Endangered and Threatened Wildlife and Plants; Determination of Endangered Species Status for 15 Species on Hawaii Island

DEPARTMENT OF THE INTERIOR

Fish and Wildlife Service

50 CFR Part 17

[Docket No. FWS-R1-ES-2012-0070; 4500030113]

RIN 1018-AY09

Endangered and Threatened Wildlife and Plants; Determination of Endangered Species Status for 15 Species on Hawaii Island

AGENCY:

Fish and Wildlife Service, Interior.

ACTION:

Final rule.

SUMMARY:

We, the U.S. Fish and Wildlife Service (Service), determine endangered species status under the Endangered Species Act of 1973 (Act), as amended, for 15 species on the island of Hawaii. In addition, we are recognizing a taxonomic change for one Hawaiian plant currently listed as an endangered species and revising the List of Endangered and Threatened Plants accordingly. The effect of this regulation is to conserve these species under the Act.

DATES:

This rule is effective on November 29, 2013.

ADDRESSES:

This final rule is available on the Internet at

http://www.regulations.gov

and

http://www.fws.gov/pacificislands.

Comments and materials received, as well as supporting documentation used in preparing this final rule, are available for public inspection, by appointment, during normal business hours, at U.S. Fish and Wildlife Service, Pacific Islands Fish and Wildlife Office, 300 Ala Moana Boulevard, Room 3-122, Honolulu, HI 96850; by telephone at 808-792-9400; or by facsimile at 808-792-9581.

FOR FURTHER INFORMATION CONTACT:

Loyal Mehrhoff, Field Supervisor, U.S. Fish and Wildlife Service, Pacific Islands Fish and Wildlife Office, 300 Ala Moana Boulevard, Room 3-122, Honolulu, HI 96850; by telephone at 808-792-9400; or by facsimile at 808-792-9581. If you use a telecommunications device for the deaf (TDD), call the Federal Information Relay Service (FIRS) at 800-877-8339.

SUPPLEMENTARY INFORMATION:

Executive Summary

Why we need to publish a rule.

This is a final rule to list 15 species (13 plants, 1 insect (picture-wing fly), and 1 crustacean (anchialine pool shrimp)) from the island of Hawaii, in the State of Hawaii, as endangered species. In addition, in this final rule, we also recognize a taxonomic change for one endangered plant species, and revise the List of Endangered and Threatened Plants accordingly.

The basis for our action.

Under the Act, we determine that a species is an endangered or threatened species based on any of five factors: (A) The present or threatened destruction, modification, or curtailment of its habitat or range; (B) overutilization for commercial, recreational, scientific, or educational purposes; (C) disease or predation; (D) the inadequacy of existing regulatory mechanisms; or (E) other natural or manmade factors affecting its continued existence. We have determined that the 15 Hawaii Island species are currently in danger of extinction throughout all their ranges as the result of ongoing threats that include the destruction and modification of habitat from nonnative feral ungulates (e.g., pigs, goats); competition with nonnative plant and animal species; agricultural and urban development; wildfire, erosion, drought, and hurricanes; climate change; predation and herbivory; the inadequacy of existing regulatory mechanisms; human dumping of nonnative fish and trash; small numbers of individuals and populations; hybridization; the lack of reproduction in the wild; loss of host plants; and competition with nonnative tipulid flies (large crane flies). We fully considered comments from the public, including comments we received during a public hearing, and comments we received from peer reviewers, on the proposed rule.

Peer reviewers support our methods.

We obtained opinions from 11 knowledgeable individuals with scientific expertise to review our technical assumptions, to review our analysis, and to determine whether or not we used the best available information. Nine (2 plant reviewers, 2 picture-wing fly reviewers, and 5 of the 7 anchialine pool shrimp reviewers) of these 11 peer reviewers generally concurred with our methods and provided additional information, clarifications, and suggestions to improve this final rule. One shrimp peer reviewer recommended further surveys for the anchialine pool shrimp, and a second shrimp reviewer commented that we should proceed with caution regarding listing the shrimp due to the lack of biological information. A response to all peer review comments is provided elsewhere in this final rule.

The final critical habitat designation for

Bidens micrantha

ssp.

ctenophylla, Isodendrion pyrifolium,

and

Mezoneuron kavaiense,

as proposed in the

Federal Register

(77 FR 63928; October 17, 2012), is still under development and undergoing agency review. It will publish in the near future in the

Federal Register

under Docket No. FWS-R1-ES-2013-0028.

Previous Federal Actions

Federal actions for these species prior to October 17, 2012, are outlined in our proposed rule (77 FR 63928), which was published on that date. Publication of the proposed rule opened a 60-day comment period, which closed on December 17, 2012. In addition, we published a public notice of the proposed rule on October 20, 2012, in the local Honolulu Star Advertiser, West Hawaii Today, and the Hawaii Tribune Herald newspapers. On April 30, 2013, we published in the

Federal Register

a document (78 FR 25243) that made available and requested public comments on the draft economic analysis for the October 17, 2012, proposed critical habitat designation (77 FR 63928); announced a public information meeting and hearing to be held in Kailua-Kona, Hawaii Island, on May 15, 2013; and reopened the comment period on the October 17, 2012, proposed rule for an additional 30 days. This second comment period closed on May 30, 2013. In total, we accepted public comments on the October 17, 2012, proposed rule for 90 days.

Background

Hawaii Island Species Addressed in This Final Rule

The table below (Table 1) provides the scientific name, common name, and listing status for the species that are the subjects of this final rule.

Table 1—The Hawaii Island Species Addressed in This Final Rule

[Note that many of the species share the same common name]

Scientific name

Common name(s)

Listing status

Plants:

Bidens hillebrandiana

ssp.

hillebrandiana

kookoolau

Endangered.

Bidens micrantha

ssp.

ctenophylla

kookoolau

Endangered.

Cyanea marksii

haha

Endangered.

Cyanea tritomantha

aku

Endangered.

Cyrtandra nanawaleensis

haiwale

Endangered.

Cyrtandra wagneri

haiwale

Endangered.

Mezoneuron kavaiense

(taxonomic change accepted) (Formerly listed as

Caesalpinia kavaiense

)

uhiuhi

Endangered—Listed in 1986.

Phyllostegia floribunda

NCN

1

Endangered.

Pittosporum hawaiiense

hoawa, haawa

Endangered.

Platydesma remyi

NCN

Endangered.

Pritchardia lanigera

loulu

Endangered.

Schiedea diffusa

ssp.

macraei

NCN

Endangered.

Schiedea hawaiiensis

NCN

Endangered.

Stenogyne cranwelliae

NCN

Endangered.

Animals:

Drosophila digressa

picture-wing fly

Endangered.

Vetericaris chaceorum

anchialine pool shrimp

Endangered

1

NCN = no common name.

Taxonomic Change Since Listing for One Plant Species

We listed

Mezoneuron kavaiense

as an endangered species in 1986 (51 FR 24672; July 8, 1986), based on the taxonomic treatment of Hillebrand (1888, pp. 110-111). Following the reduction of

Mezoneuron

to

Caesalpinia

by Hattink (1974, p. 5), Geesink

et al.

(1990, pp. 646-647) changed the name to

Caesalpinia kavaiensis.

In 1989, the List of Endangered and Threatened Plants (List) was revised to identify the listed entity as

Caesalpinia kavaiense,

although the specific epithet was misspelled in the List (at that time the correct spelling for this entity was

Caesalpinia kavaiensis

). Recent phylogenetic studies support separation of

Mezoneuron

from

Caesalpinia

(Bruneau

et al.

2008, p. 710). The recognized scientific name for this species is

Mezoneuron kavaiense

(Wagner

et al.

2012, p. 37). The range of the species between the time of listing and now has not changed. Therefore, we recognize the listed species as

Mezoneuron kavaiense.

We are amending the List to reflect this taxonomic change, but this amendment does not in any way change the listed entity or its protections under the Act (16 U.S.C. 1531

et seq.

).

An Ecosystem-Based Approach to Listing 15 Species on Hawaii Island

On the island of Hawaii, as on most of the Hawaiian Islands, native species that occur in the same habitat types (ecosystems) depend on many of the same biological features and the successful functioning of that ecosystem to survive. We have therefore organized the species addressed in this final rule by common ecosystem. Although the listing determination for each species is analyzed separately, we have organized the individual analysis for each species within the context of the broader ecosystem in which it occurs to avoid redundancy. In addition, native species that share ecosystems often face a suite of common factors that may be a threat to them, and ameliorating or eliminating these threats for each individual species often requires the exact same management actions in the exact same areas. Effective management of these threats often requires implementation of conservation actions at the ecosystem scale to enhance or restore critical ecological processes and provide for long-term viability of those species in their native environment. Thus, by taking this approach, we hope not only to organize this final rule efficiently, but also to more effectively focus conservation management efforts on the common threats that occur across these ecosystems. Those efforts would facilitate restoration of ecosystem functionality for the recovery of each species, and provide conservation benefits for associated native species, thereby potentially precluding the need to list other species under the Act that occur in these shared ecosystems. In addition, this approach is in accord with the primary stated purpose of the Act (see section 2(b)): “to provide a means whereby the ecosystems upon which endangered species and threatened species depend may be conserved.”

We are listing the plants

Bidens hillebrandiana

ssp.

hillebrandiana, Bidens micrantha

ssp.

ctenophylla, Cyanea marksii, Cyanea tritomantha, Cyrtandra nanawaleensis, Cyrtandra wagneri, Phyllostegia floribunda, Pittosporum hawaiiense, Platydesma remyi, Pritchardia lanigera, Schiedea diffusa

ssp.

macraei, Schidea hawaiiensis,

and

Stenogyne cranwelliae;

and the animals

Drosophila digressa

and

Vetericaris chaceorum,

from Hawaii Island as endangered species. These 15 species (13 plants, 1 anchialine pool shrimp, and 1 picture-wing fly) are found in 10 ecosystem types: anchialine pool, coastal, lowland dry, lowland mesic, lowland wet, montane dry, montane mesic, montane wet, dry cliff, and wet cliff (Table 2).

Table 2—The 15 Hawaii Island Species and the Ecosystems Upon Which They Depend

Ecosystem

Species

Plants

Animals

Anchialine Pool

Vetericaris chaceorum.

Coastal

Bidens hillebrandiana

ssp.

hillebrandiana

Lowland Dry

Bidens micrantha

ssp.

ctenophylla

Lowland Mesic

Pittosporum hawaiiense

Drosophila digressa.

Pritchardia lanigera

Lowland Wet

Cyanea marksii

Cyanea tritomantha

Cyrtandra nanawaleensis

Cyrtandra wagneri

Phyllostegia floribunda

Platydesma remyi

Pritchardia lanigera

Montane Dry

Schiedea hawaiiensis

Montane Mesic

Phyllostegia floribunda

Drosophila digressa.

Pittosporum hawaiiense

Montane Wet

Cyanea marksii

Drosophila digressa.

Cyanea tritomantha

Phyllostegia floribunda

Pittosporum hawaiiense

Platydesma remyi

Pritchardia lanigera

Schiedea diffusa

ssp.

macraei

Stenogyne cranwelliae

Dry Cliff

Bidens hillebrandiana

ssp.

hillebrandiana

Wet Cliff

Cyanea tritomantha

Pritchardia lanigera

Stenogyne cranwelliae

For each species, we identified and evaluated those factors that adversely impact the species and that may be common to all of the species at the ecosystem level. For example, the degradation of habitat by nonnative ungulates is considered a threat to all 15 species, and is likely a threat to many, if not most or all, of the native species within a given ecosystem. We consider such a threat factor to be an “ecosystem-level threat,” as each individual species within that ecosystem faces a threat that is essentially identical in terms of the nature of the impact, its severity, its timing, and its scope. Beyond ecosystem-level threats, we further identified and evaluated threat factors that may be unique to certain species and that do not apply to all species under consideration within the same ecosystem. For example, the threat of predation by nonnative wasps is unique to the picture-wing fly

Drosophila digressa,

and is not applicable to any of the other 14 species. We have identified such threat factors, which apply only to certain species within the ecosystems addressed here, as “species-specific threats.”

Please refer to the proposed rule (77 FR 63928; October 17, 2012) for a description of the island of Hawaii and associated map, and for a description of the 10 ecosystems on Hawaii Island that support the 15 species. We have made minor revisions to our description of the anchialine pool ecosystem described in the proposed rule (77 FR 63928; October 17, 2012); therefore, we have included the revised version in its entirety in this final rule (see

Hawaii Island Ecosystems,

below).

Hawaii Island Ecosystems

There are 12 different ecosystems (anchialine pool, coastal, lowland dry, lowland mesic, lowland wet, montane dry, montane mesic, montane wet, subalpine, alpine, dry cliff, and wet cliff) recognized on the island of Hawaii. The 15 species addressed in this final rule occur in 10 of these 12 ecosystems (none of the 15 species are reported in subalpine and alpine ecosystems). The 10 Hawaii Island ecosystems that support the 15 species are described in the proposed rule (77 FR 63928; October 17, 2012), with the exception of a revised description of the anchialine pool ecosystem below; see Table 2 (above) for a list of the species that occur in each ecosystem type.

Anchialine Pools

Anchialine pools are land-locked bodies of water that have indirect underground connections to the sea, contain varying levels of salinity, and show tidal fluctuations in water level. Anchialine pool habitats can be distinguished from similar systems (i.e., tidal pools) in that they are land-locked with no surface connections to water sources either saline or fresh, but have subterranean hydrologic connections to both fresh and ocean water where water flows through cracks and crevices, and remain tidally influenced (Holthuis 1973, p. 3; Stock 1986, p. 91). Anchialine habitats are ecologically distinct and unique, and while widely distributed throughout the world, they only occur in the United States in the Hawaiian Islands (Brock 2004, pp. i, 2, and 12). In Hawaii, the anchialine pool ecosystem has been reported from Oahu, Molokai, Maui, Kahoolawe, and Hawaii Island. In the Hawaiian Islands, there are estimated to be 600 to 700 anchialine pools, with the majority occurring on the island of Hawaii (Brock 2004, p. i). Over 80 percent of the State's anchialine pools are found on the island of Hawaii, with a total of approximately 520 to 560 pools distributed over 130 sites along all but the island's northernmost and steeper northeastern shorelines. Characteristic animal species include crustaceans (e.g., shrimps, prawns, amphipods, isopods, etc.), several fish species, mollusks, and other invertebrates adapted to the pools' surface and subterranean habitats (Brock 2004, p. i; The Nature Conservancy (TNC) 2009, pp. 1-3). Generally, vegetation within the anchialine pools consists of various types of algal forms (blue-green, green, red, and golden-brown). The majority of Hawaii's anchialine pools occur in bare or

sparsely vegetated lava fields, although some pools occur in areas with various groundcover, shrub, and tree species (Chai

et al.

1989, pp. 2-24; Brock 2004, p. 35). The anchialine pool shrimp in this final rule,

Vetericaris chaceorum,

occurs in this ecosystem (Kensley and Williams 1986, pp. 417-437).

Description of the 15 Species

Below is a brief description of each of the 15 species, presented in alphabetical order by genus. Plants are presented first, followed by animals.

Plants

In order to avoid confusion regarding the number of locations of each species (a location does not necessarily represent a viable population, as in some cases there may only be one or a very few representatives of the species present), we use the word “occurrence” instead of “population.” Each occurrence is composed only of wild (i.e., not propagated and outplanted) individuals.

Bidens hillebrandiana

ssp.

hillebrandiana

(kookoolau), a perennial herb in the sunflower family (Asteraceae), occurs only on the island of Hawaii (Ganders and Nagata 1999, pp. 275-276). Historically,

B. hillebrandiana

ssp.

hillebrandiana

was known from two locations along the windward Kohala coastline, in the coastal and dry cliff ecosystems, often along rocks just above the ocean (Degener and Wiebke 1926, in litt.; Flynn 1988, in litt.). Currently, there are two known occurrences of

B. hillebrandiana

ssp.

hillebrandiana

totaling 40 or fewer individuals along the windward Kohala coast, in the coastal and dry cliff ecosystems. There are 30 individuals on the Pololu seacliffs, and 5 to 10 individuals on the seacliffs between Pololu and Honokane Nui (Perlman 1998, in litt.; Perlman 2006, in litt.). Biologists speculate that this species may total as many as 100 individuals with further surveys of potential habitat along the Kohala coast (Mitchell

et al.

2005b; PEPP 2006, p. 3).

