabaceae, pea family. In Manual of the Flowering Plants of Hawaii, Revised Edition, Vol. 1, W.L. Wagner, D.R. Herbst, and S.H. Sohmer (eds.), University of Hawaii Press and Bishop Museum Press, Honolulu. Pp. 629– 721. [GISD] Global Invasive Species Database. 2006. Adenanthera pavonina (plant), http://www.issg.org/database/species/ecology.asp?si=201&fr=1&sts=& ang=EN, accessed 27 APR 2015. [GISD] Global Invasive Species Database. 2008. Falcataria moluccana (plant), http://www.issg.org/database/species/ecology.asp?si=1249&fr=1&sts=& ang=EN, accessed 16 JUN 2015. [GISD] Global Invasive Species Database. 2010. Spathodea campanulata (plant), http://www.issg.org/database/species/ecology.asp?si=75&fr=1&sts=& ang=EN, accessed 27 APR 2015. Goldin, M.R. 2002. Field guide to the Samoan archipelago: fish, wildlife, and protected areas. Bess Press, Inc. 320 pp. Hess, S.C. and P.C. Banko. 2006, in litt. Feral cats: Too long a threat to Hawaiian wildlife. U.S. Geological Survey, Pacific Islands Ecosystems Research Center, FS 2006-3006.

34

Higgins, P.J., L. Christidis, and H.A. Ford. 2008. Family Meliphagidae (Honeyeaters). Pages 498-691 in del Hoyo, J., A. Elliott, and D.A. Christie editors. 2008. Handbook of the Birds of the World. Volume 13. Penduline-tits to Shrikes. Lynx Edicions, Barcelona. Hoshide, H. 1996. Preliminary assessment of feral pig problem American Samoa. Report for the National Park of American Samoa and AS Environmental Protection Agency. Final Report 29 MARCH 1996. 11 pp. Hughes, R.F., Uowolo, A.L., and T.P. Togia. 2012. Recovery of native forest after removal of an invasive tree. Biol. Invasions DOI 10.1007/s10530-011-0164-y. Published online: 22 JAN 2012 Jarvi, S.I., M.E.M. Farias, H. Baker, H.B. Freifeld, P.E. Baker, E. VanGelder, J.G. Massey, C. T. Atkinson. 2003. Detection of avian malaria (Plasmodium sp.) in native land birds of American Samoa. Conservation Genetics 4: 629-637. Kessler, C.C. 2011. Invasive species removal and ecosystem recovery in the Mariana Islands; challenges and outcomes on Sarigan and Anatahan. In Island Invasives: Eradication and Management, Veitch, C.R., M.N. Clout, and D.R. Towns (eds.), IUCN, Gland. Pp. 320−324. Knapp, K. R., M. C. Kruk, D. H. Levinson, H. J. Diamond, and C. J. Neumann, 2010: The International Best Track Archive for Climate Stewardship (IBTrACS): Unifying tropical cyclone best track data. Bulletin of the American Meteorological Society, 91, 363-376. doi:10.1175/2009BAMS2755.1 Lane, S.G., and R.G. Lonnon. 1967. Longevity of Fuscous Honeyeaters. The Australian Bird Bander. June:33. Leishman, A.J. 2012. Results from the long-term mist-netting project at North Ryde, commenced by the late S.G. (Bill) Lane in 1959. Corella 36(1):12-16. Levin, M. J., and P. A. Wright. 1974. “Report on the 1974 Census of American Samoa: Part II.” Honolulu, Hawaii: University of Hawaii at Manoa, East_West Population Institute. http:// [LINZ] Land Information New Zealand. 2015a, in litt. sw pacific built up area areas [Data File]. Accessed 31 May 2019 from https://data.linz.govt.nz/data/ [LINZ] Land Information New Zealand. 2015b, in litt. sw pacific crop land areas [Data File]. Accessed 31 May 2019 from https://data.linz.govt.nz/data/ [LINZ] Land Information New Zealand. 2015c, in litt. sw pacific grassland areas [Data File]. Accessed 31 May 2019 from https://data.linz.govt.nz/data/ [LINZ] Land Information New Zealand. 2015d, in litt. sw pacific island areas [Data File]. Accessed 31 May 2019 from https://data.linz.govt.nz/data/ [LINZ] Land Information New Zealand. 2015e, in litt. sw pacific mangrove swamp areas [Data File]. Accessed 31 May 2019 from https://data.linz.govt.nz/data/ [LINZ] Land Information New Zealand. 2015f, in litt. sw pacific marsh areas [Data File]. Accessed 31 May 2019 from https://data.linz.govt.nz/data/ [LINZ] Land Information New Zealand. 2015g, in litt. sw pacific swamp areas [Data File]. Accessed 31 May 2019 from https://data.linz.govt.nz/data/

