Endangered and Threatened Wildlife and Plants; Determination of Status for 12 Species of Picture-Wing Flies From the Hawaiian Islands, 26835-26852 [06-4299]

Agencies

• Fish and Wildlife Service
• DEPARTMENT OF THE INTERIOR

[Federal Register Volume 71, Number 89 (Tuesday, May 9, 2006)]
[Rules and Regulations]
[Pages 26835-26852]
From the Federal Register Online via the Government Printing Office [www.gpo.gov]
[FR Doc No: 06-4299]


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DEPARTMENT OF THE INTERIOR

Fish and Wildlife Service

50 CFR Part 17

RIN 1018 AG23


Endangered and Threatened Wildlife and Plants; Determination of 
Status for 12 Species of Picture-Wing Flies From the Hawaiian Islands

AGENCY: Fish and Wildlife Service, Interior.

ACTION: Final rule.

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SUMMARY: We, the U.S. Fish and Wildlife Service (Service), determine 
endangered status pursuant to the Endangered Species Act of 1973, as 
amended (Act), for 11 species of Hawaiian picture-wing flies--
Drosophila aglaia, D. differens, D. hemipeza, D. heteroneura, D. 
montgomeryi, D. musaphilia, D. neoclavisetae, D. obatai, D. ochrobasis, 
D. substenoptera, and D. tarphytrichia. We determine threatened status 
pursuant to the Act for one species of Hawaiian picture-wing fly--D. 
mulli. This final rule implements the Federal protections provided by 
the Act for these 12 species of Hawaiian picture-wing flies.

DATES: This final rule is effective June 8, 2006.

ADDRESSES: Comments and materials received, as well as supporting 
documentation used in the preparation of this final rule, will be 
available for public inspection, by appointment, during normal business 
hours at the Pacific Islands Fish and Wildlife Office, U.S. Fish and 
Wildlife Service, 300 Ala Moana Boulevard, Room 3-122, Box 50088, 
Honolulu, HI 96850.

FOR FURTHER INFORMATION CONTACT: Patrick Leonard, Field Supervisor, 
Pacific Islands Fish and Wildlife Office (see ADDRESSES section) 
(telephone 808/792-9400; facsimile 808/792-9581). Persons who use a 
telecommunications device for the deaf (TDD) may call the Federal 
Information Relay Service (FIRS) at 800/877-8339, 24 hours a day, 7 
days a week.

SUPPLEMENTARY INFORMATION:

Background

    Many of the major ecological zones of the earth are represented in 
Hawaii, from coral reef systems through rain forests to high alpine 
deserts, in less than 10,800 square kilometers (6,500 square miles) of 
land. The range of topographies creates a great diversity of climates. 
Windward (northeastern) slopes can receive up to 1,000 cm (400 in) of 
rain per year, while some leeward coasts that lie in the rain shadow of 
the high volcanoes are classified as deserts, receiving as little as 25 
cm (10 in) of rain annually. This topographic and climatic regime has 
given rise to a rich diversity of plant communities, including coastal, 
lowland, montane, subalpine, and alpine; dry, mesic, and wet; and 
herblands, grasslands, shrublands, forests, and mixed communities 
(Gagne and Cuddihy 1990). These habitats and plant communities in turn 
support one of the most unique arthropod faunas in the world, with an 
estimated 10,000 endemic species (Howarth 1990). Unusual 
characteristics of Hawaii's native arthropod fauna include the presence 
of relict species; the absence of social insects, such as ants and 
termites; endemic genera; extremely small geographic ranges; adaptation 
of species to very specific conditions or environments; novel 
ecological shifts; flightlessness; and loss of certain antipredator 
behaviors (Zimmerman 1948, 1970; Simon et al. 1984; Howarth 1990). 
Native vegetation on all the main Hawaiian Islands has undergone 
extreme alteration because of past and present land management 
practices, including ranching, introduction of nonnative plants and 
animals, and agricultural development (Cuddihy and Stone 1990).
    Each species of Hawaiian picture-wing fly described in this 
document is

[[Page 26836]]

found only on a single island, and the larvae of each are dependant 
upon only a single or a few related species of plants (see Table 1). 
These host plant species are threatened by a variety of factors, 
including their direct destruction by pigs, goats, cattle, rats, and 
competition with nonnative plants, and the indirect effects of soil 
disturbance which further promotes the spread of nonnative species (see 
Factors A and C below). In addition to the habitat alteration, the 
picture-wing flies included in this rule are threatened by a variety of 
introduced predatory species including yellow jackets and several ant 
species. This suite of threats to the picture-wing flies and its 
habitat are discussed in more detail in the Summary of Factors 
Affecting the Species section.
    Flies in the Drosophilidae family in Hawaii represent one of the 
most remarkable cases of specific adaptation to local conditions that 
has been found in any group of animals (Hardy and Kaneshiro 1981). 
These insects are distributed throughout the eight main Hawaiian 
Islands (i.e., Hawaii, Maui, Oahu, Kauai, Molokai, Lanai, Niihau, and 
Kahoolawe), and each species is typically found on a single island 
(Carson and Yoon 1982).
    The general life cycle of Hawaiian Drosophilidae is typical of that 
of most flies: After mating, females lay eggs from which larvae 
(immature stage) hatch; as larvae grow they molt (shed their skin) 
through three successive stages (instars); when fully grown, the larvae 
change into pupae (a transitional form) in which they metamorphose and 
emerge as adults.
    Breeding generally occurs year-round, but egg laying and larval 
development increase following the rainy season as the availability of 
decaying matter, which the flies feed on, increases in response to the 
heavy rains (K. Kaneshiro, in litt., 2005b). In general, Drosophila lay 
between 50 and 200 eggs in a single clutch. Eggs develop into adults in 
about a month, and adults generally become sexually mature one month 
later. Adults generally live for one to two months.
    As a group, Hawaiian Drosophilidae can be found in most of the 
natural communities in Hawaii. They have developed and adapted 
ecologically to a tremendous diversity of ecosystems ranging from 
desert-like habitats, to rain forests, to swampland (Kaneshiro and 
Kaneshiro 1995). While the larval stages of most species are 
saprophytic (feeding on decaying vegetation, such as rotting leaves, 
bark, flowers, and fruits), some have become highly specialized, being 
carnivorous on egg masses of spiders, or feeding on green algae growing 
underwater on boulders in streams (Kaneshiro and Kaneshiro 1995).
    Hawaiian Drosophila, and in particular picture-wing Drosophila, are 
unique among living organisms because adaptive radiation (the evolution 
of an ancestral species, which was adapted to a particular way of life, 
into many diverse species, each adapted to a different habitat) has 
resulted in unparalleled biological diversity within a single large, 
closely related group of species (Foote and Carson 1995). The banding 
patterns of all five major chromosome arms among 106 species of 
Hawaiian picture-winged Drosophila revealed a 5 million-year-old 
evolutionary history rooted to species on the island of Kauai (Carson 
1992). This work on the evolutionary history of Hawaiian Drosophila 
augments an extensive systematic treatment of the genus (Hardy 1965; 
Kaneshiro 1976).
    Unlike numerous Hawaiian insects known only from their original 
taxonomic descriptions, many aspects of Hawaiian Drosophilidae biology 
have been researched, including their internal and external morphology, 
behavior, ecology, physiology, biochemistry, the banding sequence of 
giant chromosomes, and the structure of their DNA (Kaneshiro and 
Kaneshiro 1995). More than 80 research scientists and over 350 
undergraduates, graduate students, and postdoctoral fellows have 
participated in research on many species of the Hawaiian Drosophilidae, 
resulting in over 600 scientific publications.
    Because a large number of sites across the Hawaiian Islands have 
been surveyed since the 1960s using bait stations that are not species-
specific, researchers have a relatively good understanding of the 
distribution of Drosophila species and how that distribution has 
changed over time. Biologists have observed a general decline of the 
Hawaiian Drosophilidae along with other components of the native 
ecosystem. As noted by Spieth (1980), during the early part of the 
century, the Tantalus area (northeast of Honolulu) was a major spot for 
collecting Drosophila species. Since 1971, routine sampling in the 
Tantalus area has documented dramatic declines in the abundance of some 
Drosophila species and in other cases local extirpations (Foote and 
Carson 1995).
    All 12 species described below belong to the species group commonly 
known as the picture-wing Drosophila. This group consists of 106 known 
species, most of which are relatively large with elaborate markings on 
the otherwise clear wings of both sexes, the pattern of which varies 
among species (Hardy and Kaneshiro 1981; Carson 1992). The picture-wing 
Drosophila have been referred to as the ``birds of paradise'' of the 
insect world because of their relatively large size, colorful wing 
patterns, and the males' elaborate courtship displays and territorial 
defense behaviors.
    Males occupy territories that serve as mating arenas, or leks, to 
which receptive females are attracted. The male Drosophila use 
different techniques to ward off competing suitors. One species, 
Drosophila heteroneura, butts heads like bighorn sheep. Others grasp 
one another with legs and wings in a wrestling match. Yet another tries 
to intimidate with noise, creating a buzzing roar with muscles from its 
abdomen. When the male has secured his position in the lek, he performs 
a detailed choreography of behaviors for the females visiting that 
site. If he does not convey the right moves and messages, she leaves 
without mating. Each species has its own ritual; some include dancing 
around the female, buzzing of wings at a specific pitch, placing the 
male's head under the female's wing, tongue-tasting, or dousing the 
female with pheromone.
    The primary dataset we used to document observations of these 
picture-wing flies spans the years 1965 to 1999 (K. Kaneshiro, in 
litt., 2005a). Additional data were obtained from individuals familiar 
with particular species and locations. Many sites were surveyed 
infrequently or have not been surveyed in a long time while others have 
relatively complete records from 1966 to 1999. In this rule, when we 
state the date a species was last observed in a particular year, we do 
not intend to imply that comprehensive surveys have been conducted in 
subsequent years, only that the specified year was the last year that 
the species was located.