Bidens micrantha

ssp.

ctenophylla

(kookoolau), a perennial herb in the sunflower family (Asteraceae), occurs only on the island of Hawaii (Ganders and Nagata 1999, pp. 271, 273). Historically,

B. micrantha

ssp.

ctenophylla

was known from the north Kona district, in the lowland dry ecosystem (HBMP 2010b). Currently, this subspecies is restricted to an area of less than 10 square miles (sq mi) (26 square kilometers (sq km)) on the leeward slopes of Hualalai volcano, in the lowland dry ecosystem in 6 occurrences totaling fewer than 1,000 individuals. The largest occurrence is found off Hina Lani Road with over 475 individuals widely dispersed throughout the area (Zimpfer 2011, in litt.). Another occurrence at Kealakehe was reported to have been abundant and common in 1992, but by 2010 had declined to low numbers (Whister 2007, pp. 1-18; Bio 2008, in litt.; HBMP 2010b; Whister 2008, pp. 1-11). In addition, there are three naturally occurring individuals in Kaloko-Honokohau National Historical Park (NHP) (Beavers 2010, in litt.), and three occurrences within close proximity to each other to the northeast of the park: Five individuals in an exclosure at Puuwaawaa Wildlife Sanctuary (HBMP 2010b); a few scattered individuals at Kaupulehu; and a few individuals on private land at Palani Ranch (Whistler 2007, pp. 1-18; Whistler 2008, pp. 1-11).

Bidens micrantha

ssp.

ctenophylla

has also been outplanted within Kaloko-Honokohau NHP (49 individuals), Koaia Tree Sanctuary (1 individual), and Puuwaawaa (5 individuals) (Boston 2008, in litt.; HBMP 2010b; Billings 2012, in litt.).

Cyanea marksii

(haha), a shrub in the bellflower family (Campanulaceae), is found only on the island of Hawaii. Historically,

C. marksii

was known from the Kona district, in the lowland wet and montane wet ecosystems (Lammers 1999, p. 457; HBMP 2010e). Currently, there are 27 individuals distributed among 3 occurrences in south Kona, in the lowland wet and montane wet ecosystems (PEPP 2007, p. 61). There is an adult and 20 to 30 juveniles (each approximately 1 inch (in) (2.54 centimeters (cm) tall)) in a lava tube in the Kona unit of the Hakalau National Wildlife Refuge (NWR) (PEPP 2007, p. 61), 3 adult individuals and 6 seedlings in the Kaohe pit crater in the South Kona FR (Perry 2012, in litt.), and 25 individuals on private land in south Kona (PEPP 2007, p. 61; Bio 2011, pers. comm.). Fruit has been collected from the individuals on private land, and 11 plants have been successfully propagated at the Volcano Rare Plant Facility (VRPF) (PEPP 2007, p. 61; Bio 2011, pers. comm.).

Cyanea tritomantha

(aku), a palmlike shrub in the bellflower family (Campanulaceae), is known only from the island of Hawaii (Pratt and Abbott 1997, p. 13; Lammers 2004, p. 89). Historically, this species was known from the windward slopes of Mauna Kea, Mauna Loa, Kilauea, and the Kohala Mountains, in the lowland wet, montane wet, and wet cliff ecosystems (Pratt and Abbott 1997, p. 13). Currently, there are 16 occurrences of

Cyanea tritomantha

totaling fewer than 400 individuals in the lowland wet, montane wet, and wet cliff ecosystems: 10 occurrences (totaling fewer than 240 individuals) in the Kohala Mountains (Perlman 1993, in litt.; Perlman 1995a, in litt.; Perlman and Wood 1996, pp. 1-14; HBMP 2010f; PEPP 2010, p. 60); 2 occurrences (totaling fewer than 75 individuals) in the Laupahoehoe Natural Area Reserve (NAR) (HBMP 2010f; Bio 2011, pers. comm.); 1 occurrence (20 adults and 30 juveniles) at Puu Makaala NAR (Perlman and Bio 2008, in litt.; Agorastos 2010, in litt.; HBMP 2010f; Bio 2011, pers. comm.); 1 occurrence with 10 to 20 individuals off Tom's Trail in the Upper Waiakea Forest Reserve FR (Perlman and Bio 2008, in litt.; Perry 2012, in litt.); and 2 occurrences (totaling fewer than 11 individuals) in Olaa Tract in Hawaii Volcanoes National Park HVNP (Pratt 2007a, in litt.; Pratt 2008a, in litt.; Orlando 2012, in litt.). In 2003, over 75 individuals were outplanted in HVNP's Olaa Tract and Small Tract; however, by 2010, less than one third of these individuals remained (Pratt 2011a, in litt.). In addition, a few individuals have been outplanted at Puu Makaala NAR and Upper Waiakea FR (Hawaii Department of Land and Natural Resources (HDLNR) 2006; Belfield 2007, in litt.; Agorastos 2010, in litt.).

Cyanea tritomantha

produces few seeds, and their viability tends to be low (Moriyasu 2009, in litt.)

Cyrtandra nanawaleensis

(haiwale), a shrub or small tree in the African violet family (Gesneriaceae), is known only from the island of Hawaii (Wagner and Herbst 2003, p. 29; Wagner

et al.

2005a—

Flora of the Hawaiian Islands database

). Historically,

C. nanawaleensis

was known only from the Nanawale FR and the adjacent Malama Ki FR in the Puna district, in the lowland wet ecosystem (St. John 1987, p. 500; Wagner

et al.

1988, in litt.; HBMP 2010g; Pratt 2011b, in litt.). Currently,

C. nanawaleensis

is known from 5 occurrences with approximately 160 individuals in the lowland wet ecosystem: 2 occurrences in Malama Ki FR totaling 70 individuals (Lau 2011, pers. comm.); 1 occurrence in Keauohana FR (with 56 individuals) (Magnacca 2011a, in litt.); 2 occurrences in the Halepuaa section of Nanawale FR (one with 28 mature and 65 immature plants at 200 feet (ft) (61 meters (m)) elevation, and a second occurrence with 9 mature and 57 immature plants at 270 ft (82 m)) (Johansen 2012, in litt.; Kobsa 2012, in litt.; Perry 2012, in litt.); and 1 occurrence with an unknown number of individuals on private lands in lower Puna (Perry 2012, in litt.). A total of

approximately 56 individuals have been outplanted in Halepuaa and Keauhana (Perry 2012, in litt.).

Cyrtandra wagneri

(haiwale), a shrub or small tree in the African violet family (Gesneriaceae), occurs only on the island of Hawaii (Lorence and Perlman 2007, p. 357). Historically,

C. wagneri

was known from a few individuals along the steep banks of the Kaiwilahilahi Stream in the Laupahoehoe NAR, in the lowland wet ecosystem (Perlman

et al.

1998, in litt.). In 2002, there were 2 known occurrences totaling fewer than 175 individuals in the Laupahoehoe NAR: One occurrence (totaling 150 individuals (50 adults and 100 juveniles)) along the steep banks of the Kilau Stream (Lorence

et al.

2002, in litt.; Perlman and Perry 2003, in litt.; Lorence and Perlman 2007, p. 359), and a second occurrence (with approximately 10 sterile individuals) along the slopes of the Kaiwilahilahi stream banks (Lorence and Perlman 2007, p. 359). Currently, there are no individuals remaining at Kaiwilahilahi Stream, and the individuals at Kilau Stream appear to be hybridizing with the endangered

Cyrtandra tintinnabula.

Biologists have identified only eight individuals at Kilau Stream that express the true phenotype of

Cyrtandra wagneri,

and only three of these individuals are reproducing successfully (PEPP 2010, p. 102; Bio 2011, pers. comm.).

Phyllostegia floribunda

(NCN), a perennial herb in the mint family (Lamiaceae), is found only on the island of Hawaii (Wagner 1999, p. 268; Wagner

et al.

1999b, p. 815). Historically,

P. floribunda

was reported in the lowland wet, montane mesic, and montane wet ecosystems at scattered sites along the slopes of the Kohala Mountains; southeast through Hamakua, Laupahoehoe NAR, Waiakea FR, and Upper Waiakea FR; and southward into Hilo, HVNP, and Puna. One report exists of the species occurring from north Kona and a few occurrences in south Kona (Cuddihy

et al.

1982, in litt.; Wagner

et al.

2005b—

Flora of the Hawaiian Islands database;

Perlman

et al.

2008, in litt.; HBMP 2010h; Bishop Museum 2011—

Herbarium Database

). Currently, there are 12 known occurrences of

P. floribunda

totaling fewer than 100 individuals, in the lowland wet, montane mesic, and montane wet ecosystems (Bruegmann 1998, in litt.; Giffin 2009, in litt.; HBMP 2010h): 2 occurrences within HVNP, at Kamoamoa (1 individual) (HBMP 2010h) and near Napau Crater (4 individuals) (Pratt 2005, in litt.; Pratt 2007b, in litt.; HBMP 2010h); 1 occurrence behind the Volcano solid waste transfer station (10 to 50 individuals) (Flynn 1984, in litt.; Perlman and Wood 1993—

Hawaii Plant Conservation Maps database;

Pratt 2007b, in litt.; HBMP 2010h); 1 occurrence (with an unknown number individuals) in the Wao Kele O Puna NAR (HBMP 2010h); 1 occurrence with 20 individuals in a fenced exclosure in the Upper Waiakea FR (Perry 2012, in litt.); at least 1 occurrence each (with a few individuals each) in the Puu Makaala NAR, Waiakea FR, and TNC's Kona Hema Preserve (PR) (Perry 2006, in litt.; Perlman 2007, in litt.; Giffin 2009, in litt.; PEPP 2008, pp. 106-107; Perlman

et al.

2008, in litt.; Pratt 2008a, in litt.; Pratt 2008b, in litt.; Agorastos 2010, in litt.); 2 occurrences (each with an unknown number of individuals) from the South Kona FR; 1 occurrence (one individual) in the Kipahoehoe NAR; and 1 occurrence (with an unknown number of individuals) in the Lapauhoehoe NAR (Moriyasu 2009, in litt.; HBMP 2010h; Agorastos 2010, in litt.). Since 2003, over 400 individuals have been outplanted at HVNP, Waiakea FR, Puu Makaala NAR, Honomalino in TNC's Kona Hema PR, and Kipahoehoe NAR (Bruegmann 2006, in litt.; HDLNR 2006, p. 38; Tangalin 2006, in litt.; Belfield 2007, in litt.; Pratt 2007b, in litt.; VRPF 2008, in litt.; VRPF 2010, in litt.; Bio 2008, in litt.; Agorastos 2010, in litt.). However, for reasons unknown, approximately 90 percent of the outplantings experience high seedling mortality (Pratt 2007b, in litt.; Van DeMark

et al.

2010, pp. 24-43).

Pittosporum hawaiiense

(hoawa, haawa), a small tree in the pittosporum family (Pittosporaceae), is known only from the island of Hawaii (Wagner

et al.

1999c, p. 1,044). Historically,

P. hawaiiense

was known from the leeward side of the island, from the Kohala Mountains south to Kau, in the lowland mesic, montane mesic, and montane wet ecosystems (Wagner

et al.

1999c, p. 1,044). Currently, there are 14 known occurrences totaling fewer than 175 individuals, from HVNP to Puu O Umi NAR, and south Kona, in the lowland mesic, montane mesic, and montane wet ecosystems: 1 occurrence in Puu O Umi NAR (several scattered individuals) (Perlman 1995b, in litt.); 1 occurrence (with a least one individual) in TNC's Kona Hema PR (Oppenheimer

et al.

1998, in litt.); 1 occurrence with 50 to 100 individuals at Kukuiopae in the South Kona FR (Perlman and Perry 2002, in litt.; Perry 2012, in litt.); 1 occurrence (with a few individuals) in the Manuka NAR (Perry 2011, in litt.); 8 occurrences (totaling fewer than 58 individuals) scattered within the Kahuku unit of HVNP; 1 occurrence in the Olaa FR (at least one individual), just adjacent to the Olaa Tract in HVNP; and 1 occurrence (with fewer than 6 individuals) at the Volcano solid waste transfer station (Wood and Perlman 1991, in litt.; McDaniel 2011a, in litt.; McDaniel 2011b, in litt.; Pratt 2011d, in litt.). Biologists have observed very low regeneration in these occurrences, which is believed to be caused, in part, by rat predation on the seeds (Bio 2011, pers. comm.).

Platydesma remyi

(NCN), a shrub or shrubby tree in the rue family (Rutaceae), occurs only on the island of Hawaii (Stone

et al.

1999, p. 1210; USFWS 2010, pp. 4-66—4-67, A-11, A-74). Historically,

P. remyi

was known from a few scattered individuals on the windward slopes of the Kohala Mountains and several small populations on the windward slopes of Mauna Kea, in the lowland wet and montane wet ecosystems (Stone

et al.

1999, p. 1210; HBMP 2010i). Currently,

P. remyi

is known from 8 occurrences totaling fewer than 40 individuals, all of which are found in the Laupahoehoe NAR or in closely surrounding areas, in the lowland wet and montane wet ecosystems: Along the banks of Kaiwilahilahi Stream in the Laupahoehoe NAR (unknown number of individuals) (Perlman and Perry 2001, in litt.; Bio 2008, in litt.; HBMP 2010i); near the Spencer Hunter Trail in the Laupahoehoe NAR (fewer than 17 individuals) (PEPP 2010, p. 102); in the central part of the Laupahoehoe NAR (5 to 6 scattered individuals) (HBMP 2010i); near Kilau (1 to 3 individuals) and Pahale (1 to 3 individuals) Streams in Laupahoehoe NAR; in the southeastern region of Laupahoehoe NAR (1 individual); in the Hakalau unit of the Hakalau NWR (1 individual) (USFWS 2010, p. 4-74—4-75); and in the Humuula region of the Hilo FR (2 individuals) (Bruegmann 1998, in litt.; Bio 2008, in litt.; PEPP 2008, p. 107; HBMP 2010i). According to field biologists, this species appears to be declining with no regeneration believed to be caused, in part, by rat predation on the seeds (Bio 2011, pers. comm.). In 2009, 29 individuals of

P. remyi

were outplanted in Laupahoehoe NAR (Bio 2008, in litt.). Their current status is unknown.

Pritchardia lanigera

(loulu), a medium-sized tree in the palm family (Arecaceae), is found only on the island of Hawaii (Read and Hodel 1999, p. 1,371; Hodel 2007, pp. 10, 24-25). Historically,

P. lanigera

was known from the Kohala Mountains, Hamakua district, windward slopes of Mauna Kea,

and southern slopes of Mauna Loa, in the lowland mesic, lowland wet, montane wet, and wet cliff ecosystems (Read and Hodel 1999, p. 1,371; HBMP 2010c). Currently,

P. lanigera

is known from 8 occurrences totaling fewer than 230 individuals scattered along the windward side of the Kohala Mountains, Kau FR, and TNC Kau Preserve, in the lowland mesic, lowland wet, montane wet, and wet cliff ecosystems. Approximately 100 to 200 individuals are scattered over 1 sq mi (3 sq km) in Waimanu Valley and surrounding areas (Wood 1995, in litt.; Perlman and Wood 1996, p. 6; Wood 1998, in litt.; Perlman

et al.

2004, in litt.; HBMP 2010c). There are at least five individuals in the back rim of Alakahi Gulch in Waipio Valley (HBMP 2010c), and five individuals in the Kau FR (Perry 2013, in litt.) According to field biologists, pollination rates appear to be low for this species, and the absence of seedlings and juveniles at known locations suggests that regeneration is not occurring, which they believe to be caused, in part, by beetle, rat, and pig predation on the fruits, seeds, and seedlings (Bio 2011, pers. comm.; Crysdale 2013, pers. comm.).

Schiedea diffusa

ssp.

macraei

(NCN), a perennial climbing herb in the pink family (Caryophyllaceae), is reported only from the island of Hawaii (Wagner

et al.

2005c—

Flowering Plants of the Hawaiian Islands database;

Wagner

et al.

2005d, p. 106). Historically,

S. diffusa

ssp.

macraei

was known from the Kohala Mountains, the windward slopes of Mauna Loa, and the Olaa Tract of HVNP, in the montane wet ecosystem (Perlman

et al.

2001, in litt.; Wagner

et al.

2005d, p. 106; HBMP 2010j). Currently, there is one individual of

S. diffusa

ssp.

macraei

on the slopes of Eke in the Kohala Mountains, in the montane wet ecosystem (Wagner

et al.

2005d, p. 106; Bio 2011, pers. comm.).

Schiedea hawaiiensis

(NCN), a perennial herb or subshrub in the pink family (Caryophyllaceae), is known only from the island of Hawaii (Wagner

et al.

2005d, pp. 92-96). Historically,

S. hawaiiensis

was known from a single collection by Hillebrand (1888, p. 33) from the Waimea region, in the montane dry ecosystem (Wagner

et al.

2005d, pp. 92-96). Currently,

S. hawaiiensis

is known from 25 to 40 individuals on the U.S. Army's Pohakuloa Training Area (PTA) in the montane dry ecosystem, in the saddle area between Moana Loa and Mauna Kea (Gon III and Tierney 1996 in Wagner

et al.

2005d, p. 92; Wagner

et al.

2005d, p. 92; Evans 2011, in litt.). In addition, there are over 150 individuals outplanted at PTA (Kipuka Alala and Kalawamauna), Puu Huluhulu, Puu Waawaa, and Kipuka Oweowe (Evans 2011, in litt.).