35

[LINZ] Land Information New Zealand. 2015h, in litt. sw pacific thicket areas [Data File]. Accessed 31 May 2019 from https://data.linz.govt.nz/data/ [LINZ] Land Information New Zealand. 2015i, in litt. sw pacific wood areas [Data File]. Accessed 31 May 2019 from https://data.linz.govt.nz/data/ Loope, L.L., Medeiros, A.C. and B.H. Gagne. 1991. 77. Recovery of vegetation of a montane bog following protection from feral pig rooting. Technical report 76-78, Cooperative National Park Resources Studies Unit, University of Hawaii at Manoa, Honolulu. 23 pp. Loope, L.L., Hughes, R.F. and J-Y Meyer. 2013. Plant invasions in protected areas of tropical Pacific Islands, with special reference to Hawaii. Pp. 313- 348 in L.C. Foxcroft et al. (eds.), Plant Invasions in Protected Areas: Patterns, Problems and Challenges, Invading Nature - Springer Series in Invasion Ecology 7, DOI 10.1007/978-94-007-7750-7_15. Springer Science+Business Media Dordrecht. Lovegrove, T.; Bell, B.; Hay, R. 1992. The indigenous wildlife of Western Samoa: impacts of Cyclone Val and a recovery management strategy. New Zealand Department of Conservation, Wellington. 53 pp. MacDonald, M. 2015, in litt. Electronic message from DMWR wildlife biologist Mark MacDonald to FWS Biologist James Kwon regarding absence of mao on Tutuila, 13 JUL 2015. Matson, P. 1990. Plant-soil interactions in primary succession at Hawaii Volcanoes National Park. Oecologia 85: 241–246. Medina, F.M., Bonnaud, E., Vidal, E., Tershy, B.R., Zavaleta, E.S., Donlan, C.J., Keitt, B.S., Le Corre, M., Horwath, S.V. and M. Nogales. 2011. Global Change Biology doi: 10.1111/j.1365-2486.2011.02464.x. Meyer, R. J. Seamon, S. Faaumu, and I. Lalogafuafua. 2017. Classification and mapping of wildlife habitat in American Samoa: An object-based approach using high resolution orthoimagery and LIDAR remote sensing data. American Samoa Department of Marine and Wildlife Resources. 141 pp. [MNRE] Ministry of Natural Resources & Environment. 2006. Recovery Plan for the ma’oma’o or mao (Gymnomyza samoensis). Ministry of Natural Resources & Environment, Government of Samoa, Samoa. 46 pp. [MNRE] Ministry of Natural Resources & Environment. 2013. Samoa state of the environment report 2013. Ministry of Natural Resources & Environment, Government of Samoa, Samoa. 150 pp. [MNRE] Ministry of Natural Resources & Environment. 2014, in litt. Table 1: Samoa Land Cover by Island 2014. Accessed 6 June 2019 from http://www.sbs.gov.ws/index.php/new- document-library?view=download&fileId=1749 Mueller-Dombois, D. and R.F. Fosberg. 1998. Vegetation of the tropical Pacific Islands. Springer Verlag, New York. Nagle, B. 2010. Invasive plant project management, training workshop report. 27 pp.