[[Page 26837]]



   Table 1.--Distribution of 12 Hawaiian Picture-Wing Flies by Island, General Habitat Type, and Primary Host
                                                    Plant(s)
----------------------------------------------------------------------------------------------------------------
              Species                        Island           General habitat type      Primary host plant(s)
----------------------------------------------------------------------------------------------------------------
Drosophila aglaia..................  Oahu..................  Mesic forest..........  Urera glabra
D. differens.......................  Molokai...............  Wet forest............  Clermontia sp.
D. hemipeza........................  Oahu..................  Mesic forest..........  Cyanea sp., Lobelia sp.,
                                                                                      and Urera kaalae
D. heteroneura.....................  Hawaii................  Mesic to wet forest...  Cheirodendron sp.,
                                                                                      Clermontia sp., Delissea
                                                                                      sp.
D. montgomeryi.....................  Oahu..................  Mesic forest..........  Urera kaalae
D. mulli...........................  Hawaii................  Wet forest............  Pritchardia beccariana
D. musaphilia......................  Kauai.................  Mesic forest..........  Acacia koa
D. neoclavisetae...................  Maui..................  Wet forest............  Cyanea sp.
D. obatai..........................  Oahu..................  Dry to mesic forest...  Pleomele aurea and Pleomele
                                                                                      forbesii
D. ochrobasis......................  Hawaii................  Mesic to wet forest...  Clermontia sp., Marattia
                                                                                      sp., and Myrsine sp.
D. substenoptera...................  Oahu..................  Wet forest............  Cheirodendron sp. and
                                                                                      Tetraplasandra sp.
D. tarphytrichia...................  Oahu..................  Mesic forest..........  Charpentiera sp.
----------------------------------------------------------------------------------------------------------------

Discussion of the Species

Drosophila aglaia

    Drosophila aglaia was first recorded in 1946, on Mount Kaala on the 
island of Oahu, and described by Hardy (1965). D. aglaia is a small 
species, 0.15 inches (in) (4.0 millimeters (mm)) in length, with wings 
0.2 in (5.0 mm) long. It has a yellow head that is approximately one-
third wider than long. The eyes are brown, and the antennae are yellow, 
tinged with brown. The thorax is clear yellow with three broad brown 
stripes on the top, and the legs are yellow. The abdomen is brown with 
a large yellow spot on each of the hind corners. The wings are 
predominantly clear with irregular but characteristic brown markings, 
and are about two and three-quarter times longer than wide.
    Drosophila aglaia is historically known from five localities in the 
Waianae Mountains of Oahu between 1,400 and 2,800 feet (ft) (427 to 853 
meters (m)) above sea level. During 50 survey dates between 1966 and 
1990, 28 individuals were observed (Kaneshiro in litt., 2005a). The 5 
sites include: One lowland mesic Diospyros sp. and Metrosideros sp. 
(ohia) forest site in Makaleha Valley; two lowland mesic Acacia koa 
(koa) and ohia forest sites at Peacock Flats (Kapuahikahi Gulch) and 
Palikea; one site in diverse mesic forest at Puu Kaua; and a lowland, 
dry to mesic forest site at Puu Pane (K. Kaneshiro, in litt., 2005a).
    The last observation of this species occurred in 1997 during the 
last survey of the Palikea site. The species has not been observed at 
the other four historic sites since 1970 or 1971 despite subsequent 
surveys. However, two of the sites (Kapuahikahi Gulch and Makaleha 
Valley) have not been surveyed since the 1970s and one site, Puu Pane, 
was surveyed only once again in 1991 (K. Kaneshiro, in litt., 2005a).
    Drosophila aglaia is restricted to the natural distribution of its 
host plant, Urera glabra (family Urticaceae), which is a small shrub-
like endemic tree. The larvae of D. aglaia develop in the decomposing 
bark and stem of  U. glabra. This plant does not form large stands, but 
is infrequently scattered throughout slopes and valley bottoms in mesic 
and wet forest habitat on Oahu. In the Waianae Mountains on the west 
side of Oahu, this tree occurs infrequently in mesic forest.

Drosophila differens

    Drosophila differens was described by Hardy and Kaneshiro (1975) 
from specimens first recorded at South Hanalilolilo, Molokai, in 1972. 
This species is larger than most picture-wings, approximately 0.3 in 
(7.0 mm) in length, with wings 0.3 in (8.3 mm) long. D. differens has 
an entirely or predominantly yellow face and characteristic markings 
extending to the tip of the wings.
    Drosophila differens is historically known from three sites on 
private land between 3,800 and 4,500 ft (1,158 to 1,372 m) above sea 
level, within montane wet ohia forest (HBMP, in litt., 2005; K. 
Kaneshiro, in litt., 2005a). During 40 surveys between 1965 and 1999, 
63 individuals were recorded. At Hanalilolilo, the species was observed 
on eight survey dates between 1967 and 1983, but was not observed on 
three subsequent survey dates, the most recent being 1999. At a second 
site, Kaunuohua, which was only surveyed twice, individuals were 
observed in 1969 but not in 1999. At the third site, Puu Kolekole, 
individuals were documented in 1969 and again in 1999 (K. Kaneshiro, in 
litt., 2005a). An estimated 75 to 90 percent of D. differens' total 
potential habitat has been surveyed (K. Kaneshiro, pers. comm. 2006).
    Montgomery (1975) found that Drosophila differens larvae inhabit 
the bark and stems of Clermontia sp. (family Campanulaceae) in wet 
rainforest habitat (Kaneshiro and Kaneshiro 1995). Approximately 10 to 
25 percent of D. differens' potential habitat on steep, difficult to 
access areas and on State Natural Reserve lands surrounding its known 
range remains unsurveyed for the species (Science Panel 2005; K. 
Kaneshiro, pers. comm. 2006).