Stenogyne cranwelliae

(NCN), a vine in the mint family (Lamiaceae), is known only from the island of Hawaii. Historically,

S. cranwelliae

was known from the Kohala Mountains, in the montane wet and wet cliff ecosystems (Weller and Sakai 1999, p. 837). Currently, there are 6 occurrences of

S. cranwelliae

totaling fewer than 160 individuals in the Kohala Mountains, in the montane wet and wet cliff ecosystems: Roughly 1.5 sq mi (2.5 sq km) around the border between the Puu O Umi NAR and Kohala FR, near streams and bogs (ranging from 3 to 100 scattered individuals) (Perlman and Wood 1996, pp. 1-14; HBMP 2010k); Opaeloa, in the Puu O Umi NAR (3 individuals) (Perlman and Wood 1996, pp. 1-14; HBMP 2010k); Puukapu, in the Puu O Umi NAR (6-by-6-ft (2-by-2-m) “patch” of individuals) (HBMP 2010k); the rim of Kawainui Gulch (1 individual) (Perlman and Wood 1996, pp. 1-14; HBMP 2010k); along Kohakohau Stream, in the Puu O Umi NAR (a few individuals) (Perlman and Wood 1996, pp. 1-14; HBMP 2010k); and Waimanu Bog Unit in the Puu O Umi NAR (a “patch” of individuals) (Agorastos 2010, in litt.)

Animals

Drosophila digressa

(picture-wing fly), a member of the family Drosophilidae, was described in 1968 by Hardy and Kaneshiro and is found only on the island of Hawaii (Hardy and Kaneshiro 1968, pp. 180-1882; Carson 1986, p. 3-9). This species is small, with adults ranging in size from 0.15 to 0.19 in (4.0 to 5.0 mm) in length. Adults are brownish yellow in color and have yellow-colored legs and hyaline (shiny-clear) wings with prominent brown spots. Breeding generally occurs year round, but egg laying and larval development increase following the rainy season as the availability of decaying matter, which picture-wing flies feed on, increases in response to heavy rains. In contrast to most continental Drosophilidae, many endemic Hawaiian species are highly host-plant-specific (Magnacca

et al.

2008, p. 1).

Drosophila digressa

relies on the decaying stems of

Charpentiera

spp. and

Pisonia

spp. for oviposition (to deposit or lay eggs) and larval substrate (Magnacca

et al.

2008, pp. 11, 13; Magnacca 2013, in litt.). The larvae complete development in the decaying tissue before dropping to the soil to pupate (Montgomery 1975, pp. 65-103; Spieth 1986, p. 105). Pupae develop into adults in approximately 1 month, and adults sexually mature 1 month later. Adults live for 1 to 2 months. The adult flies are generalist microbivores (microbe eating) and feed upon a variety of decomposing plant material.

Drosophila digressa

occurs in elevations ranging from approximately 2,000 to 4,500 ft (610 to 1,370 m), in the lowland mesic, montane mesic, and montane wet ecosystems (Magnacca 2011a, pers. comm.). Historically,

D. digressa

was known from six sites: Moanuiahea pit crater on Hualalai, Papa in South Kona, Manuka FR, Kipuka 9 along Saddle Road, Bird Park in HVNP, and Olaa FR (Montgomery 1975, p. 98; Magnacca 2006, pers. comm.; HBMP 2010d; Magnacca 2011b, in litt.; Kaneshiro 2013, in litt.). Currently,

D. digressa

is known from only two locations, one population in the Manuka NAR within the Manuka FR, in the lowland mesic and montane mesic ecosystems, and a second population in the Olaa FR in the montane wet ecosystem (Magnacca 2011b, in litt.). The current number of individuals at each of these locations is unknown (Magnacca 2011b, in. litt.).

Vetericaris chaceorum

(anchialine pool shrimp) is a member of the family Procarididae, and is considered one of the most primitive shrimp species in the world (Kensley and Williams 1986, pp. 428-429). Currently known from only two locations on the island of Hawaii,

V. chaceorum

is one of seven described species of hypogeal (underground) shrimp found in the Hawaiian Islands that occur in anchialine pools (Brock 2004, p. 6). Relatively large in size for a hypogeal shrimp species, adult

Vetericaris chaceorum

measure approximately 2.0 in (5.0 cm) in total body length, excluding the primary antennae, which are approximately the same length as the adult's body length (Kensley and Williams 1986, p. 419). The species lacks large chelapeds (claws) (Kensley and Williams 1986, p. 426), which are a key diagnostic characteristic of all other known shrimp species.

V. chaceorum

is largely devoid of pigment and lacks eyes, although eyestalks are present (Kensley and Williams 1986, p. 419). Observations of

Vetericaris chaceorum

indicate the species is a strong swimmer and propels its body forward in an upright manner with its appendages held in a basket formation below the body. Forward movement is produced by a rhythmic movement of the thoracic and abdominal appendages, and during capture of some specimens,

V. chaceorum

escape tactics included only forward movement and a notable lack of tail flicking, which would allow backward movement and which is common to other shrimp species

(Kensley and Williams 1986, p. 426). No response was observed when the species was exposed to light (Kensley and Williams 1986, p. 418).

The feeding habits of

Vetericaris chaceorum

were unknown for decades with the only published data from Kensley and Williams (1986, p. 426), who reported that the gut contents of a captured specimen included large quantities of an orange-colored oil and fragments of other crustaceans, indicating that the species may be carnivorous upon its associated anchialine pool shrimp species. Sakihara (2012, in litt.) recently confirmed that

V. chaceorum

is carnivorous after observing

V. chaceorum

collected from Manuaka Natural Area Reserve actively feeding on

Halocaridina rubra

in the laboratory. In general, hypogeal shrimp occur within both the illuminated part of their anchialine pool habitat as well as within the cracks and crevices in the water table below the surface (Brock 2004, p. 6). The relative abundance of some Hawaii species is directly tied to food abundance (Brock 2004, p. 10). The lighted environment of anchialine pools offers refugia of high benthic productivity, resulting in higher population levels for the shrimp compared to the surrounding interstitial spaces often occupied by these species, albeit in lower numbers (Brock 2004, p. 10; Wada 2013, pers. comm.).

Although over 400 of the estimated 520 to 560 anchialine pool habitats have been surveyed on the island of Hawaii,

Vetericaris chaceorum

has only been documented from two locations: Lua o Palahemo, which is a submerged lava tube located on the southernmost point of Hawaii Island in an area known as Ka Lae (South Point) (Kensley and Williams 1986, pp. 417-418; Brock 2004, p. 2; HBMP 2010), and at Manuka, where only recently

V. chaceorum

was discovered in a series of pristine shallow anchialine pool complexes within and adjacent to the NAR, approximately 15 mi (25 km) northwest of Lua o Palahemo (Sakihara 2012, in litt.). The Service has concluded that the lack of detection of this species in the several hundred anchialine pools surveyed on the island of Hawaii since the 1970s suggests this species has a very limited range (Holthius 1973, pp. 1-128 cited in Sakihara 2012, pp. 83, 91, and 93; Maciolek and Brock 1974, pp. 1-73; Maciolek 1983, pp. 606-618; Kensley and Williams 1986, pp. 417-426; Maciolek 1987, pp. 1-23; Chai

et al.

1989, pp. 1-37; Chan 1995, pp. 1-31; Brock and Kam 1997, pp. 1-109; Bozanic 2004, p. 1; Brock 2004, pp. 1-60; Sakihara 2009, pp. 1-35; Sakihara 2012, pp. 83-95; Wada 2012, pers. comm.; Wada

et al.

2012, pp. 1-2; Sakihara 2013 in litt.). In total, only five individuals have been observed during one survey period in 1985 at Lua o Palahmo, and a total of seven individuals were observed in four pools during surveys conducted between 2009 and 2010 at Manuka. These two locations are described below.

Lua o Palahemo Site:

Age estimates for Lua o Palahemo range from as young as 11,780 years to a maximum of age of 25,000 years, based upon radio carbon data and timing of geophysical climatic events (Kensley and Williams 1986, pp. 417-418). Brock (2004, p. 18) states this lava tube is probably the second most important anchialine pool habitat in the State because of its unique connection to the ocean, the vertical size (i.e., depth), and the presence of a total of five different species including

Halocaridina palahemo, H. rubra, Procaris hawaiiana, Calliasmata pholidota,

and

Vetericaris chaceorum

. Lua o Palahemo is a naturally occurring opening (i.e., a surface collapse) into a large lava tube below. The opening measures approximately 33 ft (10 m) in diameter and is exposed to sunlight. Unlike most anchialine pools in the Hawaiian Islands, which have depths less than 4.9 ft (1.5 m) (Brock 2004, p. 3), Lua o Palahemo's deep pool includes a deep shaft with vertical sides extending downward about 46 ft (14 m) into the lava tube below, which branches in two directions, both ending in blockages (Holthuis 1974, p. 11; Kensley and Williams 1986, p. 418). At the subterranean level at the base of the opening, the lava tube runs generally north and south, extending northward for 282 ft (86 m) and southward for 718 ft (219 m), to a depth of 108 ft (33 m) below sea level (Kensley and Williams 1986, p. 418).

Manuka Site:

The anchialine pools at Manuka were first surveyed 1972 (Macioleck and Brock 1972, p. iii); however, this survey primarily covered only the southern extremity of the site. A more thorough survey of the Manuka coastline was conducted between 1989 and 1992 (20 pools along the southern coast of Manuka, which included both diurnal and nocturnal observations (Chan 1995, p. 1). These pools were then diurnally surveyed in 2004 (80 pools along the entire Manuka coastline) (Brock 2004, pp. 1-60), and again between 2008 and 2009 (80 pools along the entire Manuka coastline) (Sakihara 2009, pp. 1-35). The most recent and most comprehensive surveys of Manuka were conducted between 2009 and 2010, when Hawaii State biologists surveyed 81 pools at Manuka both day and night, which resulted in the discovery of

Vetericaris chaceorum

in 4 of the pools surveyed. Three of the pools are within Manuka NAR, and one pool is adjacent to the NAR, on unencumbered State land (collectively referred to as Manuka throughout this final rule) (Sakihara 2013, in litt.). This discovery documents the first observation of this species in almost three decades (Sakihara 2012, in litt.). Visual accounts made by the biologists estimate that

V. chaceorum

is established in four anchialine pools along the southern section of the NAR, approximately 15 mi (25 km) from Lua o Palahemo. A total of seven individuals of this species were observed in four pools around Awili Point and Keawaiki (Sakihara 2012, p. 89; Sakihara 2013, in litt.), although estimates of the total number of individuals are undeterminable due to the cryptic nature of this species (Sakihara 2012, in litt.). Sakihara (2012, in litt.) stated that the anchialine habitat at Manuka is considerably different than that of Lua o Palahemo, and is considered to be one of the most biologically valuable habitats of this type (Sakihara 2012, in litt.; Sakihara 2013, in litt.). The Manuka anchialine pools are characterized by shallow (less than 2 ft (0.5 m)) open pools dispersed throughout barren basaltic terrain. This observation expands the known habitat conditions that support

V. chaceorum

(Sakihara 2012, in litt.). According to Sakihara (2013, in litt.), it appears that three of the Manuka pools (the three pools closest to a jeep road) have a subterranean connection, although this has not been confirmed. Although anchialine pools have been surveyed in the Manuka area in the past (Maciolek and Brock 1974, pp. 1-80; Chan 1995, pp. 1-34; Brock 2004, pp. i-iv; Sakihara 2009, pp. 1-35; Sakihara 2012, pp. 83-95; Sakihara 2013 in litt.), the surveys conducted between 2009 and 2010 were the first to document the presence of

V. chaceorum

in this anchialine pool complex. In 1995, an anchialine pool shrimp matching the description of

V. chaceorum

was observed in at least one pool at Manuka NAR, but its identification was never confirmed (Brock 2004, p. 31; Sakihara 2012, p. 89).

Four surveys have been conducted at Lua o Palahemo (Maciolek and Brock 1974, pp. 1-73; Kensley and Williams 1986, pp. 417-426; Bozanic 2004, p. 1-3; Wada 2012, pers. comm.; Wada

et al.

2012, pp. 1-2), with five individuals observed during one survey in 1985. Five surveys have been conducted at Manuka (Maciolek and Brock 1974, pp.

1-73; Chan 1995, pp. 1-34; Brock 2004, pp. i-iv, 1-60; Sakihara 2009, pp. 1-35; Sakihara 2012, pp. 83-95; Sakihara 2013 in litt.), with seven individuals observed in four pools between 2009 and 2010. Because of the ability of hypogeal shrimp species to inhabit the interstitial and crevicular spaces in the water table bedrock surrounding anchialine pools, it is very difficult to estimate population size of a given species within a given area (Brock 2004, pp. 10-11). We are unable to estimate the population size of either occurrence of

Vetericaris chaceorum

given this behavior.

Summary of Comments and Recommendations

On October 17, 2012, we published a proposed rule to list 15 Hawaii Island species (13 plants, 1 picture-wing fly, and 1 anchialine pool shrimp) as endangered throughout their ranges, and to designate critical habitat for 3 plant species (77 FR 63928). The comment period for the proposal opened on October 17, 2012, for 60 days, ending on December 17, 2012. We requested that all interested parties submit comments or information concerning the proposed rule. We contacted all appropriate State and Federal agencies, county governments, elected officials, scientific organizations, and other interested parties and invited them to comment. In addition, we published a public notice of the proposed rule on October 20, 2012, in the local Honolulu Star Advertiser, West Hawaii Today, and the Hawaii Tribune Herald newspapers, at the beginning of the comment period. We received four requests for public hearings. On April 30, 2013, we published a document (78 FR 25243) reopening the comment period on the October 17, 2012, proposed rule (77 FR 63928), announcing the availability of our draft economic analysis (DEA) on the proposed critical habitat, and requesting comments on both the proposed rule and the DEA. In addition, in that same document (78 FR 25243; April 30, 2013), we announced a public information meeting and hearing, which was held in Kailua-Kona, Hawaii, on May 15, 2013.

During the comment periods, we received 33 comment letters, including the 11 peer review comment letters, on the proposed listing of 15 species, proposed taxonomic change for 1 endangered plant species, and proposed designation of critical habitat. In this final rule, we address only the comments regarding the proposed listing of 15 species and proposed taxonomic change for 1 plant species. Comments addressing the proposed critical habitat designation will be fully addressed in a separate rulemaking action, and published in the

Federal Register

at a later date.

Two commenters were State of Hawaii agencies ((1) Hawaii Department of Business, Economic Development, and Tourism's Hawaii Housing Finance and Development Corporation, and (2) Hawaii Department of Hawaiian Home Lands); one was a county agency (County of Hawaii Planning Department); two were Federal agencies; and 28 were nongovernmental organizations or individuals. During the May 15, 2013, public hearing, no individuals or organizations made comments on the proposed listing.

All substantive information related to the listing of the 15 species or the taxonomic change for 1 species provided during the comment periods has either been incorporated directly into this final determination or is addressed below. Comments received were grouped into general issues specifically relating to the proposed listing status of the 13 plants, or the picture-wing fly or anchialine pool shrimp, or the proposed taxonomic change for 1 plant species, and are addressed in the following summary and incorporated into the final rule as appropriate.

Peer Review

In accordance with our peer review policy published in the

Federal Register

on July 1, 1994 (59 FR 34270), we solicited expert opinions from 14 knowledgeable individuals with scientific expertise on the Hawaii Island plants, picture-wing fly, and anchialine pool shrimp, and their habitats, including familiarity with the species, the geographic region in which these species occur, and conservation biology principles. We received responses from 11 of these peer reviewers. Nine of these 11 peer reviewers generally supported our methodology and conclusions. One peer reviewer expressed concern regarding the lack of more recent survey data for the anchialine pool shrimp at Manuka, and was unaware of the recent surveys (between 2009 and 2010) conducted by Hawaii State biologists. Another commented that we should proceed with caution due to the lack of biological information regarding the shrimp. Three peer reviewers supported the Service's ecosystem-based approach for organizing the rule and for focusing on the actions needed for species conservation and management, and all 11 reviewers provided information on one or more of the Hawaii Island species, which was incorporated into this final rule (see also Summary of Changes from Proposed Rule). We reviewed all comments received from the peer reviewers for substantive issues and new information regarding the listing of 15 species and taxonomic change for 1 plant species. Peer reviewer comments are addressed in the following summary and incorporated into the final rule as appropriate.

Peer Review Comments on Plants

(1)

Comment:

One peer reviewer recommended that we include inundation by high surf and subsequent erosion, and the nonnative plant

Wedelia

[

Sphagneticola

]

trilobata

(wedelia), as threats to the plant

Bidens hillebrandiana

ssp.

hillebrandiana.

Our Response:

We have incorporated this information, as appropriate, into Summary of Changes from Proposed Rule, Table 3, and in the sections “Nonnative Plants in the Coastal Ecosystem” and “Habitat Destruction and Modification Due to Rockfalls, Treefalls, Landslides, Heavy Rain, Inundation by High Surf, Erosion, and Drought” under

Factor A. The Present or Threatened Destruction, Modification, or Curtailment of Habitat or Range

in this final rule (see below).

(2)

Comment:

One peer reviewer recommended that we include vandalism and trash dumping as threats to the plant

Bidens micrantha

ssp.

ctenophylla,

in the Kaloko Makai area.