36

[NOAA-IBTrACS] National Oceanic and Atmospheric Administration, International Best Track Archive for Climate Stewardship. 2019. IBTrACS version 4.Accessed 6 June 2019 from https://www.ncdc.noaa.gov/ibtracs/. Nogales, M., Vidal, E., Medina, F.M., Bonnaud, E., Tershy, B.R., Campbell, K.J. and E.S. Zavaleta. 2013. Feral cats and biodiversity conservation: the urgent prioritization of island management. BioScience 63: 804-810. Nogueira-Filho, S.L., S.S.C. Noguieira, and J.M.V. Fragoso. 2009. Ecological impacts of feral pigs in the Hawaiian Islands. Biodiversity and Conservation 18: 3677–3683. [NPSA] National Park of American Samoa. 2012, in litt. Invasive plant field guide. 32 pp. O’Donnell, C.F.J., Clapperton, B.K., and J. M. Monks. 2015. Impacts of introduced mammalian predators on indigenous birds of freshwater wetlands in New Zealand. New Zealand Journal of Ecology 39: 19-33. Paulson, D.D. 1994. Understanding tropical deforestation: the case of Western Samoa. Environmental Conservation 21: 326–332. [PCCSP] Pacific Science Program. 2019. Pacific Climate Change Data Portal, Samoa and American Samoa, http://www.bom.gov.au/climate/pccsp/. Accessed 31 May 2019. Pearsall, S.H. and W.A. Whistler. 1991. Terrestrial ecosystem mapping for Western Samoa: summary, project report, and proposed national parks and reserves plan. Prepared for the Government of Western Samoa by the South Pacific Regional Environment Program. 80 pp. [PIER] Pacific Islands Ecosystems at Risk. 2013. Spathodea campanulata. PIER species information, http://www.hear.org/pier/species/spathodea_campanulata.htm. Accessed 27 APR 2015. Pratt, H.D., and J.C. Mittermeier. 2016. Notes on the natural history, , and conservation of the endemic avifauna of the Samoan archipelago. Wilson Journal of Ornithology 128:217-241. Pratt, H. D., P. L. Bruner, and D. G. Berrett. 1987. A field guide to the birds of Hawaii and the tropical Pacific. Princeton University Press, Princeton, NJ. 409 pp. + plates. Public Law. 1988. An Act to Establish the National Park of American Samoa. Public Law. 2006. Boundary Adjustment of the National Park of American Samoa. 100-336. Reed, S. 1980. The birds of Savaii, Western Samoa. Notornis 27:151-159. Robertson, H.A., Rod Hay, J., Saul, E.K., and G.V. McCormack. 1994. Recovery of the kakekori: an endangered forest bird of the Cook Islands. Conservation Biology 8: 1,078- 1,086. Samoa Bureau of Statistics (SBS). 2011. Population and housing census 2011: Analytical Report. Government of Samoa. 252 pp. Scott, J.M., Mountainspring, S., Ramsey, F.L., and C.B. Kepler. 1986. Forest bird communities of the Hawaiian Islands: their dynamics, ecology, and conservation. Studies in Avian Biology 9: 1-429.