Drosophila hemipeza

    Hardy (1965) described Drosophila hemipeza from specimens recorded 
at Pupukea, Oahu, in 1952. The thorax of D. hemipeza is predominantly 
yellow with two brown stripes on the top, and the legs are entirely 
yellow. This species is 0.2 in (5.0 mm) long; the front legs are very 
slender with short straight bristles; and the wings are 0.2 in (6.0 mm) 
in length, slender, and somewhat pointed.
    Drosophila hemipeza is restricted to the island of Oahu where it is 
historically known from seven localities between 1,600 and 2,800 ft 
(488 to 853 m) above sea-level (not including the Pupakea site of 
discovery which is considered an extripated population). Since formal 
surveys began for the species, 49 individuals were recorded during a 
total of 56 different survey dates between 1965 and 1999 (K. Kaneshiro, 
in litt., 2005a). The species has been documented from seven sites, 
with survey history at these sites as follows: (1) The species was 
documented in 1969 but not in subsequent surveys spanning until 1972 in 
the Makaleha Valley; (2) individuals were detected at Puu Kaua in 1971 
but not in subsequent surveys as recently as 1999; (3) at Kaluaa Gulch, 
the species was observed in 1971 but not in 1972; (4) in Makaha Valley, 
the species was detected in 1971 and no surveys have been conducted 
since; (5) at Palikea the last observation occurred in 1997, also the 
date of the last survey; and (6) the species has not been detected at 
the Mauna Kapu site since 1975 despite

[[Page 26838]]

subsequent surveys spanning until 1983; (7) the species was detected at 
Pauoa Flats in the Koolau Range that was surveyed three times between 
1973 and 1974, with one observation of one individual during the last 
survey in 1974 (K. Kaneshiro, in litt., 2005a).
    Montgomery (1975) determined that Drosophila hemipeza larvae feed 
within decomposing portions of several different mesic forest plants. 
The larvae inhabit the decomposing bark of Urera kaalae (family 
Urticaceae), a federally-endangered plant (USFWS 1991, 1995) that grows 
on slopes and in gulches of diverse mesic forest. In 2004, only 41 
individuals of U. kaalae were known to remain in the wild (USFWS, in 
litt., 2004). The larvae also feed within the decomposing stems of 
Lobelia sp. (family Campanulaceae) and the decomposing bark and stems 
of Cyanea sp. (family Campanulaceae) in mesic forest habitat (Kaneshiro 
and Kaneshiro 1995; Science Panel 2005).

Drosophila heteroneura

    R.C.L. Perkins initially described this species as Idiomyia 
heteroneura, based on specimens from Olaa on the island of Hawaii 
(Perkins 1910). This taxon was later transferred to the genus 
Drosophila (Hardy 1969), forming its presently accepted name. 
Drosophila heteroneura has very large spots on the bases of the wings 
and the males have a broad head with the eyes situated laterally, 
giving them a hammerhead appearance. The hammer-shaped head and 
entirely yellow face differentiate it from D. silvestris, a closely 
related species. The thorax is predominantly yellow with several black 
streaks and markings on top. The legs are yellow except for slight 
tinges of brown on the ends of the middle and hind femora and tibiae. 
The wings are hyaline (transparent) and are very similar in markings 
and venation (vein markings) to those of D. silvestris, except that the 
marking in the front margin of the wing of D. heteroneura extends 
nearly to the marking at the end of the wing. The abdomen is shiny 
black with a large yellow spot on the top of each segment. This species 
is about 0.22 in (5.7 mm) in length with wings approximately 0.3 in 
(7.0 mm) long (Kaneshiro and Kaneshiro 1995).
    Drosophila heteroneura has been the most intensely studied of the 
12 species discussed in this rule (Kaneshiro and Kaneshiro 1995). This 
species is restricted to the island of Hawaii where, historically, it 
was known to be relatively widely distributed between 3,800 and 5,500 
ft (1,158 to 1,675 m) above sea level. D. heteroneura has been recorded 
from 24 localities on 4 of the island's 5 volcanoes (Hualalai, Mauna 
Kea, Mauna Loa, and Kilauea) in 5 different montane environments 
(Kaneshiro and Kaneshiro 1995; HBMP, in litt., 2005; K. Kaneshiro, in 
litt., 2005a).
    Based on the relatively extensive survey data, the population 
decline of Drosophila heteroneura has been demonstrated clearly. For 
example, D. heteroneura was recorded 760 times during surveys between 
1975 and 1979. In the early 1980s, the first disappearance of a D. 
heteroneura population was recorded from the Olaa Forest site in Hawaii 
Volcanoes National Park (Carson 1986; Foote and Carson 1995). 
Subsequently, the absence of the species was noted in several other 
locations in southern and western parts of the island where D. 
heteroneura had previously been relatively common. By the late 1980s, 
D. heteroneura was believed to be extinct until an extremely small 
population was discovered on private land at Hualalai Volcano in 1993 
(Kaneshiro and Kaneshiro 1995). The species was not observed again 
until 1998 when Foote (2000) recorded six specimens of D. heteroneura 
inhabiting a site at approximately 4,436 ft (1,352 m) above sea level 
near a host plant species, Clermontia clermontioides. D. heteroneura 
was last observed in 2001, at the refuge (D. Foote, pers. comm., 2005).
    Drosophila heteroneura larvae primarily inhabit the decomposing 
bark and stems of Clermontia sp. (family Campanulaceae), including C. 
clermontioides, and Delissea sp. (family Campanulaceae), but it is also 
known to feed within decomposing portions of Cheirodendron sp. (family 
Araliaceae) in open mesic and wet forest habitat (Kaneshiro and 
Kaneshiro 1995).

Drosophila montgomeryi

    Drosophila montgomeryi was described by Hardy and Kaneshiro (1971) 
from specimens collected in the Waianae Mountains of Oahu in 1970. 
Morphologically, this species appears to be most closely related to D. 
pisonia from the island of Hawaii. It can be distinguished by the 
narrow, pale brown stripe on each side of the top of the thorax, the 
long hairs on the front legs, and the second antennal segment, which is 
yellow, tinged with brown on the top.
    Drosophila montgomeryi is historically known from three localities 
in the Waianae Mountains on western Oahu between 2,000 and 2,800 ft 
(610 to 853 m) above sea level. The best available information 
concerning the status of the species at these sites is as follows: (1) 
One individual was recorded from Kaluaa Gulch during the last survey in 
1972; (2) at Palikea, one individual was observed on the last survey 
date in March 1997; and (3) at Puu Kaua, historically the site with the 
highest number of total individuals observed, the species was last 
detected in 1971 despite five subsequent surveys between 1997 and 1999 
(K. Kaneshiro, in litt., 2005a).
    Montgomery (1975) reported that the larvae of this species feed 
within the decaying bark of Urera kaalae, a federally-endangered plant 
(USFWS 1991, 1995) that grows on slopes and in gulches of diverse mesic 
forest (Kaneshiro and Kaneshiro 1995). In 2004, only 41 individuals of 
U. kaalae were known to remain in the wild (USFWS, in litt., 2004).