Our Response:

We are aware that vandalism and trash dumping has occurred in the Kaloko Makai area near the individuals of

Bidens micrantha

ssp.

ctenophylla

in the past, although it has not been recently observed (Ball 2013, pers. comm.). We will continue to monitor this potential threat in that area.

(3)

Comment:

One peer reviewer informed us of an act of vandalism where approximately 150 ft (46 m) of fencing was removed from a fenced exclosure in the Upper Waiakea FR where individuals of the plant

Phyllostegia floribunda

are found. The fencing was repaired later in the same month (November 2012), and the plants appeared to suffer no adverse impacts.

Our Response:

We agree that vandalism is a potential threat to all fenced species. However, vandalism is not considered an imminent threat at this time because the frequency at which vandalism occurs and the degree of impact cannot be determined in advance of the incident occurring. We will continue to monitor the area and gather information on this potential threat.

(4)

Comment:

One peer reviewer suggested that we identify the nonnative plant

Paederia foetida

(skunk weed) as a threat to the plant

Cyrtandra

nanawaleensis

because it completely covers and smothers understory vegetation and outcompetes low-growing plants and small shrubs for light and space and that we identify

Psidium cattleianum

(strawberry guava) as a threat to

Cyanea tritomantha

because it forms dense stands in which few other plants can grow, displacing native vegetation through competition.

Our Response:

We have included this information in this final rule (see Summary of Changes from Proposed Rule, below).

(5)

Comment:

One peer reviewer supported the listing of the plants

Schiedea diffusa

ssp.

macraei, S. hawaiiensis,

and

Stenogyne cranwelliae

as endangered, and stated that we did a very thorough job of outlining the threats for these three species. In addition, this peer reviewer expressed appreciation for our emphasis on the anticipated effects of climate change in the proposed rule.

Our Response:

We appreciate the support from this peer reviewer regarding our threats analysis, and our discussion on the anticipated threats from climate change. All 15 species we are listing in this final rule may be especially vulnerable to the effects of climate change due to their small number of populations and individuals, as well as highly restricted ranges. Environmental changes that may affect these species are expected to include habitat loss or alteration and changes in disturbance regimes (e.g., storms, hurricanes, and drought).

(6)

Comment:

One peer reviewer stated that climate change appears to be having especially serious effects on

Schiedea

species occurring in dry habitats due to death of adult plants, presumably through drought, failure to regenerate due to drought, and increased fire frequency. Drought may have a pronounced effect on

Schiedea hawaiiensis.

Our Response:

We agree that drought is a threat to

Schiedea hawaiiensis,

for the reasons mentioned above (see also “Habitat Destruction and Modification by Fire” and “Habitat Destruction and Modification Due to Rockfalls, Treefalls, Landslides, Heavy Rain, Inundation by High Surf, Erosion, and Drought” under

Factor A. The Present or Threatened Destruction, Modification, or Curtailment of Habitat or Range,

below).

(7)

Comment:

One peer reviewer stated that

Schiedea diffusa

ssp.

macraei

and

S. hawaiiensis

are obligate autogamous species (i.e., reproduces by self-pollination) and facultative autogamous (i.e., reproduces by self- and cross-pollination), respectively. Because both of these species are hermaphroditic and autogamous, they are capable of regenerating from single individuals, and may not be severely hampered by inbreeding depression. Unfortunately, autogamous species of

Schiedea

also appear to be short-lived, emphasizing the importance of appropriate conditions for regeneration.

Our Response:

We agree that the obligate and facultative autogamous nature of

Schiedea diffusa

ssp.

macraei

and

S. hawaiiensis,

respectively, in addition to being hermaphroditic, afford these species the ability to regenerate from single individuals and may not be severely hampered by inbreeding depression. However, there are other negative impacts that can result from low number of individuals (e.g., random demographic fluctuations; climate change effects; and localized catastrophes, such as hurricanes, drought, rockfalls, landslides, and disease outbreaks (Pimm

et al.

1988, p. 757; Mangel and Tier 1994, p. 607). Any of these stressors represent threats that can lessen the chances of survival for these species in the wild. We agree that the short-lived nature of these species increases the importance for appropriate conditions for regeneration, and have added this information to our files.

(8)

Comment:

One peer reviewer pointed out that it was incorrect to state, in our proposed rule (77 FR 63928; October 17, 2012) on page 63931, that

Mezoneuron

was listed in error as

Caesalpinia kavaiense

in 50 CFR 17.12, because at the time of the listing (51 FR 24672; July 8, 1986), this was the accepted name applied to the taxon. The peer reviewer stated that it is important to emphasize that names of taxa typically may change during the course of standard taxonomic investigations, and these changes do not affect the validity of conservation concerns for the taxon in question.

Our Response:

We wish to clarify the error described in the October 17, 2012 (77 FR 63928), proposed rule regarding

Mezoneuron kavaiense.

The error described in the proposed rule refers to the entry in the 1989 List of Endangered and Threatened Plants (50 CFR 17.12), where this taxon was revised and the specific epithet was misspelled as

Caesalpinia kavaiense

(instead of

Caesalpinia kavaiensis

). Subsequent taxonomic revision resulted in the currently recognized scientific name for the listed entity,

Mezoneuron kavaiense,

which we accept in this final rule.

(9)

Comment:

One peer reviewer pointed out that under our description of the lowland dry ecosystem, we incorrectly wrote “high rates of diversity and endemism” when technically it should read “high levels of diversity and endemism,” as rate is a process occurring over time.

Our Response:

We agree with the peer reviewer.

Peer Review Comments on the Picture-Wing Fly

(10)

Comment:

One peer reviewer provided additional information regarding the host plants for

Drosophila digressa.

Although

D. digressa

has only been reared from

Charpentiera

spp., at Manuka,

D. digressa

was found in a

Pisonia sandwicensis

treefall with a considerable number of rotten branches. A large number of individuals of

D. digressa

were found in a small area, indicating a local breeding group rather than vagrant individuals. The only

Charpentiera

spp. in this area are a few trees in a pit crater, over 0.62 mi (1 km) from the known location of

D. digressa

on

Pisonia sandwicensis.

This reviewer further stated that many native

Drosophila

species that breed in either

Charpentiera

spp. or

Pisonia

spp. are also able to use both plants. According to the reviewer, while this ability of

D. digressa

to use both tree species as host plants expands its potential habitat slightly, it does not do so by a great deal, as

Pisonia sandwicensis

and

P. brunoniana

[two of the three species of

Pisonia

on Hawaii Island] are only found on Hawaii Island at the sites where

D. digressa

is already known (Olaa and Manuka), or where the forest is currently too open and dry to support this species of picture-wing fly (Kipuka Pualulu and Puu Waawaa cone).

Pisonia umbellifera

can be found at lower elevations on the windward side of the island, such as gulches on the east slopes of Kohala and Mauna Kea below 1,500 ft (457) m, but

D. digressa

has never been recorded from these areas or elevation. Species of

Pisonia

face most of the same threats as species of

Charpentiera

(i.e., goat and cattle browsing of leaves and seedlings, pig rooting of seedlings, and desiccation of habitat from drought and subsequent fires at Manuka). The reviewer concludes that even if

Pisonia

spp. at Manuka survive the [ongoing] drought, the habitat will likely be too dry to support

D. digressa.

Our Response:

We appreciate this information regarding

Drosophila digressa

and have incorporated this new information, as appropriate, in this final rule (see above, Description of the 15 Species; see below, Summary of Changes from Proposed Rule, “Habitat Destruction and Modification by Introduced Ungulates” (

Factor A. The Present or Threatened Destruction, Modification, or Curtailment of Habitat or Range

), “Predation and Herbivory”

(

Factor C. Disease or Predation

), and “Loss of Host Plants” (

Factor E. Other Natural or Manmade Factors Affecting Their Continued Existence

)).

(11)

Comment:

One peer reviewer stated that the drought-associated ohia [

Metrosideros polymorpha

] dieback occurring at Manuka adversely affects

Drosophila digressa

by allowing more sunlight into the understory, increasing the temperature and lowering humidity. This increases the stress on the picture-wing flies and their host plants, as well as increasing opportunities for invasive plants to become established. The extraordinary amount of dead wood accumulation at Manuka means that any fire that occurs there likely would be extremely damaging. A fire resulting from a similar scenario at Kealakekua Ranch a year or two ago produced smoke that covered most of the island and burned for weeks because it is nearly impossible to fight fire in such dense brush.

Our Response:

We appreciate the additional information provided regarding the drought-associated ohia dieback at Manuka and

Drosophila digressa,

and we have included this new information in our final rule, as appropriate, in “Habitat Destruction and Modification Due to Rockfalls, Treefalls, Landslides, Heavy Rain, Inundation by High Surf, Erosion, and Drought” (

Factor A. The Present or Threatened Destruction, Modification, or Curtailment of Habitat or Range

) in this final rule (see below).

Peer Review Comments on the Anchialine Pool Shrimp

(12)

Comment:

One peer reviewer commented that the field surveys cited in our proposed rule are not adequate, and that more surveys should be conducted at other sites such as Manuka, Hawaii. The peer reviewer also recommended that the analysis of listing

Vetericaris chaceorum

as endangered should be based on the number of field surveys conducted, the number of pools surveyed, the number of locations surveyed, trapping surveys, day and night surveys, and seasonal surveys.

Our Response:

We are required to make listing determinations solely on the basis of the best scientific and commercial data available, and, for the reasons described here, we have concluded that the number and locations of surveys are adequate to determine that

Vetericaris chaceorum

appears to be restricted to a limited number of pools in the southern portion of the island of Hawaii, and that

V. chaceorum

faces threats from habitat degradation and destruction and from predation such that it is in danger of extinction throughout its range. There are between 600 and 700 anchialine pools in the Hawaiian Islands and approximately 80 percent (approximately 520 to 560) occur on Hawaii Island. Over 400 pools have been surveyed on Hawaii Island alone since the 1970s, and

V. chaceorum

has only been documented from two locations: Lua o Palahemo and Manuka, where

V. chaceorum

was recently (between 2009 and 2010) discovered in a series of pristine shallow anchialine pool complexes within and adjacent to Manuka NAR (Holthius 1973, pp. 1-128 cited in Sakihara 2012, pp. 83, 91, and 93; Maciolek and Brock 1974, pp. 1-73; Maciolek 1983, pp. 606-618; Maciolek 1987, pp. 1-23; Chai

et al.

1989, pp. 1-37; Chan 1995, pp. 1-31; Brock and Kam 1997, pp. 1-109; Brock 2004, pp. 1-60; Sakihara 2009, pp. 1-35; Sakihara 2012, pp. 83-95; Wada

et al.

2012, pp. 1-2). This reviewer was apparently unaware that Hawaii State biologists conducted surveys at Manuka between 2008 and 2009, and again between 2009 and 2010 (Sakihara 2009, pp. 1-35; Sakihara 2012, pp. 83-95). Several other peer reviewers stated that the Service used the best available scientific and commercial data to document the presence or absence of

V. chaceorum

in anchialine pools around Hawaii Island.

Under the Act, we determine whether a species is an endangered species or a threatened species because of any of five factors (see Summary of Factors Affecting the 15 Species, below), and we are required to make listing determinations solely on the basis of the best scientific and commercial data available, pursuant to section 4(b)(1)(A) of the Act. Based on the best available information we determined that

V. chaceorum

faces threats from habitat destruction and modification by feral goats and cattle at Lua o Palahemo; dumping of trash and introduction of nonnative fish at Lua o Palahemo; and introduction of nonnative fish at the pools at Manuka (see Summary of Factors Affecting the 15 Species, below).

(13)

Comment:

One peer reviewer questioned the importance of flushing to the functioning of the anchialine pool ecosystem and its relationship to the effects of excessive siltation and sedimentation on the population of

Vetericaris chaceorum

and its associated species and the anchialine pool ecosystem at Lua o Palahemo. The commenter referenced the occurrence of large numbers of individuals of

Halocaridina rubra, Procaris hawaiiana,

and

V. chaceorum

during the 1985 survey (Kensley and Williams 1985, pp. 417-426) despite a reduction in visibility (few centimeters) as a result of the disturbance of ceiling sediments caused by exhalation bubbles during an exit phase of a dive. The commenter also stated that “there is no reason to discount the opposite idea that increased flushing has mobilized the sediment, allowed the movement of native predators and competitors into the system, and resulted in the decline or perhaps extirpation of

Vetericaris.”

The commenter then suggested that the thick sediment cone just below the opening was not a problem for the dense populations of native species detected directly beneath the surface of the pool during the 1985 surveys.

Our Response:

We acknowledge the peer reviewer's statement that

Vetericaris chaceorum

and other native species may be able to coexist with a certain level sedimentation in the anchialine pool ecosystem at Lua o Palahemo. However, the water clarity has declined since earlier surveys (Kensley and Williams 1986, pp. 417-437; Bozanic 2004, pp. 1-3; Wada 2010, in litt.; Wada et al. 2012, in litt.; Wada 2012, pers. comm.; Wada 2013, in litt.), which took place in the 1970s and 1980s, despite the presence of silt in the system at that time. Further, we disagree that the reduced visibility created by a diver's exhalation bubbles or similar human-initiated disturbance during those early surveys is comparable to the low visibility levels apparent in recent surveys before surveyors even enter the water. Flushing is necessary for the successful functioning of an anchialine pool ecosystem (Brock 2004, pp. 11, 35-36). We have concluded that continued excessive siltation into and additional collapse of the lava tube system at Lua o Palahemo is causing degradation of the anchialine pool ecosystem. These factors, combined with the system's diminished ability to flush, have resulted in the degradation of water quality, which has also led to the drastic decline in two of the other hypogeal shrimp species within the pool (i.e.,

Procaris hawaiiana

numbered in the thousands, and

Halocaridina

numbered in the tens of thousands (Kensley and Williams 1986, p. 418), and the most recent survey counted 7

Procaris hawaiiana

and zero

Halocaridina

(Wada

et al.

2012, in litt.; Wada 2013, pers. comm.)). These shrimp are considered food sources for

V. chaceorum,

and their decline may affect the survival of

V. chaceorum.

(14)

Comment:

One peer reviewer requested that the discussion of Lua o Palahemo clarify land ownership and the attitude of the landowner toward the anchialine pool and its fauna.

Our Response:

Lua o Palahemo is located on land owned by the State of

Hawaii Department of Hawaiian Homelands (DHHL). We hope to work with DHHL to address the threats to

Vetericaris chaceorum

and the anchialine pool ecosystem at Lua o Palahemo from ungulates, recreational vehicles, dumping of trash, the intentional introduction of nonnative fish, and sedimentation, as identified in this final rule.

(15)

Comment:

One peer reviewer suggested that additional data on phylogenetic or biogeographical relationships on the ancestor(s) to

Vetericaris chaceorum

could have very important implications about the spatial extent of potential habitat, specific features of the habitat that may be critical to the species, and other possible sites where the species may occur. However, the peer reviewer also stated that this information is not currently available.

Our Response:

We agree that such information would provide additional insights on the species' distribution and range, as well as the physical and biological habitat features required for the conservation of

Vetericaris chaceorum.

However, as the peer reviewer noted, such information is not currently available. The documented observation of

V. chaceorum

less than 19 mi (25 km) from Lua o Palahemo in the shallow water pools at Manuka, Hawaii, may be explained by Maciolek's (1983, p. 615) hypothesis that habitats may be colonized from long-existing subterranean populations.

(16)

Comment:

One peer reviewer suggested that we add nonnative plants (e.g.,

Prosopis pallida

(kiawe)) as a threat to the anchialine pool shrimp

Vetericaris chaceorum,

as any nonnative canopy or peripheral vegetation may result in changes in anchialine habitat conditions such as increased senescence, changes in water quality, and potential increases in nutrient availability that may alter primary production and the community structure of the algae. This peer reviewer further stated that these impacts may primarily affect the predominant endemic faunal species

Halocaridina rubra,

which is considered to be a key species in maintaining the ecological integrity of the anchialine pools, and that this may ultimately lead to an overall degradation of the anchialine pool ecosystem, and therefore impact

V. chaceorum.

However, this peer reviewer also noted that both Lua o Palahemo and Manuka are either very sparse or entirely free of peripheral vegetation, but that this does not preclude the possibility of

P. pallida

or any other type of nonnative vegetation from establishing itself within these areas.

Our Response:

The Act and our regulations direct us to consider the “present” or “threatened” destruction, modification, or curtailment of the species' habitat or range. At this time, there are insufficient data to determine the impacts on

Vetericaris chaceorum

from nonnative plants such as

Prosopis pallida.

Therefore, we cannot address nonnative plants as threats to

V. chaceorum

(i.e., we cannot identify a future condition that may or may not occur as a threat) in this final rule. We will consider the need to address nonnative plants in our future recovery planning efforts for this species, should new information become available indicating nonnative plants are a threat to

V. chaceorum

at Lua o Palahemo or Manuka.