37

Seamon, J.O. 2004, in litt. Email message from Joshua Seamon, DMWR biologist, to Holly Freifeld, U.S. Fish and Wildlife Service, regarding candidate status updates for American Samoa, 31 MAY 2004. Siemann, E., J.A. Carrillo, C.A. Gabler, R. Zipp, and W.E. Rogers. 2009. Experimental test of the impacts of feral hogs on forest dynamics and processes in the southeastern US. Forest Ecology and Management 258: 546–553. Smith, C.W. 1985. Impact of alien plants on Hawaii’s native biota. In Hawaii’s Terrestrial Ecosystems: Preservation and Management, C.P. Stone and J.M. Scott (eds.), Cooperative National Park Resources Studies Unit, University of Hawaii, Honolulu. Pp. 180–250. Smith, C.W. 1989. Non-native plants. In Conservation Biology in Hawaii, Stone, C.P., D.B. Stone, LW. Cuddihy, and M.E. Land (eds.), University of Hawaii Press, Honolulu. Pp. 60– 69. Space, J.C. and T. Flynn. 2000. Observations on invasive plant species in American Samoa. 50 pp Space, J.C. and T. Flynn. 2002. Report to the Government of Samoa on invasive plant species of environmental concern. U.S.D.A. Forest Service, Pacific Southwest Research Station, Institute of Pacific Islands Forestry, Honolulu, Hawaii, USA. 80 pp. Stirnemann, R. 2015a. Ecology, population dynamics and drivers of decline in a tropical island honeyeater: the Ma’oma’o. PhD. Thesis. Massey University, New Zealand. 127 pp. Stirnemann, R. 2015b. in litt. Email from from Rebecca Stirnemann, researcher, providing information on mao population estimate, 1 APR 2015. Stirnemann, R., M.A. Potter, D. Butler, and E. Minot. 2015a. Acoustic differences enable sex discrimination in a species with high sexual and morphological overlap. Wilson Journal of Ornithology 127:375-386. Stirnemann, R., M.A. Potter, D. Butler, and E. Minot. 2015b. Slow life history traits in an endangered tropical island bird, the Ma’oma’o. Bird Conservation International 26:366-379. Stirnemann, R., M.A. Potter, D. Butler, and E. Minot. 2015c. Nest success does not predict reproductive success in a tropical island honeyeater. Pages 67-83 in R. Stirnemann. Ecology, population dynamics and drivers of decline in a tropical island honeyeater: the Ma’oma’o. PhD. Thesis. Massey University, New Zealand. 127 pp. Stone, C.P. 1985. Alien animals in Hawaii’s native ecosystems: toward controlling the adverse effects of introduced vertebrates. In Hawaii’s Terrestrial Ecosystems: Preservation and Management, Cooperative National Park Resources Studies Unit, University of Hawaii, Honolulu. Pp. 254–255, 261–264. Tauiliili, P. and A.M. Vargo. 1993. History of biological control in American Samoa. Micronesica 4(Suppl.): 57–60. Thornberry-Ehrlich, T. 2008. National Park of American Samoa Geologic Resource Evaluation Report. Natural Resource Report NPS/NRPC/GRD/NRR—2008/025. National Park Service, Denver, Colorado.