Drosophila mulli

    Drosophila mulli was described by Perreira and Kaneshiro (1990) and 
named for William P. Mull, the Hawaiian naturalist who first discovered 
this species. The head of D. mulli is yellow on the front and covered 
with light, silvery grey fuzz. The face of the male is 
characteristically white, while that of the female is brown. The top of 
the thorax is brownish yellow and lacks conspicuous markings or 
stripes. The legs are predominantly yellow, and the front legs of males 
bear three distinct rows of long, curled hairs. The wings are two and 
one-half times longer than wide, with distinct brown markings at the 
base and the tip. The length of the body is 0.17 to 0.2 in (4.3 to 5.0 
mm), and the wings are 0.17 to 0.19 in (4.3 to 4.8 mm) long (Kaneshiro 
and Kaneshiro 1995).
    Drosophila mulli is restricted to the island of Hawaii and is 
historically known from two locations between 3,200 and 4,000 ft (985 
to 1,220 m) above sea level. Adult flies are found only on the leaf 
undersides of the endemic fan palm, Pritchardia beccariana (family 
Arecaceae) which is the only known association of a Drosophila species 
with a native Hawaiian palm species. Individual P. beccariana are long-
lived (approximately 100 years). Current regeneration of the host plant 
has been compromised by feral ungulates, rats, and scolytid beetles 
(see Summary of Factors Affecting the Species section below). The 
larval feeding site on the plant remains unknown because attempts to 
rear this species from decaying parts of P. beccariana have thus far 
been unsuccessful (W. P. Mull, Volcano, Hawaii, pers. comm., 1994; 
Science Panel 2005).
    The site of the discovery for Drosophila mulli is located within a

[[Page 26839]]

State-owned montane wet ohia forest at Olaa Forest Reserve at 
approximately 3,200 ft (985 m) above sea level. This site was surveyed 
at least 62 times between 1965 and 2001, with fewer than 10 individuals 
observed on 4 different dates. The last recorded observation at this 
site occurred in 2001 (K. Kaneshiro, in litt., 2005a; D. Foote, in 
litt., 2006). A second locality was discovered in 1999, approximately 
9.3 mi (15 km) from the original site within a State-owned montane wet 
ohia forest site at Upper Waiakea Reserve at approximately 4,000 ft 
(1,219 m) above sea level (Science Panel 2005; S. Montgomery, pers. 
comm., 2005a).

Drosophila musaphilia

    Hardy (1965) formally described Drosophila musaphilia from 
specimens collected at Kokee, Kauai, in 1952. Although Hardy (1965) 
originally indicated that D. musaphilia is very similar to D. 
villosipedis, more recent work indicates D. musaphilia is most closely 
related to D. hawaiiensis (Kaneshiro et al. 1995).
    Drosophila musaphilia is characterized by a predominantly black 
thorax with gray fuzz and a very narrow gray stripe extending down the 
top. The legs are dark brown to yellow, with the front tibia devoid of 
ornamentation, and the tips of the legs have abundant long, black hairs 
on top. The wings are three times longer than wide with characteristic 
markings of the D. hawaiiensis group. The abdomen is dark brown to 
black and densely covered with brown fuzz. The body length is about 0.2 
in (5.0 mm) and the wings 0.207 in (5.25 mm) long (Kaneshiro and 
Kaneshiro 1995).
    Drosophila musaphilia is historically known from only four sites, 
one at 1,900 ft (579 m) above sea level, and three sites between 3,000 
and 3,500 ft (915 to 1,065 m) above sea level. The species has been 
observed a total of 11 times during 52 different survey dates since its 
discovery (Kaneshiro and Kaneshiro 1995; K. Kaneshiro, in litt., 
2005a). Researchers estimate that 75 percent of D. musaphilia's total 
potential habitat has been surveyed (K. Kaneshiro, pers. comm. 2006). 
The best available information concerning the status of the species at 
these sites is as follows: (1) A single observation of D. musaphilia 
was recorded from one lowland, wet ohia forest site at Wahiawa 
(Alexander Reservior) in 1968 (this population is believed to be 
extirpated); (2) at the Halemanu site, the species was observed in 1970 
and last observed in 1972 but not in subsequent surveys as recent as 
1996; (3) one individual was observed in 1968 at the Kokee (Nualolo 
Trail) site and not again during numerous surveys through 1999; and (4) 
individuals were last observed in 1988 at the Pihea Trail site located 
at 3,000 ft (915 m), but was not relocated in five subsequent surveys 
between 1989 and 1999 in that area (HBMP, in litt., 2005; Kaneshiro, in 
litt., 2005a).
    Montgomery (1975) determined that the host plant for Drosophila 
musaphilia is Acacia koa. The females lay their eggs upon, and the 
larvae develop in, the moldy slime flux (seep) that occasionally 
appears on certain trees with injured plant tissue and seeping sap. 
Understanding the full range of D. musaphilia is difficult because its 
host plant, Acacia koa, is fairly common and stable within, and 
surrounding, its known range on Kauai; however, the frequency of 
suitable slime fluxes occurring on the host plant appears to be much 
more restricted and unpredictable (Science Panel 2005).

Drosophila neoclavisetae

    Drosophila neoclavisetae was described by William Perreira and 
Kenneth Kaneshiro (1990) from specimens collected at Puu Kukui, West 
Maui, in 1969. It was named for its obvious affinities with D. 
clavisetae from East Maui. Both species are similar in wing and thorax 
markings, and they share a specialized part of the courtship behavior. 
The male bends its abdomen up over its head, produces a bubble of 
liquid (believed to be a sex pheromone) from its anal gland and then 
vibrates the abdomen, fanning the scent toward the female. Both D. 
neoclavisetae and D. clavisetae are members of the D. adiastola species 
group (Perreira and Kaneshiro 1990), and while other species in this 
group perform similarly unusual mating dances, the behavior is highly 
exaggerated in D. clavisetae and D. neoclavisetae (Kaneshiro and 
Kaneshiro 1995).
    Drosophila neoclavisetae is between 0.2 and 0.25 in (6.0 and 6.4 
mm) in length, with wings 0.26 to 0.3 in (6.5 to 7.0 mm) long. It is 
distinguished by its amber brown head and yellow face, with the middle 
portion raised to form a prominent ridge. The thorax is predominantly 
reddish brown with a distinct brown median stripe, bordered on each 
side by two brown stripes. The legs are yellow, with brown on the 
femora and a distinct brown band on the tips of the tibiae. The wings 
are broad and rounded, more than twice as long as wide, and with the 
front portion covered with brown markings and large clear spots tinged 
light yellow. It shares with D. clavisetae an extra cross-vein in the 
wing, which distinguishes both these species from the other species of 
the D. adiastola group. The abdomen is dark brown and black with 
numerous long hairs on the hind segments of the male (Kaneshiro and 
Kaneshiro 1995).
    Two populations of Drosophila neoclavisetae were found historically 
along the Puu Kukui Trail within montane wet ohia forests on State land 
in West Maui. One habitat site was found in 1969 at 4,440 ft (1,353 m) 
and the other in 1975 at 3,500 ft (1,067 m) above sea level (Kaneshiro 
and Kaneshiro 1995; HBMP, in litt., 2005; K. Kaneshiro, in litt., 
2005a). Fewer than 10 individuals have been observed despite attempts 
to relocate the species through 1997 (Kaneshiro and Kaneshiro 1995; K. 
Kaneshiro, in litt., 2005a; K. Kaneshiro pers. comm. 2006). Researchers 
estimate that between 90 and 95 percent of D. neoclavisetae's total 
potential range has been surveyed (K. Kaneshiro, pers. comm., 2006).
    The host plant of Drosophila neoclavisetae has not yet been 
confirmed, although it is likely associated with Cyanea sp. (family 
Campanulaceae). Because both collections of this species occurred 
within a small patch of Cyanea sp. and because many other species in 
the D. adiastola species group use species in this genus and other 
plants in the family Campanulaceae, researchers believe the Cyanea sp. 
found at Puu Kukui is likely the correct host plant for D. 
neoclavisetae (Science Panel 2005; Kaneshiro and Kaneshiro 1995). Due 
to its inaccessibility, some potential habitat surrounding the known 
range of D. neoclavisetae remains unsurveyed for the species (Science 
Panel 2005).