(17)

Comment:

Two peer reviewers suggested that we add native marine fish species (e.g., aholehole (

Kuhlia

sp.) or papio (

Caranx

sp.)) not normally found in anchialine pools as a threat to

Vetericaris chaceorum,

from either natural events (e.g., high surf and storm surges) or deliberate introduction by people to the Lua o Palahemo anchialine pool ecosystem. According to these reviewers, the introduction of native marine fish in anchialine pools could result in the same deleterious impacts to

V. chaceorum

and its pool habitat as the intentional introduction of nonnative fish (see “Dumping of Trash and Introduction of Nonnative Fish” under

Factor E. Other Natural or Manmade Factors Affecting Their Continued Existence,

below). One peer reviewer later suggested that it was possible, although unlikely, that native marine fish would be intentionally introduced to the four pools at Manuka.

Our Response:

We agree that the introduction of native marine species, normally isolated from the anchialine pool environment, into the anchialine pool at Lua o Palahemo that supports

Vetericaris chaceorum

may be possible. For the reasons described below, we believe it is unlikely that natural events such as high surf and storm surges will introduce native marine fish to either location (Lua o Palahemo or Manuka) of

V. chaceorum,

although one peer reviewer suggested that the 2005 earthquake on Hawaii Island may have reopened or improved the connection between the ocean and Lua o Palahemo, thus allowing natural recruitment of native marine fish into and out of the pool (Kinzie 2012, in litt.). The intentional introduction of native marine fish is possible at its two known locations.

Nonnative fish have been intentionally introduced to Lua o Palahemo in the past (see “Dumping of Trash and Introduction of Nonnative Fish” under

Factor E. Other Natural or Manmade Factors Affecting Their Continued Existence,

below), and it is not unreasonable to assume that native marine fish may be deliberately introduced to the pool. In our 2012 snorkel survey of this pool, we observed a tropical marine goby in the pool (Wada

et al.

2012, in litt.). However, it is unclear how this fish gained access to the pool. The accidental introduction or natural recruitment of native marine fish due to natural events such as storm surge and high surf is unlikely at Lua o Palahemo due to its elevation above the coast (approximately 25 ft (8 m)) and its distance from the coast (490 ft (150 m)) (Kensley and Williams 1986, p. 418). Although a massive landslide or earthquake may trigger a local tsunami that generates waves that may sweep over and deposit native marine fish in the pool, these events are purely speculative.

The intentional introduction of native marine fish is possible at the Manuka pools that support

V. chaceorum

because there is evidence that at least one pool in this area harbors nonnative freshwater poeciliids (see Factors Affecting the 15 Species, below) and marine fish, likely introduced by fishermen. This pool is located near a popular coastal fishing spot. Three of the four pools that support

V. chaceorum

at Manuka are located between 10 and 33 ft (3 and 10 m) from a jeep road that provides access to coastal fishing and recreational locations frequented by the public (Sakihara 2013, in litt.). The fourth pool is approximately 60 ft (18 m) from the jeep road (Sakihara 2013, in litt.). However, the accidental introduction or natural recruitment of native marine fish, due to natural events such as storm surge and high surf, is unlikely at the four pools that support

V. chaceorum

at Manuka because these pools are located at least 98 ft (30 m) from the coast (Sakihara 2013, in litt.), and storm surge and high surf that would cover this distance is improbable. Although a massive landslide or earthquake may trigger a tsunami that generates waves that may sweep over and deposit native marine fish in the pools, these events are purely speculative.

On Maui, both aholehole and papio have been found in the larger anchialine pools closest to the ocean at Ahihi Kinau NAR, where high surf and storm waves appear to wash those and other native marine fish into the pools (Wada 2013, in litt.). However, these pools are

subject to coastal influences due to natural events such as storm surge and high surf due to their proximity to the ocean. We are unaware of any data documenting the impacts of native marine fish that may be swept into the pools at Ahihi Kinau NAR on native anchialine pool shrimp.

Native marine fish species have a purely marine (pelagic) larval stage, so a population of native fishes in an anchialine pool is likely to be individuals that are introduced to pools post larvae-stage (Sakihara 2013, in litt.). According to Brock (2004, p. 9), native marine fish are typically found in pools in close proximity to the ocean and it is believed that the biological status of these pools changes with successful colonization or mortality of marine fishes in these pools. The presence of native fish in Hawaiian anchialine pools usually signals the lack of hypogeal shrimp (Brock 2004, p. 9). Brock (2004, p. 29) also states that native marine fish are not able to complete their life cycles in anchialine pools, so the impacts to hypogeal shrimp are temporary (i.e., only as long as the fish occupy the pool) and that hypogeal shrimp may successfully hide in crevices from predatory fish and thus possibly recolonize a pool after the fish die off. Therefore, although

V. chaceorum

is a hypogeal shrimp and three species upon which it is known to feed in Lua o Palahemo are hypogeal shrimp, we are unable to determine the impact of marine fish on

V. chaceorum

at this time.

(18)

Comment:

Two peer reviewers mentioned the presence of aggressive biting isopods and an eel at Lua o Palahemo, and the possibility of the eel, specifically, as a predator of

Vetericaris chaceorum.

Our Response:

We are aware that eels have been seen periodically in other anchialine pools, including pools at Manuka NAR on Hawaii Island and Ahihi Kinau on Maui. At this time, however, there are insufficient data to determine the impacts on

Vetericaris chaceorum

from biting isopods and an unidentified eel at Lua o Palahemo. Therefore, we are unable to address these animals as threats to

V. chaceorum

in this final rule. We will consider the need to address biting isopods and eels in our future recovery planning efforts for this species, should new information become available indicating these animals are threats to

V. chaceorum.

(19)

Comment:

Two peer reviewers suggested that earthquakes and subsequent landslides and rockfalls are threats to

Vetericaris chaceorum,

due to destruction or degradation of its pool habitat. This peer reviewer believes that given a large enough earthquake, the Lua o Palahemo anchialine pool could potentially lose its connection to the ocean by boulder “chokes” that block off movement of ocean water to and from the pool, or by a complete or partial collapse of the tube itself. This peer reviewer then added that we would need an engineer to make a more definitive assessment regarding the pool's vulnerability to collapse.

Our Response:

We agree that earthquakes and subsequent landslides and rockfalls are potential threats to

Vetericaris chaceorum

and its habitat. We also agree that an engineer or other professional with the necessary skills is needed to assess the vulnerability of the lava tubes within the Lua o Palahemo anchialine pool to the threat of earthquakes. We do not have enough data to include earthquakes as a threat at this time.

(20)

Comment:

Two peer reviewers commented that our analysis of the threats to

Vetericaris chaceorum

seemed too focused on the surface of the anchialine pool rather than on the depths within Lua o Palahemo (where

V. chaceorum

is reported to occur). One of the peer reviewers questioned the relevance of threats at the opening when the species is so far below the surface, while the other peer reviewer stated that any impacts at the surface of the pool may lead to degradation of the habitat within the recesses of the lava tube by causing shifts in water quality, physical conditions, and flushing, and therefore causing shifts in biological characteristics (i.e., benthic algae and primary consumer abundance and assemblage). As such, these threats may extend beyond the immediately impacted areas at Lua o Palahemo.

Our Response:

Based on the best scientific and commercial data available, we believe

Vetericaris chaceorum

faces threats from habitat loss or degradation from sedimentation in Lua o Palahemo due to degradation of the immediate area surrounding the pool. Feral goats and cattle trample and forage on both native and nonnative plants around and near the pool opening (Magnacca 2012, in litt.; Richardson 2012, in litt.), increasing erosion resulting in sediment entering the pool (see “Habitat Destruction and Modification by Introduced Ungulates” under

Factor A. The Present or Threatened Destruction, Modification, or Curtailment of Habitat or Range,

below). In addition,

V. chaceorum

faces threats from the intentional dumping of trash (at Lua o Palahemo) and introduction of nonnative fish (at Lua o Palahemo and Manuka NAR), activities which originate at the pool openings and result in impacts to

V. chaceorum

(within the deep recesses of Lua o Palahemo and within the shallower pools at Manuka NAR) (see “Dumping of Trash and Introduction of Nonnative Fish” under

Factor E. Other Natural or Manmade Factors Affecting Their Continued Existence,

below).

(21)

Comment:

One peer reviewer commented that the proposed rule presents a good summary of potential threats to the shrimp and its habitat, and it clearly makes the point that the population at Lua o Palahemo is exceedingly small and probably declining, if not extinct.

Our Response:

We appreciate this reviewer's concurrence and have considered that the shrimp may no longer be extant at Lua o Palahemo; however, since anchialine pool shrimp are known to spend much of their time within the crevices of pools, we believe the species may still be present in the pool, but in very low numbers.

(22)

Comment:

One peer reviewer commented that they had observed items that humans dumped into Lua o Palahemo, including a bicycle, boom box, and large cement block, but that they were uncertain whether or not these items had a deleterious or observable effect on

V. chaceorum.

Our Response:

The impact of human dumping of trash into an anchialine pool is directly related to the proportion between the size of the pool and the amount and type of trash dumped. For example, a large trash bag in a small, shallow anchialine pool will negatively impact habitat quality, whereas the negative effect from same trash bag in a larger, deeper anchialine pool will not reach the same magnitude of effect. In addition, if the boom box had decaying batteries in it, contaminants such as lead, mercury and cadmium could have leached into the pool (Center for Disease Control—Agency for Toxic Substances and Disease Registry (CDC-ATSDR) 2011—

Toxic Substance Database

). In addition, there is risk from exposure to general electronic waste contaminants, which contain various hazardous materials and are harmful to the environment (e.g., polyvinyl chloride, polychlorinated biphenyls, and chromium) (CDC-ATSDR 2011—

Toxic Substance Database

). These toxins produce varying effects on biological organisms that include, but are not limited to, deoxyribose nucleic acid (DNA) damage, mucous membrane damage, cancer, and organ failure (CDC-ATSDR 2011—

Toxic Substance Database

).

(23)

Comment:

Five peer reviewers commented on the likelihood of

whether or not

Vetericaris chaceorum

has a niched habitat deep within the darkness of the lava tube at Lua o Palahemo where it was observed in 1985, or whether it has a broader habitat that extends throughout the matrix of the lava tube of Lua o Palahemo. The first of these peer reviewers commented that, due to insufficient data and the challenging conditions of assessing the particular habitat(s) of Lua o Palahemo, it would be difficult to determine whether this species would likely occur throughout Lua o Palahemo or only be limited to the area where it was originally collected from within the lava tube. The second peer reviewer commented that literature suggested that

Vetericaris chaceorum

did not have a uniform distribution throughout Lua o Palahemo when it was first observed and collected, so that would suggest that it does have a limited niche and that it is highly likely that it would be still limited to the area where it was originally collected within the lava tube. The third of these peer reviewers commented that it has been confirmed that the range of

Vetericaris chaceorum

extends beyond Lua o Palahemo, although only approximately 25 km away. Therefore, it is plausible that its distribution within Lua o Palahemo also extends beyond where it was originally collected. Furthermore, the habitat in which

Vetericaris chaceorum

was found at Manuka is considerably different than that of Lua o Palahemo, which was characterized by shallow (less than 0.5 m deep), open pools dispersed throughout barren basaltic terrain. Accordingly, its range does not seem to be limited to the deep recesses of the anchialine habitat, but may also roam freely throughout shallow exposed areas. The fourth peer reviewer commented that

Vetericaris chaceorum

likely has a wider lateral distribution in the Lua o Palahemo lava tube and that it is likely found in adjacent hypogeal habitat. The fourth peer reviewer also commented that it is unclear if

Vetericaris chaceorum

venture into the lighted, mixohaline portion of Lua o Palahemo. The fifth peer reviewer commented that there is no reason to believe that the shrimp's range did not extend, at least, to the ends of that lava tube, and possibly into other openings connecting to it. As the boundaries of Lua o Palahemo were not defined in the proposed rule, an answer to the question about “throughout Lua o Palahemo” is not clear.

Our Response:

We agree and are aware that it is difficult to know exactly where this species occurs within Lua o Palahemo, and whether or not it favors the depth at which it was observed or if it utilizes the greater part of the lava tube. The newly discovered occurrence in the shallow pools at Manuka suggests that the habitat is not limited to the area it was originally collected from deep within the lava tube at Lua o Palahemo, and that it is likely

Vetericaris chaceorum

occupies areas along the matrices of Lua o Palahemo at varying depths. Because hypogeal shrimp often spend much of their time in crevices, and it is possible that

V. chaceorum

can occur throughout the lava tube, we retain the status of extant for the population of

V. chaceorum

at this location, despite the fact that

V. chaceorum

was not observed in recent surveys. Regarding the boundaries of Lua o Palahemo, we do not currently have any data that lay out the entire matrix of the lava tube, nor are we aware that such data exist.

(24)

Comment:

Three peer reviewers commented that the threats to the habitat of Lua o Palahemo expand throughout the entire lava tube matrix. One of these three peer reviewers also said that the historical differences documented for Lua o Palahemo, primarily in water clarity and quality, and the absence of other shrimp species that were common (such as

Halocaridina

) suggests the habitat has undergone serious degradation in the last 30 to 40 years that is likely to get worse if actions are not taken.

Our Response:

We agree that the threats to the species' habitat at Lua o Palahemo are not limited to any particular area and span the scope of the entire lava tube matrix. We also agree that more surveys and monitoring efforts are needed to determine how best to recover this habitat. The Service has conducted surveys in 2010 and 2012 (Wada 2012, pers. comm.; Wada et al. 2012, in litt.), and will continue to monitor and research this habitat in the future, in addition to conservation methodologies to recover

Vetericaris chaceorum

at this site.

(25)

Comment:

One peer reviewer commented that it is unclear that the best available scientific data and methodologies currently available can determine rarity vs. human accessibility to the

Vetericaris chaceorum.

This commenter also stated that a dark-adapted organism could potentially be found anywhere within the hypogeal environment of the Hawaiian Islands, and that the Service may be drawing its listing conclusion of this species based on lack of biological knowledge. In addition, this reviewer commented that the lack of information may not enable practical management decisions.

Our Response:

We agree that it is difficult to determine the entire range that is occupied by

Vetericaris chaceorum

on Hawaii Island or elsewhere in the Hawaiian Islands. We have based our determination on the number of estimated pools throughout the Hawaiian Islands and the percentage of these pools that have been surveyed. Despite surveys throughout the islands,

Vetericaris chaceorum

has only been observed in two pool complexes on Hawaii Island: Lua o Palahemo and Manuka. In addition, the fact that these two habitats are so different informs us that

Vetericaris chaceorum

is not solely a dark-adapted organism, but that it is has a range of suitable habitat that also includes shallow pools in full sunlight. This increase in suitable habitat types, the number of surveys throughout the Hawaiian Islands, and the fact that in total only 12 shrimp (5 at Lua o Palahemo and 7 at Manuka) have ever been observed suggest that

Vetericaris chaceorum

is not occurring in high numbers. We do not currently have methodologies that afford us the opportunity to search cracks and crevices within the anchialine pool environment; however, if this type of survey technology equipment becomes available, it will certainly enhance our understanding of the population dynamics of hypogeal shrimp, including

Vetericaris chaceorum.

The Service agrees that additional information will benefit management decisions.

(26)

Comment:

Two peer reviewers commented on the connection of Lua o Palahemo to the marine environment. One of these reviewers commented that the further collapse of the lava tube and increased siltation may have the effect of decreasing the slight flow of colder water into the depth of the lava tube, and that the further collapse may actually have a beneficial effect, such as isolation from human access. The second peer reviewer commented that the lava tube may be connected to a deep water marine habitat and associated fauna.

Our Response:

Kensley and Williams (1986, p. 435) state that it is probable that neither temperature nor salinity imposes a barrier to the dispersal of hypogeal shrimp. They reported a surface temperature of 24 degrees Celsius, but they did not report the temperature at the depth they observed

Vetericaris chaceorum

(Kensley and Williams 1986, p. 418). During the surveys conducted by the Service in 2012, the temperature of the water at a depth of 7.5 m from the surface ranged from 23.8 degrees Celsius at noon to 26.4 Celsius at 4:50 a.m. (Wada

et al.

2012, in litt.). The data suggest

temperature is not currently a determining factor in the presence or absence of

Vetericaris chaceorum

at Lua o Palahemo.

The definition of an anchialine pool includes being tidally influenced due to a subterranean connection to the ocean, so we agree that the lava tube is connected to a marine habitat and fauna, although to what extent and what depth is not known at this time. The size (i.e., a smaller cracks versus a wide diameter lava tube) of the connection to the marine environment will determine to some extent the species present in a given anchialine pool; the better the connection to the sea, the more likely a pool will have marine organisms (Brock 2004, p. 9). For example, the unusual ecotypic variant of the moray eel (

Gymnothorax pictus,

puhi) is often found in pools with better connections to the sea (Brock 2004, p. 9). Regarding relationship between a further collapse of the lava tube and human access, we have no data to support or deny a benefit from limiting human access to the depths of Lua o Palahemo.

(27)

Comment:

One peer reviewer commented that since so little is known about

Vetericaris chaceorum,

most considerations of threats are conjectural, and that because no apparent observations have been made of this species in the upper reaches of Lua o Palahemo, purported threats to other anchialine species may not be a limiting factor or relevant to life in the lightless marine environment.