38

Togia, T. 2015, in litt. Email from Tavita Togia, NPSA terrestrial ecologist, to James Kwon, U.S. Fish and Wildlife Service, regarding nonnative plant and feral pig control, dated 24 APR 2015. Tomich, P.Q. 1986. Mammals in Hawaii: a synopsis and notational bibliography. Bishop Museum Press, Honolulu. Pp. 37–45, 120–134, 140–168. Tye, A. 2012, in litt. Restoration of Nuutele and Nuulua Islands (Aleipata Group), Samoa through the management of introduced rats and ants. CEPF final project completion report for Secretariat of the Pacific Regional Environment Programme (SPREP). 33 pp. Unkown. 1956. “Census of American Samoa: 1930-1956.” Census. http://pacificweb.org/categories/Statistical%20Activities/Census/AmericanSamoaCensus. html. U. S. Census Bureau. 1982. “Chapter A: Number of Inhabitants. Volume 1: Characteristics of the Population. 1980 Census of Population, Part 56: American Samoa.” Census. U.S. Census Bureau, Economics and Statistical Administration, U.S. Department of Commerce. http://pacificweb.org/categories/Statistical%20Activities/Census/AmericanSamoaCensus. html. U. S. Census Bureau. 1992. “Social Economic, and Housing Characteristics: American Samoa. 1990 Census of Population and Housing.” Census. U.S. Census Bureau, Economics and Statistical Administration, U.S. Department of Commerce. http://pacificweb.org/categories/Statistical%20Activities/Census/AmericanSamoaCensus. html. U.S. Census Bureau. 2010, in litt. Census 2000 and 2010 for American Samoa [Data File]. Retrieved from https://www.census.gov/2010census/news/xls/cb11cn177_as.xls [USDA] U.S. Department of Agriculture-Agricultural Research Service. National Genetic Resources Program. 2006. Germplasm Resources Information Network – (GRIN) [Online Database]. National Germplasm Resources Laboratory, Beltsville Maryland, http://www.ars-grin.gov/cgi-bin/npgs/html/taxon.pl?70913, accessed on 13 JUL 2015. [USFWS] U.S. Fish and Wildlife Service. 2016a. Endangered and threatened wildlife and plants; endangered status for five species from American Samoa; Final Rule. Federal Register 81:65466-65508. [USFWS] U.S. Fish and Wildlife Service. 2016b. USFWS Species Status Assessment Framework: an integrated analytical framework for conservation. Version 3.4. Dated August 2016. VanderWerf, E. 2009. Importance of nest predation by alien rodents and avian poxvirus in conservation of Oahu elepaio. Journal of Wildlife Management 73: 737-746. Van Riper, C., III, S.G. Van Riper, M.L. Goff, and M. Laird. 1986. The epizootiology and ecological significance of malaria in Hawaiian land birds. Ecological Monographs 56:327- 344.

39

Vitousek, P.M., C.M. D’Antonio, L.L. Loope, M. Rejmanek, and R. Westerbrooks. 1997. Introduced species: a significant component of human-caused global change. New Zealand Journal of Ecology 21: 1–16. Vitousek, P., G.P. Asner, O.A. Chadwick, and S. Hotchkiss. 2009. Landscape-level variation in forest structure and biogeochemistry across a substrate age gradient in Hawaii. Ecology 90: 2074–3086. Wagner, W. L., D. R. Herbst, and R. S. N. Yee. 1985. Status of the native flowering plants of the Hawaiian Islands. Pages 23-74 in C. P. Stone and J. M. Scott, editors. Hawai‘i’s terrestrial ecosystems: preservation and management. Univ. Hawai‘i Coop. Natl. Park Resources Stud. Unit. Univ. Hawai‘i Press, Honolulu, HI. Warner, R.E. 1968. The role of introduced diseases in the extinction of the endemic Hawaiian avifauna. Condor 70:101-120. Watling, D. 2001. A guide to the birds of Fiji and Western Polynesia including American Samo, Niue, Samoa, Tokelau, Tonga, Tuvalu and Wallis and Futuna. Environmental Consultants, Suva, Fiji. Whistler, W.A. 1992. Botanical inventory of the proposed Ta’u Unit of the National Park of American Samoa. Technical Report 83. Cooperative National Park Resources Studies Unit, University of Hawaii at Manoa. 85 pp. + figures. Whistler, W.A. 1994. Botanical inventory of the proposed Tutuila and Ofu Units of the National Park of American Samoa. Technical Report 87. Cooperative National Park Resources Studies Unit, University of Hawaii at Manoa. 142 pp. Whistler, W.A. 2002. The Samoan rainforest, vegetation of the Samoan archipelago. Isla Botanica. 168 pp. Wiley, J. W., and Wunderle, J. M. 1993. The effects of hurricanes on birds, with special reference to Caribbean islands. Bird Conservation International 3(04): 319-349. Wunderle Jr., J. M., and Wiley, J. W. 1996. Effects of hurricanes on wildlife: implications and strategies for management. In: Conservation of faunal diversity in forested landscapes (pp. 253-264). Springer Netherlands.

40