Drosophila obatai

    Drosophila obatai was described by Hardy and Kaneshiro in 1972, 
from specimens collected in the Waianae Mountains of Oahu. D. obatai 
resembles D. sodomae from Maui and Molokai and is distinguished by 
small differences in wing markings and the black coloration of the 
abdomen.
    Drosophila obatai is historically known from two localities between 
1,500 and 2,200 ft (457 to 670 m) above sea level. Nine individuals 
were recorded during ten surveys between 1970 and 1991 (Kaneshiro, in 
litt., 2005a). Individuals of the species were detected in November 
1971 at the time of the last survey at Wailupe Gulch. The second site 
(Puu Pane), has been surveyed eight times between 1970 and 1991, with 
the last detection occurring in March 1971 (Kaneshiro, in litt., 
2005a).
    Drosophila obatai larvae feed within decomposing portions of 
Pleomele forbesii, a candidate for Federal listing

[[Page 26840]]

(90 FR 24870), and Pleomele aurea (both in the family Agavaceae) 
(Kaneshiro and Kaneshiro 1995; Montgomery 1975). These host plants grow 
on slopes in dry forest and diverse mesic forest, and occur singly or 
in small clusters, rarely forming large stands (Wagner et al. 1999).

Drosophila ochrobasis

    Drosophila ochrobasis was originally described by Hardy and 
Kaneshiro (1968) based on a specimen collected from Puu Hualalai on the 
island of Hawaii at an elevation of 5,550 ft (1,692 m) above sea level. 
Based on chromosomal studies, D. ochrobasis is a member of the D. 
adiastola group and appears to be most closely related to D. 
setosimentum (Kaneshiro et al. 1995).
    Both the body and wings of Drosophila ochrobasis are approximately 
0.18 in (4.6 mm) in length. The head is yellow in front and brown on 
top, and the face is white with a prominent ridge running down the 
middle. The thorax is yellow except for a large brown spot on each 
side. The legs are yellow tinged with brown. In males, the basal three-
fifths of the wings are predominantly clear to translucent with faint 
transverse streaks of brown. The outer two-thirds of the wing is dark 
brown with large clear spots similar to that portion of the wings in D. 
setosimentum. The females of D. ochrobasis are virtually 
indistinguishable from D. setosimentum females (Kaneshiro and Kaneshiro 
1995).
    Historically, Drosophila ochrobasis was relatively widely 
distributed between 3,900 and 5,300 ft (1,189 to 1,615 m) above sea 
level. D. ochrobasis has been recorded from 10 localities on 4 of the 
island's 5 volcanoes (Hualalai, Mauna Kea, Mauna Loa, and the Kohala 
mountains).
    Recorded almost every year from 1967 to 1975, sometimes in 
relatively large numbers (135 occurrences in the period between 1970 
and 1974), Drosophila ochrobasis is now largely absent from its 
historical localities. A single individual of D. ochrobasis was last 
observed at the 1855 lava flow (Kipuka 9 and Kipuka 14) in 1986 
(Kaneshiro and Kaneshiro 1995; K. Kaneshiro, in litt., 2005a). Several 
surveys between 1995 to 1997 failed to locate the species at many of 
its historical sites (K. Kaneshiro, in litt., 2005a).
    The larvae of this species have been reported to use the 
decomposing portions of three different host plant groups--Myrsine sp. 
(family Myrsinaceae), Clermontia sp. (family Campanulaceae), and 
Marattia sp. (family Marattiaceae) (Montgomery 1975; Kaneshiro and 
Kaneshiro 1995).

Drosophila substenoptera

    Hardy (1965) originally described this species as Idiomyia 
substenoptera. He later determined the genus Idiomyia to be synonymous 
with Drosophila (Hardy 1969), thus creating the current name of 
Drosophila substenoptera. This species is closely related to D. 
planitibia and its relatives (Kaneshiro et al. 1995), but is 
distinguished by its wing markings, narrow wing shape, and complexity 
of the male genitalia. D. substenoptera is predominantly yellow with 
two black stripes extending down the entire length of the top surface 
of the thorax. The legs are yellow and lack long hairs on the dorsal 
surfaces. Body length is 0.171 in (4.35 mm), and the wings are 0.2 to 
0.21 in (5.0 to 5.3 mm) long (Kaneshiro and Kaneshiro 1995).
    Drosophila substenoptera is historically known from seven 
localities in both the Koolau and Waianae Mountains at elevations 
between 1,300 and 3,900 ft (396 to 1,189 m) above sea level. Drosophila 
substenoptera is now only known to occur on the summit of Mt. Kaala. 
Drosophila researchers have devoted intensive efforts to relocating 
this species at other sites because the species is considered important 
for genetic studies of the D. planitibia phylogeny group; 
unfortunately, these efforts have failed to relocate this species at 
other sites (Kaneshiro and Kaneshiro 1995; Science Panel 2005).
    Montgomery (1975) determined that Drosophila substenoptera larvae 
inhabit only the decomposing bark of Cheirodendron sp. trees (family 
Araliaceae) and Tetraplasandra sp. trees (family Araliaceae) in 
localized patches of wet forest habitat.

Drosophila tarphytrichia

    Drosophila tarphytrichia was described by Hardy (1965) from 
specimens collected from Manoa Falls on Oahu in 1949. This species is 
closely related to D. vesciseta based on the structure of the male 
genitalia (Kaneshiro et al.1995), but can be differentiated by distinct 
wing markings and the ornamentation of the front legs of the male. The 
thorax is almost entirely yellow to red with a tinge of brown on the 
top. The legs are yellow, with the tip of the front leg strongly 
flattened laterally and with a dense clump of black hairs. This species 
is 0.148 in (3.70 mm) long with wings 0.2 in (4.0 mm) long (Kaneshiro 
and Kaneshiro 1995).
    Drosophila tarphytrichia was historically known from both the 
Koolau and the Waianae Mountains between 2,000 and 2,800 ft (610 to 853 
m) above sea level. A total of 31 individuals were recorded on 36 
survey dates between 1965 and 1999 (Kaneshiro, in litt., 2005a). 
Drosophila tarphytrichia is now apparently extirpated from the Koolau 
range where it was originally discovered near Manoa Falls, and is 
presently known from four localities in the Waianae Mountains 
(Kaneshiro and Kaneshiro 1995; HBMP, in litt., 2005; K. Kaneshiro, in 
litt., 2005a).
    The larvae of Drosophila tarphytrichia feed only within the 
decomposing portions of the stems and branches of Charpentiera sp. 
trees (family Amaranthaceae) in mesic forest habitat (Montgomery 1975).