Our Response:

As described earlier,

Vetericaris chaceorum

was initially discovered in 1985, in complete darkness within one of the lava tubes at Lua o Palahemo, at a location 180 m (590 ft) from the opening, at a depth of 30 m (98 ft). We agree that there is still much to be learned about

V. chaceorum's

life history and biology. It was recently confirmed that the species is not confined to the dark depths of Lua o Palahemo. In addition, Sakihara (2013, in litt.) observed

V. chaceorum

feeding on other anchialine pool shrimp species. Considering the new information, threats to other anchialine pool shrimp at varying depths are directly relevant to the survival of

V. chaceorum.

If the food supply of

V. chaceorum

is declining or diminished, it will have a direct impact on the health and survival of

V. chaceorum.

Further, the threats of dumping nonnative fish and trash can directly negatively impact the ecosystem at either Lua o Palahemo or Manuka; this is confirmed by observations at other anchialine pools around the Hawaiian Islands where nonnative fish and trash have caused the degradation of pools (Brock 2004, pp. 12-15).

(28)

Comment:

One peer reviewer questioned the value of comparing

Vetericaris chaceorum

with the anchialine pool shrimp

Halocaridina rubra.

This peer reviewer commented that

Vetericaris chaceorum

is likely much more specialized and that its lack of eyes, limited swimming option, and, as far as is known, very limited distribution makes comparisons between the two species uninformative for the most part. This peer reviewer further stated that the observations on the behavior of

V. chaceorum

suggests it may prey on smaller organisms by capturing them in the basket formed by its pereiopods as it swims in the dark; if this is true, the species would require large volumes of open water. The reviewer further elaborates that Kensley and Williams (1986) note the species is a strong swimmer and apparently stays in midwater, avoiding the solid walls, consistent with the filter-basket feeding hypothesis. If true, this makes this species somewhat different from other anchialine shrimp, which are generally associated with the substratum, although Maciolek observed

H. rubra

feeding in midwater “presumably grazing only on phytoplankton.” Similarly

V. chaceorum

does not appear to be very similar to the more well-studied anchialine shrimp. Its troglobitic (more correctly stygobitic) habit, large size, possibly its specialized trophic role and potentially unique evolutionary history should make comparisons with other anchialine shrimp suspect.

Our Response:

We appreciate this reviewer's comments regarding the value of comparing

Vetericaris chaceorum

and

Halocaridina rubra.

We agree that these two shrimp are not exactly the same; however,

H. rubra

is the most well-studied anchialine pool shrimp in the Hawaiian Islands, and, therefore, we used it as a surrogate species in some examples for

V. chaceorum

in regards to the negative impacts associated with human dumping of nonnative fish and trash, in addition to recognizing it as a potential food source for

V. chaceorum.

The newly discovered population of

V. chaceorum

in the four shallow pools at Manuka has broadened our understanding of the range and habitat for this species, debunking the thoughts that this species is niched to the dark depths of Lua o Palahemo. Further, this challenges the above hypothesis that this species may require large volumes of open water. As stated in the comments above, we have much to learn about

V. chaceorum,

and we base our action in this rule on the fact that the habitat is threatened by sedimentation, recreational off-road vehicles, human dumping of nonnative fish, and human dumping of trash.

(29)

Comment:

One peer reviewer commented that poeciliids are not only introduced illegally in Hawaii, State agencies introduce mosquito fish to freshwater and anchialine habitats as mosquito control. While perhaps legal, the effects are just as detrimental. However, the peer reviewer did not think that mosquito control is a concern for a site like Lua o Palahemo.

Our Response:

We agree that mosquito control is not a concern at Lua o Palahemo, and we have no information that would indicate that State agencies are introducing nonnative fish at Manuka for mosquito control.

(30)

Comment:

The proposed rule states that reduced flushing in the pool portion of Lua o Palahemo may allow an accumulation of sediment and detritus in the pool, reducing food productivity and the ability of

Vetericaris chaceorum

to move between the pool and water table. One peer reviewer commented there is no reason to discount the opposite idea that increased flushing has mobilized the sediment, allowed the movement of native predators and competitors into the system, and resulted in the decline or perhaps extirpation of

V. chaceorum.

In support of this is the statement in the October 17, 2012, proposed rule at 77 FR 63939: “During those dives, researchers made five observations of

Vetericaris chaceorum

in total darkness at a depth of 108 ft (33 m) and 590 ft (180 m) from the opening, collecting two specimens. Kensley and Williams (1986, p. 418) noted, however, that the area surveyed directly beneath the surface of the pool contained the highest density of animals (e.g., shrimps and crustaceans).” This suggests the very thick sediment cone just below the opening was not a problem for the dense populations of native species. All this just shows that there is an exceedingly limited understanding of how the system functions, and specifically what physical, chemical, and hydrologic aspects of the system promote sustaining

V.

chaceorum and its associated species. This commenter suggested that a high level of sediment is not, per se, deleterious to the shrimp, other anchialine pool species, and, by inference, the entire pool.

Our Response:

We agree it is possible that increased flushing allowed the movement of native predators and competitors into the system, resulting in the decline or perhaps extirpation of

Vetericaris chaceorum

at Lua o

Palahemo; however, we are unaware of any data to support this hypothesis. Recent surveys by the Service and State (Wada 2012, pers. comm.; Wada

et al.

2012, in litt.) have found the degradation of habitat of Lua o Palahemo is a result of excessive siltation and sedimentation of the anchialine pool system, combined with the diminished ability of the system to flush, which Brock (2004, pp. 11, 35-36) described as necessary for a functioning anchialine pool system. Long-term sedimentation accumulation leads to the senescence of anchialine pools (Ramsey 2013, in litt.). Suspended sediment within the water column of Lua o Palahemo likely reduces the capacity of the pool to produce adequate cyanobacteria and algae to support some of the pool's herbivorous hypogeal species. A decreased food supply (i.e., a reduction in cyanobacteria and algae) would likely lead to a lower abundance of herbivorous hypogeal shrimp species, as well as a lower abundance of the known carnivorous species (i.e.,

Vetericaris chaceorum

). Because lower numbers of the herbivorous hypogeal shrimp have been observed over time, the data indicate this is a contributing to, but not necessarily the sole factor in, the lack of detection of

Vetericaris chaceorum

at Lua o Palahemo.

(31)

Comment:

One peer reviewer commented that Lua o Palahemo should not be treated as a typical anchialine pool. Rather it is a singular system, or perhaps somewhat like Lake Kauhako. Extrapolating from the little we know about typical anchialine systems will probably not be productive.

Our Response:

Anchialine pools are land-locked bodies of water that have indirect underground connections to the sea, contain varying levels of salinity, and show tidal fluctuations in water level. Lua o Palahemo meets this definition. Further, Lua o Palahemo has floral and faunal characteristics of an anchialine pool ecosystem (see

Hawaii Island Ecosystems

and Description of the 15 Species, above). Lake Kauhako is situated in the crater of an extinct, late Pleistocene volcano on the north shore of Molokai, Hawaii, and reportedly not tidally influenced, although early data suggested it may have been at one time and anchialine pool shrimp were observed here in 1982 (Maciolek 1982, p. 12; Donachie

et al.

1999, p. 93). Lake Kauhako is considered one of the deepest lakes in the United States with a depth of 814 ft (248 m) (Donachie

et al.

1999, p. 93). Lake Kauhako is also meromictic (has layers of water that do not intermix) and anoxic (lacking dissolved oxygen) below 6 ft (2 m); Lua o Palahemo has not been classified as meromictic and is not noted as anoxic until a depth of 98 ft (30 m) and a distance of 180 m into one of the branches of the lava tube from the base of the surface opening (Kensley and Williams 1986, pp. 417-20). Both Lake Kauhako and Lua o Palahemo do have comparable surface dissolved oxygen and salinity and temperature gradients; however, the shape and depth of each water body, in addition to the presence or absence of tidal influence and meromictic properties, provide some distinction for these two bodies of water.

(32)

Comment:

One peer reviewer commented that the reproductive mode of

Vetericaris chaceorum

would play an important role in determining if populations could recolonize neighboring habitats after a local extirpation. Maciolek postulates that these habitats are colonized from long-existing subterranean populations, and Kensley and Williams (1986) state: “Given the relative youth of the Lua o Palahemo lava tube, the above-mentioned and unexplained absences and occurrences, and the presence of some of these shrimps in modern wells and quarries, Maciolek's postulate (1983: 615) that these habitats are colonized from long-existing subterranean populations, must be strengthened.” If this is true, the main habitat of

V. chaceorum

may be completely different from what we know about Lua o Palahemo.

Our Response:

We agree it would be beneficial to know the reproductive mode for

Vetericaris chaceorum;

however, the complete life history for this species is not known at this time. Hypogeal shrimp by definition occupy subterranean habitat. The fact that

V. chaceorum

is described as a primitive species, combined with the depth within Lua o Palahemo in which

V. chaceorum

was observed and the recent discovery of

V. chaceorum

in very different habitat at Manuka, together appear to support Maciolek's hypothesis that hypogeal shrimp colonized anchialine pool habitats from long-existing subterranean populations, but this is only conjecture at this time. The newly discovered population at Manuka supports the thought that the main habitat of

V. chaceorum

at Lua o Palahemo is likely different from what we previously thought.

Comments From the State of Hawaii

(33)

Comment:

The Hawaii Department of Business, Economic Development, and Tourism's Hawaii Housing Finance and Development Corporation challenged our proposal to list

Bidens micrantha

ssp.

ctenophylla

as an endangered species, stating that the lowland dry ecosystem covers a very large area on Hawaii Island and that the Service did not have enough studies regarding the absence or abundance of this species within this ecosystem. According to this agency, without knowing the absence or prevalence of this species, it cannot be determined whether or not this species should be designated as endangered, and the Service's findings are premature with no foundation.

Our Response:

We disagree that there is a lack of information regarding the presence or abundance of

Bidens micrantha

ssp.

ctenophylla

in the lowland dry ecosystem on the island of Hawaii and that our determination to list this species as an endangered species is premature and without foundation. Lowland dry ecosystems in the Hawaiian Islands have undergone sweeping changes over the last 100 years due to development, agriculture, and nonnative plants and animals that have resulted in the loss of over 90 percent of Hawaii's dry forests (Bruegmann 1996, pp. 26-27; Cabin

et al.

2000, pp. 439-453; Sakai

et al.

2002, pp. 276-302; Cordell

et al.

2008, pp. 279-284); however, the actual extent of native dry forest cover may be as low as 1 percent (Pau 2011, in litt.). Forty-five percent of Hawaii's dry forest plant species are at risk of endangerment (Pau

et al.

2009, p. 3,167). Twenty-five percent of the endangered plant species in the Hawaiian Islands are dry forest species, and approximately 20 percent of Hawaii's dry land plant species are believed to be extinct (Cabin

et al.

2000, pp. 439-453; Sakai

et al.

2002, pp. 276-302). One of the last remaining areas of lowland dry forest in the Hawaiian Islands is in the north Kona region of Hawaii Island, where only patches or scattered individuals of native plants remain amidst a sea of the highly flammable, nonnative fountain grass (

Pennisetum setaceum

), where over 200,000 ac (80,939 ha) of land are covered with fountain grass (HISC 2013, in litt.). North Kona is also a rapidly growing, urban area with a steady flow of new housing, roads, commercial, and industrial developments. Surveys and observations conducted over the last 90 years have detected

Bidens micrantha

ssp.

ctenophylla

from only six locations, totaling fewer than 1,000 individuals in north Kona (see Description of the 15 Species, above) (Sherff 1920, p. 97; Degener and Wiebke 1926, in litt.; Scottsberg 1926, in litt.; Borges and Degener 1929, in litt.; Degener and Iwasaki 1930, in litt.; Nishina 1931, in litt.; Krajina 1961, in litt.; Gillett 1965,

in litt.; Nagata and Ganders 1983, pp. 1-16; Pratt and Abbott 1996, p. 26; Ganders and Nagata 1999, pp. 271, 273; TNC 2007-

Ecosystem Database of ArcMap Shapefiles,

unpublished; Whistler 2007, pp. 1-18; Bio 2008, in litt.; Whistler 2008, pp. 1-11; Hawaii Forest Institute 2009, in litt.; Beavers 2010, in litt.; Faucette 2010, pp. 1-27; HBMP 2010b; Giffin 2011, pers. comm.; Pau 2011, in litt.; Wagner 2011, in litt.; Zimpfer 2011, in litt.; Kaahahui O Ka Nahelehele 2013, in litt.).

Under the Act, we determine whether a species is an endangered species or a threatened species because of any of five factors (see Summary of Factors Affecting the 15 Species, below), and we are required to make listing determinations solely on the basis of the best available scientific and commercial data

available

[emphasis ours] (sections 4(a)(1) and 4(b)(1)(A)). The threats to

B. micrantha

ssp.

ctenophylla,

as well as those that impact lowland dry ecosystems in the Hawaiian Islands, are well documented. This plant species faces threats from habitat degradation from development and nonnative ungulates (feral pigs and goats), predation by nonnative ungulates (feral pigs and goats) and rats, competition with nonnative plants, fire, drought, hurricanes, and hybridization; it also faces threats from the synergistic effects that may arise from any combination of these threats (see Summary of Factors Affecting the 15 Species, below). Therefore, in this final rule, we have made our determination to list

Bidens micrantha

ssp.

ctenophylla

as an endangered species based on the best scientific and commercial data available.

Comments From Federal Agencies

All of the comments we received from Federal agencies have been incorporated, as appropriate, in the Description of the 15 Species, above, and Summary of Changes from Proposed Rule, below.

Public Comments on the Proposed Listing of 15 Species

(34)

Comment:

One commenter, representing Laiopua 2020, stated that none of the 15 species proposed for listing occurs on parcels proposed for development of the Laiopua Community Center (Tax Map Key parcels 3-7-4-021:002, 003, and 023). The commenter provided a 2008 botanical survey report (Gerrish and Leonard Bisel Associates, LLC, 2008, entire) to confirm the absence of the 15 species on the three parcels.

Our Response:

We appreciate the information provided by the commenter and have taken it into consideration in this final listing determination. The botanical survey published by Gerrish and Leonard Bisel Associates, LLC, in 2008 was one of multiple surveys and botanical expert reports used by the Service to determine the range of

Bidens micrantha

ssp.

ctenophylla

in North Kona. Since

Bidens micrantha

ssp.

ctenophylla

is known to occur in the area of Laiopua, the Service considered this area as habitat for this species. In addition, there is likely a seed bank in the soil of the surrounding area that, if given the opportunity, can contribute toward the recovery of this species.

Summary of Changes From Proposed Rule

In preparing this final rule, we reviewed and fully considered comments from the peer reviewers and public on the proposed listing for 15 species. This final rule incorporates the following substantive changes to our proposed listing, based on the comments we received:

(1) We added inundation by high surf as a threat to the newly listed plant

Bidens hillebrandiana

ssp.

hillebrandiana

in the following locations in this final rule: Table 3 (below) and “Habitat Destruction and Modification Due to Rockfalls, Treefalls, Landslides, Heavy Rain, Inundation by High Surf, Erosion, and Drought” under

Factor A. The Present or Threatened Destruction, Modification, or Curtailment of Habitat or Range

(below), based on a peer review comment.

(2) We added the nonnative understory plant species

Sphagneticola trilobata

[

Wedelia trilobata

] (wedelia) as a threat to the plant

Bidens hillebrandiana

ssp.

hillebrandiana

in the coastal and dry cliff ecosystem, and to “Specific Nonnative Plant Species Impacts” (below), based on a peer review comment.

(3) We added the nonnative vine

Paederia foetida

(skunk weed) as a threat to the newly listed plant

Cyrtandra nanawaleensis

in the lowland wet ecosystem and to “Specific Nonnative Plant Species Impacts” (below), based on a peer review comment.

(4) We added the nonnative canopy plant species

Psidium cattleianum

(strawberry guava) as a threat to

Cyanea tritomantha

in the wet cliff ecosystem, based on a peer review comment that we include this nonnative plant species as a threat to this species in its known locations, in this final rule.

(5) We added

Pisonia

spp. as a host plant for the picture-wing fly

Drosophila digressa,

in the following locations in this final rule: Description of the 15 Species (above); “Habitat Destruction and Modification by Introduced Ungulates” and “Habitat Destruction and Modification Due to Rockfalls, Treefalls, Landslides, Heavy Rain, Inundation by High Surf, Erosion, and Drought” under

Factor A. The Present or Threatened Destruction, Modification, or Curtailment of Habitat or Range

(below); “Predation and Herbivory” under

Factor C. Disease or Predation

(below); and “Loss of Host Plants” under

Factor E. Other Natural or Manmade Factors Affecting Their Continued Existence

(below), based on a peer review comment.

(6) Hawaii State biologists discovered a population of

Vetericaris chaceorum

at Manuka NAR between 2009 and 2010. We solicited public comments on the new location in the

Federal Register

in our April 30, 2013, document announcing the availability of the draft economic analysis and reopening the comment period on the proposed rule (78 FR 25243). The new location information has been incorporated in the following sections in this final rule: Description of the 15 Species (above), “Habitat Destruction and Modification by Sedimentation” under

Factor A. The Present or Threatened Destruction, Modification, or Curtailment of Habitat or Range

(below), and “Dumping of Trash and Introduction of Nonnative Fish” (below) under

Factor E. Other Natural or Manmade Factors Affecting Their Continued Existence,

and we reassessed whether listing was warranted for

V. chaceorum

based on this additional information.