Previous Federal Action

    Ten of these 12 species were classified as candidates for listing 
in the February 28, 1996, Notice of Review of Plant and Animal Taxa 
That Are Candidates for Listing as Endangered or Threatened Species 
(Notice of Review) (61 FR 7596). The remaining two species, Drosophila 
differens and D. ochrobasis, were classified as candidates for listing 
in the Notice of Review dated September 19, 1997 (62 FR 49398). 
Candidates are those taxa for which the Service has on file substantial 
information on biological vulnerability and threats to support 
preparation of listing proposals.
    On January 17, 2001, we published a proposed rule to list as 
endangered the 12 species of Hawaiian picture-wing flies (66 FR 3964), 
which included a detailed history of Federal actions completed prior to 
the publication of the proposal. At that time, we did not propose 
critical habitat for the 12 picture-wing flies. In the proposed rule 
and associated notifications, we requested that all interested parties 
submit comments, data, or other information that might contribute to 
the development of a final rule. A 60-day comment period on the January 
17, 2001, proposal closed on March 19, 2001; we later reopened the 
comment period, as discussed below (see Summary of Comments and 
Recommendations section).
    On February 28, 2005, the Center for Biological Diversity (CBD) 
filed a lawsuit in the District of Oregon alleging that the Service 
failed to take action following issuance of a proposed rule to list 12 
species of picture-wing flies and for failure to designate critical 
habitat for the species (Center for Biological Diversity v. Allen, CV-
05-274-HA). CBD and the Service subsequently agreed to settle the case. 
Pursuant to the settlement agreement approved by the

[[Page 26841]]

United States District Court for the District of Hawaii on August 31, 
2005, the Service must make a final listing decision for these 12 
Hawaiian picture-wing flies by May 1, 2006, and if prudent and 
determinable, propose critical habitat by September 15, 2006, and 
finalize critical habitat by April 17, 2007. However, the Service will 
propose critical habitat for 12 species of picture-wing flies within 60 
days of the publication of this final rule.

Summary of Comments and Recommendations

    In the proposed rule published on January 17, 2001 (66 FR 3964), we 
requested that all interested parties submit written comments on the 
proposal by March 19, 2001. We also contacted appropriate Federal and 
State agencies, scientific experts and organizations, and other 
interested parties and invited them to comment on the proposal. 
Newspaper notices inviting general public comment were published in the 
Honolulu Advertiser. No requests for a public hearing were received.
    Because the proposed rule was published in 2001, and public 
outreach was conducted in 2001, we sought additional public comment on 
the proposed rule by reopening the public comment period from October 4 
to November 3, 2005 (70 FR 57851). We again reopened the comment period 
from November 18 to December 2, 2005 (70 FR 69922). The reopened 
comment periods (and associated notifications in local media and via 
direct mailing) gave interested parties additional time to consider the 
information in the proposed rule and provide comments and new 
information.
    During the comment periods for the proposed rule, we received nine 
written comments. Of those comments received, one commenter opposed the 
final listing, five commenters stated support for the final listing, 
one commenter expressed concern about unrestricted collecting of the 
flies, one commenter provided additional information regarding a fire 
management plan, and one commenter stated concerns about the potential 
impacts of the listing and critical habitat designation on private 
lands.

Peer Review

    In 2005, in accordance with our peer review policy published on 
July 1, 1994 (59 FR 34270), we solicited opinions from researchers, 
land managers, and State officials. All 16 individuals solicited have 
expertise with the species and the geographic regions where the species 
occur, and are familiar with conservation biology principles. We 
received written comments from two experts and incorporated their 
information into this final rule. One of the peer reviewers has a 
doctorate degree based upon study and research concerning Hawaiian 
Drosophila biology, evolution, genetics, and ecology research. The 
other holds a doctorate in insect taxonomy and has studied Hawaiian 
picture-wing flies for the past 10 years while working as a research 
scientist for the U.S. Geological Survey.
    One peer reviewer suggested the threats described in the proposed 
rule may not include all of the factors affecting the 12 flies, 
including factors causing their reduction in numbers. The reviewer 
noted that at least 3 of the 12 flies proposed for listing have 
demonstrated an apparent habitat shift upward in elevation, and 
suggested that global warming and increased temperatures on the 
Hawaiian Islands may be the cause. The reviewer suggested additional 
research was needed to validate the theory.
    This same reviewer provided a synopsis, based partly on the 
reviewer's own 35 years of Hawaiian Drosophila research, surveys, and 
personal observations in the field and laboratory while employed as a 
researcher with the University of Hawaii, emphasizing three major 
threats to the Hawaiian picture-wing flies including predation by wasps 
(Vespula sp.), habitat destruction by feral ungulates, and the effects 
of global warming.
    The other peer reviewer provided specific information about 
firsthand observations and evidence of declines in numbers and 
populations of three Drosophila species found on the island of Hawaii. 
This peer reviewer provided information and observational accounts of 
the effects of feral ungulates, rats, tipulid flies, and scolytid 
beetles upon picture-wing fly host plants and habitat and also the 
effects of predation by wasps (Vespula sp.) upon the 12 species. This 
peer reviewer also provided comments detailing the taxonomic 
differences recognized by Drosophila experts which establish the 12 
flies as separate and distinct species.
    Substantive information provided in all public comments, including 
the peer review process, either has been incorporated directly into 
this final rule or is addressed below.
    Comment 1: The U.S. Army's Schofield Barracks Integrated Wildfire 
Management Plan significantly reduces the threats to Drosophila aglaia 
and D. obatai and therefore could reduce the imminent need to list 
these species.
    Our Response: We agree that the Department of the Army's 
implementation of the completed Integrated Wildfire Management Plan 
will reduce the threat of fire caused by the Department of the Army to 
the habitat of these two picture-wing flies. However, the Integrated 
Wildfire Management Plan does not address the additional threats to 
these species' habitat within the Puu Pane area, including feral 
ungulates, nonnative weed plants, and predation by insect predators.
    Comment 2: Several commenters were concerned that the listing, and 
especially the critical habitat designation for the flies, could impact 
native Hawaiian traditional and customary gathering rights and access, 
and could jeopardize cooperative conservation efforts.
    Our Response: Private lands are likely to be important to the 
conservation of many of the picture-wing flies, and we appreciate all 
opportunities to work in partnerships with private landowners, the 
State, and others to further their conservation. The Act requires the 
listing of a species to be based solely on whether a species is 
affected by any of the five factors (see Summary of Factors section) to 
such an extent that they are in danger of becoming extinct (endangered 
status) or likely to become endangered (threatened status).
    According to the court settlement related to this final listing, we 
are required to propose critical habitat if appropriate by September 
15, 2006. The public will be invited to comment on any such proposal. 
Unlike when a species is listed, economic factors and conservation 
partnerships are considered in a critical habitat designation. Under 
the Act, the Secretary has the discretion to exclude areas from 
critical habitat designation if the benefits of exclusion outweigh the 
benefits of designation and such exclusion would not result in 
extinction of the species.
    Comment 3: The proposed listing of the 12 picture-wing flies lacks 
stringent research, detailed surveys, and up-to-date population 
assessments, and the data were spotty, hearsay, incomprehensive, and 
not empirical.
    Our Response: Since 1963, a mutli-disciplinary team of biologists 
have researched Drosophila through the University of Hawaii affiliated 
Hawaiian Drosophila Project. This effort has resulted in over 500 
scientific papers being published and the taxonomic description of over 
500 species of Drosophila. The information used to prepare this rule 
includes peer reviewed publications, unpublished literature, and 
written and verbal communications from research and field studies 
covering

[[Page 26842]]

a period of over 40 years of Hawaiian Drosophila research. In addition, 
this final rule includes information gathered after the proposed rule 
was published and a review of all available information on these 
species was made during science and managers review panels conducted in 
November 2005. While we acknowledge that additional systematic surveys 
for the picture-wing fly species and host plants would assist with 
understanding population trends and status, we believe we have ample 
information on habitat threats and trends in distribution for the 
picture-wing flies covered by this final rule.

Extinction Risk Assessment and Listing Decision Making Process

    The Service convened a panel of three scientists from outside the 
Service with expertise in Hawaiian Drosophila to help synthesize and 
address uncertainties in the scientific information available for these 
12 picture-wing flies, particularly threats to their existence (Science 
Panel 2005). A second panel made up of four Service managers and a 
State manager participated in related policy discussions and considered 
the available information including assessment of status, threats, and 
extinction risks. These two panels reviewed the available information 
and participated in a combined panel meeting in November 2005, prior to 
the close of the final comment period.

Science Panel

    The purpose of the Science Panel was to assess threats for each of 
the 12 picture-wing flies, identify and resolve areas of scientific 
uncertainty, and discuss extinction risks in a carefully structured 
format. The panelists discussed taxonomy, adaptive radiation of 
picture-wing flies, hybridization, sexual selection, survey methods, 
Drosophila lifecycle, and species' distribution (Science Panel 2005). 
They then discussed specific threats to each of the flies. Following 
this information review, each expert was asked to rank independently 
the severity of each threat on a scale of 1 to 5 and explain why they 
assigned a given rank to a threat. Then the other scientists were given 
the opportunity to change their rankings based on the rationales 
presented. In this manner three ranks (one for each scientist) were 
assigned to each threat factor for each species (Science Panel 2005). 
The scientific panel discussed the strengths and weaknesses of the 
various data and hypotheses about threats to the flies. Results from 
these exercises revealed little disagreement among the scientists 
regarding the type and degree of threats faced by each species. Each 
scientist was separately asked, based on his/her threats assessment and 
experience, to categorize extinction risk for each species as high, 
medium, or low over the next 40 years. The results of this exercise are 
presented in Table 2.

 Table 2.--Science Panel Categorization of Extinction Risk (H=High, M=Medium, L=Low) Over the Next 40 Years for
                                         12 Hawaiian Picture-Wing Flies
----------------------------------------------------------------------------------------------------------------
                 Species                              Island                         Extinction risk
----------------------------------------------------------------------------------------------------------------
Drosophila aglaia........................  Oahu........................  H H H
D. differens.............................  Molokai.....................  M H H
D. hemipeza..............................  Oahu........................  M M M
D. heteroneura...........................  Hawaii......................  H M M
D. montgomeryi...........................  Oahu........................  H M H
D. mulli.................................  Hawaii......................  M M M
D. musaphilia............................  Kauai.......................  H H H
D. neoclavisetae.........................  Maui........................  H H H
D. obatai................................  Oahu........................  H H H
D. ochrobasis............................  Hawaii......................  H H M
D. substenoptera.........................  Oahu........................  H M M
D. tarphytrichia.........................  Oahu........................  H H H
----------------------------------------------------------------------------------------------------------------

Manager Panel

    The manager panel reviewed background materials, interacted with 
the science panel during their risk assessment exercise, and 
participated in general and specific discussions about the definition 
of threatened and endangered. Following these discussions, the managers 
were asked to give their separate opinions as to whether each of the 12 
species of fly should be listed as endangered, listed as threatened, or 
withdrawn. The managers based their assessment on the information in 
the record, including comments previously received, the information 
presented by the individual mem bers of the science panel, information 
gaps and uncertainty, the number and severity of the threats affecting 
each species, and mitigating circumstances that might ameliorate one or 
more of those threats. Each manager was asked to explain their opinion 
and then the managers were given the opportunity to change their 
opinion based on the rationale presented by the other managers. The 
manager's panel presented its recommendations to the Regional Director. 
Subsequent to this, a recommendation of the Regional Director was 
forwarded to the Director for a final decision.
    This rule is based on the record of these discussions and all 
relevant and available information pertaining to the threats and status 
of the species.

Summary of Factors Affecting the Species

    Section 4 of the Act and its implementing regulations (50 CFR 424) 
set forth the procedures for adding species to the Federal list of 
endangered and threatened species. A species may be determined to be an 
endangered or threatened species due to one or more of the five factors 
described in section 4(a)(1) of the Act. The five listing factors are: 
(1) The present or threatened destruction, modification, or curtailment 
of its habitat or range; (2) overutilization for commercial, 
recreational, scientific, or educational purposes; (3) disease or 
predation; (4) the inadequacy of existing regulatory mechanisms; and 
(5) other natural or manmade factors affecting its continued existence.
A. The Present or Threatened Destruction, Modification, or Curtailment 
of its Habitat or Range
    Native vegetation on all the main Hawaiian Islands has undergone 
extreme alteration because of past and present land management 
practices, including ranching, introduction of nonnative plants and 
animals, and agricultural development (Cuddihy and Stone 1990). The 
primary threat facing

[[Page 26843]]