(7) We revised the statement that incorrectly indicated that the outplanted individuals of

Bidens micrantha

ssp.

ctenophylla

within KHNHP are fenced in Description of the 15 Species, above, based on a comment we received.

Summary of Factors Affecting the 15 Species

Section 4 of the Act (16 U.S.C. 1533) and its implementing regulations (50 CFR part 424) set forth the procedures for adding species to the Federal Lists of Endangered and Threatened Wildlife and Plants. 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: (A) The present or threatened destruction, modification, or curtailment of its habitat or range; (B) overutilization for commercial, recreational, scientific, or educational purposes; (C) disease or predation; (D) the inadequacy of existing regulatory mechanisms; and (E) other natural or manmade factors affecting its continued

existence. Listing actions may be warranted based on any of the above threat factors, singly or in combination.

If we determine that the level of threat posed to a species by one or more of the five listing factors is such that the species meets the definition of either endangered or threatened under section 3 of the Act, that species may then be listed as endangered or threatened. The Act defines an endangered species as “in danger of extinction throughout all or a significant portion of its range,” and a threatened species as “likely to become an endangered species within the foreseeable future throughout all or a significant portion of its range.” The threats to each of the individual 15 species are summarized in Table 3, and discussed in detail below.

Table 3-Summary of Primary Threats Identified for Each of the 15 Hawaii Island Species

Species

Ecosystem

Factor A

Agriculture and urban development

Ungulates

Non

native

plants

Fire

Stochastic events

Climate change

Factor B

Over-

utilization

Factor C

Disease

Predation/

herbivory by

ungulates

Predation/

herbivory by

other NN

vertebrates

Predation/

herbivory

by NN

invertebrates

Factor D

Inadequate

existing

regulatory

mechanisms

Factor E

Other

species-

specific

threats

Plants:

Bidens hillebrandiana

ssp.

hillebrandiana

CO, DC

P, G

X

H, RF, L, HS, E

Pt

P, G

R

X

LN

Bidens micrantha

ssp.

ctenophylla

LD

X

P, G

X

X

H, DR

Pt

P, G

R

X

HY

Cyanea marksii

LW, MW

P, C, M

X

H, RF, L

Pt

P, C, M

R

S

X

LN

Cyanea tritomantha

LW, MW, WC

P, C

X

H, TF

Pt

P, C

R

S

X

NR

Cyrtandra nanawaleensis

LW

P

X

H

Pt

P

R

S

X

HY

Cyrtandra wagneri

LW

P

X

H, HR, E

Pt

P

R

S

X

LN, HY

Phyllostegia floribunda

LW, MM, MW

P

X

X

H

Pt

P

X

Pittosporum hawaiiense

LM, MM, MW

P, C, M

X

H

Pt

P, C, M

R

X

NR

Platydesma remyi

LW, MW

P

X

H

Pt

P

X

LN, NR

Pritchardia lanigera

LM, LW, MW, WC

P, G, M, C

X

H

Pt

X

P, G, M

R

LH, B

X

NR

Schiedea diffusa

ssp.

macraei

MW

P, C

X

H

Pt

P, C

R

X

LN

Schiedea hawaiiensis

MD

P, G, SH, M

X

X

H, DR

Pt

P, G, SH, M

R

X

LN

Stenogyne cranwelliae

MW, WC

P

X

H

Pt

P

R

S

X

Animals

Drosophila digressa

(Picture-wing fly)

LM, MM, MW

P, G, C, M

X

X

H, DR

Pt

W, A

X

LN, LOH, F

Vetericaris chaceorum

(Anchialine pool shrimp)

AP

G, C

Pt

X

REC, SD, D

Factor A = Habitat Modification

LW = Lowland Wet

SH = Sheep

DR = Drought

HY = Hybridization

Factor B = Overutilization

MD = Montane Dry

M = Mouflon

RF = Rockfalls

NR = No Regeneration

Factor C = Disease or Predation

MM = Montane Mesic

R = Rats

L = Landslides

F = Flies

Factor D = Inadequacy of Regulatory Mechanisms

MW = Montane Wet

S = Slugs

HR = Heavy Rain

LOH = Loss of Host

Factor E = Other Species-Specific Threats

DC = Dry Cliff

W = Wasps

HS = High Surf

REC = Recreational vehicles

AP = Anchialine Pools

WC = Wet Cliff

A = Ants

E = Erosion

SD = Sedimentation

CO = Coastal

P = Pigs

LH = Leafhopper

TF = Tree Fall

Pt = Potential

LD = Lowland Dry

G = Goats

B = Beetles

D = Dumping (i.e., Human dumping of nonnative fish and trash)

X = Threat

LM = Lowland Mesic

C = Cattle

H = Hurricane

LN = Limited Numbers

Blank = Not a Threat

The following constitutes a list of ecosystem-scale threats that affect the species in this final rule in one or more of the 10 described ecosystems on Hawaii Island:

(1) Foraging and trampling of native plants by feral pigs (

Sus scrofa

), goats (

Capra hircus

), cattle (

Bos taurus

), sheep (

Ovis aries

), or mouflon sheep (

Ovis gmelini musimon

), which results in severe erosion of watersheds because these mammals inhabit terrain that is often steep and remote (Cuddihy and Stone 1990, p. 63). Foraging and trampling events destabilize soils that support native plant communities, bury or damage native plants, and have adverse water quality effects due to runoff over exposed soils.

(2) Ungulate destruction of seeds and seedlings of native plant species via foraging and trampling (Cuddihy and Stone 1990, pp. 63, 65) facilitates the conversion of disturbed areas from native to nonnative vegetative communities.

(3) Disturbance of soils by feral pigs from rooting can create fertile seedbeds for alien plants (Cuddihy and Stone 1990, p. 65), some of them spread by ingestion and excretion by pigs.

(4) Increased nutrient availability as a result of pigs rooting in nitrogen-poor soils, which facilitates establishment of alien weeds. Introduced vertebrates are known to enhance the germination of alien plants through seed scarification in digestive tracts or through rooting and fertilization with feces of potential seedbeds (Stone 1985, p, 253). In addition, alien weeds are more adapted to nutrient-rich soils than native plants (Cuddihy and Stone 1990, p. 65), and rooting activity creates open areas in forests allowing alien species to completely replace native stands.

(5) Rodent damage to plant propagules, seedlings, or native trees, which changes forest composition and structure (Cuddihy and Stone 1990, p. 67).

(6) Feeding or defoliation of native plants from alien insects, which reduces geographic ranges of some species because of damage (Cuddihy and Stone 1990, p. 71).

(7) Alien insect predation on native insects, which affects pollination of native plant species (Cuddihy and Stone 1990, p. 71).

(8) Significant changes in nutrient cycling processes because of large numbers of alien invertebrates, such as earthworms, ants, slugs, isopods, millipedes, and snails, resulting in changes to the composition and structure of plant communities (Cuddihy and Stone 1990, p. 73).

Each of the above threats is discussed in more detail below, and summarized in Table 3.

Factor A. The Present or Threatened Destruction, Modification, or Curtailment of Habitat or Range

The Hawaiian Islands are located over 2,000 mi (3,200 km) from the nearest continent. This isolation has allowed the few plants and animals that arrived in the Hawaiian Islands to evolve into many highly varied and endemic species (species that occur nowhere else in the world). The only native terrestrial mammals in the Hawaiian Islands are two bat taxa, the extant Hawaiian hoary bat (

Lasiurus cinereus semotus

) and an extinct, unnamed, insectivorous bat (Ziegler 2002, p. 245). The native plants of the Hawaiian Islands, therefore, evolved in the absence of mammalian predators, browsers, or grazers. As a result, many of the native species have lost unneeded defenses against threats such as mammalian predation and competition with aggressive, weedy plant species that are typical of continental environments (Loope 1992, p. 11; Gagne and Cuddihy 1999, p. 45; Wagner

et al.

1999d, pp. 3-6). For example, Carlquist (in Carlquist and Cole 1974, p. 29) notes that “Hawaiian plants are notably free from many characteristics thought to be deterrents to herbivores (toxins, oils, resins, stinging hairs, coarse texture).”

Native Hawaiian plants are therefore highly vulnerable to the impacts of introduced mammals and alien plants. In addition, species restricted and adapted to highly specialized locations (e.g.,

Bidens hillebrandiana

ssp.

hillebrandiana

) are particularly vulnerable to changes (e.g., nonnative species, hurricanes, fire, and climate change) in their habitat (Carlquist and Cole 1974, pp. 28-29; Loope 1992, pp. 3-6; Stone 1992, pp. 88-102).

Habitat Destruction and Modification by Agriculture and Urban Development

The consequences of past land use practices, such as agricultural or urban development, have resulted in little or no native vegetation below 2,000 ft (600 m) throughout the Hawaiian Islands (TNC 2007-

Ecosystem Database of ArcMap Shapefiles,

unpublished), largely impacting the coastal, lowland dry, lowland mesic, and lowland wet ecosystems. Although agriculture has been declining in importance, large tracts of former agricultural lands are being converted into residential areas or left fallow (TNC 2007-

Ecosystem Database of ArcMap Shapefiles,

unpublished). In addition, Hawaii's population has increased almost 7 percent in the past 10 years, further increasing demands on limited land and water resources in the islands (Hawaii Department of Business, Economic Development, and Tourism (HDBEDT) 2010).

Development and urbanization of the lowland dry ecosystem on Hawaii Island is a threat to one species in this rule,

Bidens micrantha

ssp.

ctenophylla. Bidens micrantha

ssp.

ctenophylla

is currently found in an area less than 10 sq mi (26 sq km) on the leeward slopes of Hualalai volcano in the lowland dry ecosystem. This area encompasses the increasingly urbanized region of north Kona, where there is very little undisturbed habitat (Pratt and Abbott 1997, p. 25). Approximately 25 percent (119 individuals of 475) of the largest of the 6 occurrences of this species is in the right-of-way of the Ane Keohokalole Highway Project (USFWS 2010, in litt.) and Kaloko Makai Development, although 154 ac (62 ha) will be set aside as a lowland dry forest preserve (Kaloko Makai Dryland Forest Preserve) to compensate for the loss of these individuals as a result of highway construction and prior to the Kaloko Makai Development. Individuals of

Bidens micrantha

ssp.

ctenophylla

also occur in areas where the development of the Villages of Laiopua at Kealakehe and of the Keahuolu affordable housing project (Whistler 2007, pp. 1-18; DHHL 2009, p. 15) is a threat to the species.

Habitat Destruction and Modification by Introduced Ungulates

Introduced mammals have greatly impacted the native vegetation, as well as the native fauna, of the Hawaiian Islands. The presence of introduced alien mammals is considered one of the primary factors underlying the alteration and degradation of native plant communities and habitats on the island of Hawaii. The destruction or degradation of habitat due to nonnative ungulates (hoofed mammals), including pigs, goats, cattle, sheep, and mouflon, is currently a threat to the 10 ecosystems (lowland dry, lowland mesic, lowland wet, montane dry, montane mesic, montane wet, coastal, anchialine pool, dry cliff, and wet cliff) on Hawaii Island and their associated species. Habitat degradation or destruction by ungulates is also a threat to all 13 plant species and the picture-wing fly in this final rule (Table 3). Habitat degradation or destruction by ungulates is a threat to the anchialine pool shrimp at Lua o Palahemo, but is not reported to pose a threat to the four pools that support this species at Manuka.

The destruction or degradation of habitat due to pigs is currently a threat to nine of the Hawaii Island ecosystems (coastal, lowland dry, lowland mesic, lowland wet, montane dry, montane mesic, montane wet, dry cliff, and wet cliff) and their associated species. In Hawaii, pigs have been described as the most pervasive and disruptive nonnative influence on the unique native forests of the Hawaiian Islands, and are widely recognized as one of the greatest current threats to forest ecosystems (Aplet

et al.

1991, p. 56; Anderson and Stone 1993, p. 195).

These feral animals are extremely destructive and have both direct and indirect impacts on native plant communities. While rooting in the earth in search of invertebrates and plant material, pigs directly impact native plants by disturbing and destroying vegetative cover, and by trampling plants and seedlings. It has been estimated that at a conservative rooting rate of 2 sq yards (yd) (1.7 sq m) per minute, with only 4 hours of foraging a day, a single pig could disturb over 1,600 sq yd (1,340 sq m) (or approximately 0.3 ac, or 0.12 ha) of groundcover per week (Anderson

et al.

2007, p. 2).

Pigs reduce or eliminate plant regeneration by damaging or eating seeds and seedlings (further discussion of predation by nonnative ungulates is provided under

Factor C. Disease or Predation,

below). Pigs are a major vector for the establishment and spread of competing invasive, nonnative plant species by dispersing plant seeds on their hooves and fur, and in their feces (Diong 1982, pp. 169-170), which also serves to fertilize disturbed soil (Matson 1990, p. 245; Siemann

et al.

2009, p. 547). Pigs feed on the fruits of many nonnative plants, such as

Passiflora tarminiana

(banana poke) and

Psidium cattleianum

(strawberry guava), spreading the seeds of these invasive species through their feces as they travel in search of food. Pigs also feed on native plants, such as Hawaiian tree ferns that they root up to eat the core of the trunk (Baker 1975, p. 79). In addition, rooting pigs contribute 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; Medeiros

et al.

1986, pp. 27-28; Scott

et al.

1986, pp. 360-361; Tomich 1986, pp. 120-126; Cuddihy and Stone 1990, pp. 64-65; Aplet

et al.

1991, p. 56; Loope

et al.

1991, pp. 1-21; Gagne and Cuddihy 1999, p. 52; Nogueira-Filho

et al.

2009, pp. 3,677-3,682; Dunkell

et al.

2011, pp. 175-177). Erosion impacts native plant communities by watershed degradation and alteration of plant nutrient status due to associated outcomes such as sediment build up in waterways and top soil run off, respectively, as well as damage to individual plants from landslides (Vitousek

et al.

2009, pp. 3074-3086; Chan-Halbrendt

et al.

2010, p. 252).

Pigs have been cited as one of the greatest threats to the public and private lands within the Olaa Kilauea Partnership (an area of land that includes approximately 32,000 ac (12,950 ha) in the upper sections of the Olaa and Waiakea forests above Volcano village) that comprise the lowland mesic, lowland wet, montane mesic, and montane wet ecosystems that support individuals of three of the plant species in this final rule (

Cyanea tritomantha, Phyllostegia floribunda,

and

Pittosporum hawaiiense

) (Olaa Kilauea Partnership Area Feral Animal Monitoring Report 2005, pp. 1-4; Perlman 2007, in litt.; Pratt 2007a, in litt.; Pratt 2007b, in litt.; Benitez

et al.

2008, p. 58; HBMP 2010f; HBMP 2010h; PEPP 2010, p. 60, TNC 2012, in litt.). Impacts from feral pigs are also a threat to the coastal, lowland mesic, lowland wet, montane wet, dry cliff, and wet cliff ecosystems in the northern Kohala Mountains and adjacent coastline. These ecosystems support occurrences of seven of the plant species in this final rule (

Bidens hillebrandiana

ssp.

hillebrandiana, Cyanea tritomantha,

Cyrtandra wagneri, Platydesma remyi,

Pritchardia lanigera, Schiedea diffusa

ssp.

macraei,

and

Stenogyne cranwelliae

) (Wood 1995, in litt.; Wood 1998, in litt.; Perlman

et al.

2001, in litt.; Wagner

et al.

2005d, pp. 31-33; Kohala Mountain Watershed Partnership (KMWP) 2007, pp. 54-56; Lorence and Perlman 2007, pp. 357-361; HBMP 2010a; HBMP 2010c; HBMP 2010f; HBMP 2010i; HBMP 2010j; HBMP 2010k; PEPP 2010, pp. 63, 101, 106; Bio 2011, pers. comm.). In addition, feral pigs are a threat to the lowland wet and montane wet ecosystems in south Kona, Kau, and Puna districts that support the plants

Cyanea marksii, Cyrtandra nanawaleensis,

and

Pritchardia lanigera

(Bio 2011, pers. comm.; Magnacca 2011b, pers. comm.; Maui Forest Bird Recovery Project 2011, in litt.; Crysdale 2013, pers. comm.). Feral pigs have also been reported in the lowland dry ecosystem that supports the plant

Bidens micrantha

ssp.

ctenophylla

(Bio 2011, pers. comm.) and the montane dry ecosystem that supports habitat for the only known occurrence of the plant

Schiedea hawaiiensis

(Mitchell

et al.

2005c; U.S. Army Garrison 2006, pp. 27, 34, 95-97, 100-107, 112). Although we do not have direct evidence of feral pigs threatening the particular species on Hawaii Island that are in this final rule, those threats have been documented on other islands where pigs have been introduced (Mitchell

et al.