these picture-wing flies is the ongoing loss of habitat caused by feral 
animals and nonnative plants (Kaneshiro and Kaneshiro 1995).
    Feral ungulates have devastated native vegetation in many areas of 
the Hawaiian Islands (Cuddihy and Stone 1990). Because the endemic 
Hawaiian flora evolved without the presence of browsing and grazing 
ungulates, many plant groups have lost their adaptive defenses such as 
spines, thorns, stinging hairs, and defensive chemicals (University of 
Hawaii Department of Geography 1998), and cattle (Bos taurus), goats 
(Capra hircus), pigs (Sus scrofa), sheep (Ovis aries), Mouflon sheep 
(Ovis musimon), axis deer (Axis axis), and mule deer (Odocoileus 
hemionus) readily eat these plants as well as disturbing the soil and 
distributing nonnative plant seeds that can alter the ecosystem. In 
addition to the damage these nonnative herbivores cause by browsing and 
grazing, goats, pigs, and other ungulates that inhabit steep and remote 
terrain cause severe erosion of whole watersheds due to their foraging 
and trampling behaviors (Cuddihy and Stone 1990).
Feral Pigs (Sus scrofa)
    On the island of Hawaii, feral pigs are found from dry coastal 
grasslands through rain forests and into the sub-alpine zone of Mauna 
Kea and Mauna Loa. On Maui, Kauai, Oahu, and Molokai feral pigs inhabit 
rain forests, mesic forests, and grasslands (Cuddihy and Stone 1990). 
An increase in pig densities and expansion of their distribution has 
caused widespread damage to native vegetation (Cuddihy and Stone 1990). 
Feral pigs create open areas within forest habitat by digging up, 
eating, and trampling native species (Stone 1985). These open areas 
become fertile ground for non-native plant seeds spread through their 
excrement and by transport in their hair (Stone 1985). In nitrogen-poor 
soils, feral pig excrement increases nutrient availability, enhancing 
establishment of non-native weeds that are more adapted to richer soils 
than are native plants (Cuddihy and Stone 1990). In this manner, 
largely non-native forests replace native forest habitat (Cuddihy and 
Stone 1990).
    Foote and Carson (1995) found that pig exclosures on the Big Island 
supported significantly higher relative frequencies of picture-wing 
flies compared to other native and nonnative Drosophila species (7 
percent of all observations outside of the exclosure and 18 percent of 
all observations inside the exclosure) and their native host plants. 
Loope et al. (1991) showed that excluding pigs from a montane bog on 
northeastern Haleakala, Maui, resulted in an increase in native plant 
cover from 6 to 95 percent after 6 years of protection.
Goats (Capra hircus)
    Goats native to the Middle East and India were first successfully 
introduced to the Hawaiian Islands in 1792. Feral goats now occupy a 
wide variety of habitats from lowland dry forests to montane grasslands 
on Kauai, Oahu, Molokai, Maui, and Hawaii, where they consume native 
vegetation, trample roots and seedlings, accelerate erosion, and 
promote the invasion of nonnative plants (van Riper and van Riper 1982; 
Stone 1985). On the island of Oahu, encroaching urbanization and 
hunting pressure have tended to concentrate the goat population in the 
dry upper slopes of the Waianae Mountains (Kaneshiro and Kaneshiro 
1995). The population is increasing and spreading, becoming an even 
greater threat to the native habitat (Kapua Kawelo, U.S. Army, 
Environmental Division, pers. comm., 2005).
Cattle (Bos taurus)
    Large-scale ranching of cattle on the Hawaiian Islands began in the 
middle of the 19th century on the islands of Kauai, Oahu, Maui, and 
Hawaii (Cuddihy and Stone 1990). Large ranches, tens of thousands of 
acres in size, were developed on East Maui and Hawaii (Cuddihy and 
Stone 1990) where most of the State's large ranches still exist. 
Degradation of native forests used for ranching activities became 
evident soon after full-scale ranching began. Feral cattle now occupy a 
wide variety of habitats from lowland dry forests to montane 
grasslands, where they consume native vegetation, trample roots and 
seedlings, accelerate erosion, and promote the invasion of nonnative 
plants (van Riper and van Riper 1982; Stone 1985). Cattle grazing 
continues in several lowland regions in the northern portion of the 
Waianae Mountains of Oahu, and within many areas on the island of 
Hawaii.
Rats (Rattus spp.)
    Several species of nonnative rats, including the Polynesian rat 
(Rattus exulans), the roof rat (Rattus rattus), and the Norway rat 
(Rattus norvegicus), are present on the Hawaiian Islands and cause 
considerable environmental degradation (Staples and Cowie 2001). The 
seeds, bark, and flowers of several of the picture-wing flies' host 
plants, including Clermontia sp., Pleomele sp., and Pritchardia 
beccariana, are susceptible to grazing by all the rat species (Science 
Panel 2005; K. Magnacca, in litt., 2005; S. Montgomery, pers. comm., 
2005b). The grazing by rats causes host plant mortality, diminished 
vigor, and seed predation, resulting in reduced host plant fecundity 
and viability (Science Panel 2005; K. Magnacca, in litt., 2005; S. 
Montgomery, pers. comm., 2005b).
Fire
    Fire threatens species of Hawaiian picture-wing flies living in dry 
to mesic grassland, shrubland, and forests on both the islands of 
Hawaii and Oahu. A large factor in the alteration of Hawaiian dry and 
mesic regions in the past 200 years has been the increase in fire 
frequency, a condition to which the native flora is not adapted. The 
invasion of fire-adapted alien plants, especially Melinis minutiflora 
on Oahu and Pennisetum setaceum on Hawaii, facilitated by ungulate 
disturbance, has increased the susceptibility of native areas to 
wildfire and increased wildfire frequency. These plants can quickly 
reestablish following a fire and effectively outcompete less fire-
adapted native plants. This change in fire regime has reduced the 
amount of forest cover for native species (Hughes et al. 1991; 
Blackmore and Vitousek 2000) and resulted in an intensification of 
feral ungulate herbivory in the remaining native forest areas. The 
impact of an altered wildfire regime to these areas is a serious and 
immediate threat to the viability of the dry and mesic habitats that 
support over one-third of Hawaii's threatened and endangered species as 
well as Hawaiian picture-wing flies and their host plants (Hughes et 
al. 1991; Kaneshiro and Kaneshiro 1995; Blackmore and Vitousek 2000). 
Furthermore, Hawaiian picture-wing fly habitat damaged or destroyed by 
fire is more likely to be invaded and re-vegetated by nonnative plants 
that cannot be used as host plants by picture-wing flies (Kaneshiro and 
Kaneshiro 1995).
Island of Oahu--Drosophila aglaia, D. hemipeza, D. montgomeryi, D. 
obatai, D. substenoptera, and D. tarphytrichia
    The picture-wing flies on Oahu that are addressed in this rule 
(Drosophila aglaia, D. hemipeza, D. montgomeryi, D. obatai, D. 
substenoptera, and D. tarphytrichia) are threatened by the loss of 
habitat due to a variety of factors. Feral pigs and goats have 
dramatically altered the native vegetation (Kaneshiro and Kaneshiro 
1995; Science Panel 2005). These feral ungulates destroy host plant 
seedlings and habitat by the trampling action of their hooves and 
through the spread of seeds of nonnative plants (Cuddihy and Stone 
1995). Goats

[[Page 26844]]

directly feed upon the host plants of D. aglaia, D. obatai, and D. 
substenoptera, and contribute to erosion on some steeper slopes where 
the host plants occur; rats feed upon the host plants of D. hemipeza 
and D. obatai; pigs feed upon the host plants of D. hemipeza, D. 
montgomeryi, D. obatai, and D. substenoptera; and cattle feed upon the 
host plants of D. obatai and contribute to erosion on some steeper 
slopes where the host plants occur (S. Montgomery, pers. comm., 2005b).
    The invasion of several nonnative plants, particularly Psidium 
cattleianum, Lantana camara, Melinis minutiflora, Schinus 
terebinthifolius, and Clidemia hirta, further contributes to the 
degradation of native forests and the host plants of picture-wing flies 
(Kaneshiro and Kaneshiro 1995; Wagner et al. 1999; Science Panel 2005). 
Psidium cattleianum, Lantana camara, Melinis minutiflora, and Schinus 
terebinthifolius form dense stands, thickets, or mats that shade or 
outcompete native plants. M. minutiflora is a grass that burns readily, 
often grows at the border of forests, and tends to carry fire into 
areas with woody native plants (Smith 1985; Cuddihy and Stone 1990). It 
is able to spread prolifically after a fire and effectively outcompete 
less fire-adapted native plant species, ultimately creating a stand of 
nonnative grass where forest once stood. Lantana camera produces 
chemicals that inhibit the growth of other plant species (Smith 1985; 
Wagner et al. 1999).
    Drosophila aglaia and D. obatai occur at Puu Pane, located above 
the United States Army's Schofield Barracks Military Reservation. The 
gently sloping lands below Puu Pane are used as a live firing range, 
and ordnance-induced fires have been a common occurrence in this area 
(U.S. Army, in litt., 2005). The U.S. Army recently completed and is 
implementing an Integrated Wildfire Management Plan to reduce the risk 
and improve control of training-related fires in this area. As part of 
the Integrated Wildfire Management Plan, firebreak roads have been 
constructed around the perimeter of the live-fire training area. We 
believe that the Integrated Wildfire Management Plan will reduce the 
threat and magnitude of wildfires caused by the U.S. Army; however 
wildfires caused by