2005c; U.S. Army Garrison 2006, pp. 27, 34, 95-97, 100-107, 112). We find it is reasonable to infer that feral pig threats to these species that have been observed on other Hawaiian islands would act in a similar manner on Hawaii Island, where those species interact.

Many of the most important host plants of Hawaiian picture-wing flies (

Charpentiera, Pisonia,

Pleomele, Reynoldsia, Tetraplasandra,

Urera,

and the lobelioids (e.g.,

Cyanea

spp.)) are also among the most susceptible to damage from feral ungulates, such as pigs (Foote and Carson 1995, p. 370; Kaneshiro and Kaneshiro 1995, pp. 8, 39; Magnacca

et al.

2008, p. 32; Magnacca 2013, in litt.). Feral pig browsing alters the essential microclimate in picture-wing fly (

Drosophila digressa

) habitat by opening up the canopy, leading to increased desiccation of soil and host plants (

Charpentiera

spp. and

Pisonia

ssp.), which disrupts the host plants' life cycle and decay processes, resulting in disruption of the picture-wing fly's life cycle, particularly oviposition and larvae substrate (Magnacca

et al.

2008, pp. 1, 32). Foote and Carson (1995, p. 369) have experimentally demonstrated the above detrimental effects of feral pigs on

Drosophila

spp. in wet forest habitat on the island of Hawaii. In addition, Montgomery (2005, in litt.; 2007, in litt.) and Foote (2005, pers. comm.) have observed feral pig damage to host plants (e.g.,

Charpentiera

sp.,

Cheirodendron

sp.,

Pleomele

sp.,

Tetraplasandra

sp.,

Urera kaalae

) of Hawaiian picture-wing flies on the island of Hawaii (Foote 2005, pers. comm.) and throughout the main Hawaiian Islands (Montgomery 2005, in litt.; 2007, in litt.). Magnacca (2012, pers. comm.) has observed the lack of regeneration of picture-wing fly host plants due to destruction of seedlings caused by pig rooting and herbivory.

The destruction or degradation of habitat due to goats is currently a threat to all 10 of the described ecosystems on Hawaii Island (anchialine pool, coastal, lowland dry, lowland mesic, lowland wet, montane dry, montane mesic, montane wet, dry cliff, and wet cliff) and their associated species. Goats occupy a wide variety of habitats on Hawaii Island, where they consume native vegetation, trample roots and seedlings, accelerate erosion, and promote the invasion of alien plants

(van Riper and van Riper 1982, pp. 34-35; Stone 1985, p. 261; Kessler 2011, pers. comm.). Goats are able to access, and forage in, extremely rugged terrain, and they have a high reproductive capacity (Clarke and Cuddihy 1980, pp. C-19, C-20; Culliney 1988, p. 336; Cuddihy and Stone 1990, p. 64). Because of these factors, goats have completely eliminated some plant species from islands (Atkinson and Atkinson 2000, p. 21).

Goats are be highly destructive to native vegetation, and contribute to erosion by eating young trees and young shoots of plants before they can become established, creating trails that damage native vegetative cover, promoting erosion by destabilizing substrate and creating gullies that convey water, and dislodging stones from ledges that can cause rockfalls and landslides and damage vegetation below (Cuddihy and Stone 1990, pp. 63-64). A recent study by Chynoweth

et al.

(2011, in litt.), which deployed GPS (global positioning system) satellite collars on 12 feral goats to track movement patterns every 2 hours for 1 year in Pohakuloa Training Area, found that goats prefer native-dominated shrublands in the montane dry ecosystem during the day and barren lava at night. Pohakuloa Training Area supports one of the few montane dry forest ecosystems on Hawaii Island that supports native plants in the montane dry ecosystem, including the only occurrence of the plant

Schiedea hawaiiensis

(U.S. Army Garrison 2006, pp. 27, 34; Evans 2011, in litt.). In addition, one of the two occurrences of the plant

Pritchardia lanigera

is known from an unfenced area of the Kohala Mountains, where herds of wild goats and other ungulates occur (Maly and Maly 2004

in

KMWP 2007, p. 55; KMWP 2007, pp. 54-55; Warshauer

et al.

2009, pp. 10, 24; Laws

et al.

2010, in litt.; Ikagawa 2011, in litt.). Maly and Maly (2004

in

KMWP 2007, p. 55) report that “herds of wild goats roam throughout this region, trampling, grubbing, and rending, grinding the bark of old trees and eat the young ones . . . which will destroy the beauty and alter the climate of the mountainous region of Hawaii.” There are direct observations that goats are also altering the coastal ecosystem along the Kohala Mountains, the location of the only known wild individuals of the plant

Bidens hillebrandiana

ssp.

hillebrandiana

(Warshauer

et al.

2009, p. 24; Bio 2011, pers. comm.). Goats are also found in North Kona and have been observed browsing in the lowland dry ecosystem that supports the plant

B. micrantha

ssp.

ctenophylla

(Bio 2011, pers. comm.; Knoche 2011, in litt.). Fresh seedlings from native plants attract goats to the dry and rough lava (Bio 2011, pers. comm.). Further, the host plants (

Charpentiera

spp. and

Pisonia

spp.) of the picture-wing fly in this final rule appear to be decreasing throughout their ranges due to impacts from browsing goats (Foote and Carson 1995, p. 369; Science Panel 2005, pp. 1-23; Magnacca

et al.

2008, p. 32; Magnacca 2013, in litt.). Feral goat browsing alters the picture-wing fly's (

Drosophila digressa

) essential microclimate by opening up the canopy, leading to increased desiccation of soil and host plants, which disrupts the host plants' life cycle and decay processes, resulting in the disruption of the picture-wing fly's life cycle, particularly oviposition and larvae substrate (Magnacca

et al.

2008, pp. 1, 32). Based on observations of goats and their scat (Magnacca 2012, pers. comm.) within the Ka Lae region where the Lua o Palahemo anchialine pool is located, the Service concludes that goats contribute to the degradation of the anchialine pool habitat and, thus, are a threat to the anchialine pool shrimp

Vetericaris chaceorum.

Feral goats trample and forage on both native and nonnative plants around and near the pool opening at Lua o Palahemo, and increase erosion around the pool and sediment entering the pool.

The destruction or degradation of habitat due to cattle is currently a threat to five of the described ecosystems (anchialine pool, lowland mesic, lowland wet, montane mesic, and montane wet) on Hawaii Island and their associated species. Feral cattle eat native vegetation, trample roots and seedlings, cause erosion, create disturbed areas into which alien plants invade, and spread seeds of alien plants in their feces and on their bodies. The forest in areas grazed by cattle degrades to grassland pasture, and plant cover is reduced for many years following removal of cattle from an area. In addition, several alien grasses and legumes purposely introduced for cattle forage have become noxious weeds (Tomich 1986, pp. 140-150; Cuddihy and Stone 1990, p. 29).

The wet forests of Kohala Mountain are reported to have a feral cattle population of at least 100 individuals that are causing forest degradation by trampling and browsing, which leads to subsequent increased nitrogen availability through deposition of feces (Stone 1985, p. 253), all of which contribute to the influx of nonnative plant and animal species (KMWP 2007, pp. 54-55; Laws 2010, in litt.). Feral cattle are reported from remote regions on Hawaii Island, including the back of both Pololu and Waipio Valleys (KMWP 2007, p. 55). Feral cattle are a threat to the lowland wet and montane wet ecosystems on Kohala Mountain where individuals of

Cyanea tritomantha, Pittosporum hawaiiense,

and

Pritchardia lanigera,

and the last wild individual of

Schiedea diffusa

ssp.

macraei,

are reported (PEPP 2010, pp. 59-60; Bio 2011, pers. comm.). According to a 2010 Service report (USFWS 2010, pp. 3-15, 4-86), a herd of 200 to 300 feral cattle roams the Kona unit of the Hakalau Forest NWR, where individuals of

Cyanea marksii

are reported (USFWS 2010, pp. 3-15, 4-86). Field biologists have observed cattle-induced habitat degradation at all elevations in this refuge unit, including within the montane wet ecosystem that supports individuals of

Cyanea marksii

(PEPP 2007, p. 61; USFWS 2010, pp. 1-15, 2-13, 4-10, 4-58-4-59, 4-82, 4-86; Bio 2011, pers. comm.; Krauss 2012, pers. comm.). In addition, the host plants (

Charpentiera

spp. and

Pisonia

spp.) of the picture-wing fly

Drosophila digressa

have decreased throughout their ranges due to impacts from cattle browsing in the lowland mesic and montane mesic ecosystems (Science Panel 2005, pp. 1-23; Magnacca 2011b, in litt.; Magnacca 2013, in litt.). Feral cattle browsing alters the picture-wing fly's essential microclimate by opening up the canopy, leading to increased desiccation of soil and host plants, which disrupts the host plants' life cycle and decay processes, resulting in the disruption of the picture-wing fly's life cycle, particularly oviposition and larvae substrate (Magnacca

et al.

2008, pp. 1, 32). According to Palikapu Dedman with the Pele Defense Fund, observations of feral cattle in the Ka Lae region where the Lua o Palahemo anchialine pool is located contribute to the degradation of the anchialine pool habitat (Richarson 2012, in litt.). Feral cattle trample and forage on both native and nonnative plants around and near the pool opening at Lua o Palahemo, and increase erosion around the pool and sediment entering the pool. We therefore conclude that feral cattle are a threat to the anchialine pool shrimp

Vetericaris chaceorum

(Richardson 2012, in litt., pp. 1-2). Further, cattle carcasses have been observed within the pool at Lua o Palahemo (Kinzie 2012, in litt.). Due to the steep sides of the pool, animals may fall into the water, and if they die there, their decomposing bodies could have a negative impact on the ability of the pool habitat to support

V. chaceorum

(Kinzie 2012, in litt.).

The destruction or degradation of habitat due to feral sheep is currently a

threat to the montane dry ecosystem on Hawaii Island and its associated species. Feral sheep browse and trample native vegetation, and have decimated large areas of native forest and shrubland on Hawaii Island (Tomich 1986, pp. 156-163; Cuddihy and Stone 1990, pp. 65-66). Browsing erodes top soil, which alters moisture regimes and micro-environments, and results in the loss of native plant and animal taxa (Tomich 1986, pp. 156-163; Cuddihy and Stone 1990, pp. 65-66). In addition, nonnative opportunistic plant seeds get dispersed to disturbed forest sites by adhering to sheep wool coats (Hawaii Division of Forestry and Wildlife (HDOFAW) 2002, p. 3).

In 1962, game hunters intentionally crossbred feral sheep with mouflon sheep and released them on Mauna Kea (Tomich 1986, pp. 156-163). In

Palila

v.

Hawaii Department of Land and Natural Resources

(471 F. Supp. 985 (Haw. 1979)), the Federal court ordered complete removal of feral sheep from Mauna Kea in 1979, because they were harming the endangered palila (

Loxioides bailleui

) by degrading and destroying palila habitat in the montane dry ecosystem. Throughout the past 30 years, attempts to protect the vegetation of Mauna Kea and the saddle from sheep have only been sporadically effective (Scowcroft and Conrad 1992, p. 628). Currently, a large feral population surrounds Mauna Kea and extends into the saddle and northern part of Mauna Loa, including the State forest reserves, where they trample and browse endangered plants (Hess 2008, p. 1). At the U.S. Army's Pohakuloa Training Area, located in the saddle area of the island, biologists have reported that feral sheep are a threat to the last occurrence of the plant species

Schiedea hawaiiensis,

which occurs in the montane dry ecosystem (Mitchell

et al.

2005a; U.S. Army Garrison 2006, pp. 27, 34).

Five of the described ecosystems (lowland mesic, lowland wet, montane dry, montane mesic, and montane wet) on Hawaii Island, and their associated species are currently threatened by the destruction or degradation of habitat due to mouflon sheep. The mouflon sheep (mouflon), native to Asia Minor, was introduced to the islands of Lanai and Hawaii in the 1950s, as a managed game species, and has become widely established on these islands (Tomich 1986, pp. 163-168; Cuddihy and Stone 1990, p. 66; Hess 2008, p. 1). In 1968, mouflon were introduced to Kahuku Ranch (now a unit of HVNP) on Mauna Loa for trophy hunting. By 2008, mouflon ranged over the southern part of Mauna Loa in the Kahuku area on adjacent public and private lands (Hess 2008, p. 1). According to Ikagawa (2011, in litt.), mouflon are found on the slopes of both Mauna Loa and Mauna Kea. Ikagawa (2011, in litt.) also notes that mouflon and mouflon-sheep hybrids are found from sea level to over 3,280 ft (1,000 m) elevation. Mouflon have high reproduction rates; for example, the original population of 11 individuals on the island of Hawaii has increased to more than 2,500 in 36 years, even though mouflon are hunted as a game animal (Hess 2008, p. 3). Mouflon only gather in herds when breeding, thus limiting control techniques and hunting efficiency (Hess 2008, p. 3; Ikagawa 2011, in litt.). Mouflon are both grazers and browsers, and have decimated vast areas of native forest and shrubland through browsing and bark stripping (Stone 1985, p. 271; Cuddihy and Stone 1990, pp. 63, 66; Hess 2008, p. 3). Mouflon also create trails and pathways through thick vegetation, leading to increased runoff and erosion through soil compaction. In some areas, the interaction of browsing and soil compaction has led to a change from native rainforest to grassy scrublands (Hess 2008, p. 3). Field biologists have observed habitat degradation in five of the described ecosystems (lowland mesic, lowland wet, montane dry, montane mesic, and montane wet) that support four plants (

Cyanea marksii, Pittosporum hawaiiense,

Pritchardia lanigera,

and

Schiedea hawaiiensis

) (Bio 2011, pers. comm.; Ikagawa 2011, in litt.; Pratt 2011d, in litt.), and the picture-wing fly (

Drosophila digressa

) (Magnacca 2011b, pers. comm.), in this final rule. Many of the current and proposed fenced exclosures on Hawaii Island are only 4 ft (1.3 m) in height, as they are designed to exclude feral pigs, goats, and sheep. However, a fence height of at least 6 ft (2 m) is required to exclude mouflon sheep, as they can easily jump a 4-ft (1.3-m) fence (Ikagawa 2011, in litt.). Both the increased range of mouflon, as well as the lack of adequately protected habitat, increase the threat of mouflon sheep to additional ecosystems on Hawaii Island.

Between 2010 and 2011, an unauthorized introduction of axis deer (

Axis axis

) occurred on Hawaii, for purposes of big game hunting (Kessler 2011, in litt.; Aila 2012a, in litt.). Axis deer are primarily grazers, but also browse numerous palatable plant species, including those grown as commercial crops (Waring 1996, in litt., p. 3; Simpson 2001, in litt.). They prefer the lower, more openly vegetated areas for browsing and grazing; however, during episodes of drought (e.g., from 1998-2001 on Maui (Medeiros 2010, pers. comm.)), axis deer move into urban and forested areas in search of food (Waring 1996, in litt., p. 5; Nishibayashi 2001, in litt.). Like goats, axis deer are highly destructive to native vegetation and contribute to erosion by eating young trees and young shoots of plants before they can become established, creating trails that can damage native vegetative cover, promoting erosion by destabilizing substrate and creating gullies that convey water, and by dislodging stones from ledges that cause rockfalls and landslides and damage vegetation below (Cuddihy and Stone 1990, pp. 63-64). The unauthorized introduction of axis deer on Hawaii Island is a concern due to the devastating impacts of habitat destruction by axis deer in nine ecosystems (coastal, lowland dry, lowland mesic, lowland wet, montane dry, montane mesic, montane wet, dry cliff, and wet cliff) on the islands of Kahoolawe, Lanai, and Maui (Mehrhoff 1993, p. 11; Anderson 2002, poster; Swedberg and Walker 1978, cited in Anderson 2003, pp. 124-125; Perlman 2009, in litt., pp. 4-5; Hess 2008, p. 3; Hess 2010, pers. comm.; Kessler 2010, pers. comm.; Medeiros 2010, pers. comm.). As reported on the islands of Kahoolawe, Lanai, and Maui, the spread of axis deer into nine of the described ecosystems (coastal, lowland dry, lowland mesic, lowland wet, montane dry, montane mesic, montane wet, dry cliff, and wet cliff) on Hawaii Island will lead to similar habitat degradation and destruction if the deer are not controlled. The results from the studies above, in addition to the confirmed sightings of axis deer on Hawaii Island, suggest that axis deer will significantly alter these ecosystems and directly damage or destroy native plants if they become established. Although habitat degradation due to axis deer has not yet been observed on Hawaii Island, we believe it is reasonable to assume similar habitat effects on this island. Based on the prevailing evidence of the documented impacts to native ecosystems and individual plants on the other islands, we determine that the expanding population of axis deer on the Island of Hawaii, while not currently resulting in population-level effects to native plants, is expected to do so in the future if the deer are not managed or controlled. See Factor D for further information regarding State efforts to eradicate this species.

In summary, the 15 species dependent upon the 10 ecosystems identified in this final rule (anchialine pool, coastal, lowland dry, lowland mesic, lowland

wet, montane dry, montane mesic, montane wet, dry cliff, and wet cliff) are exposed to the ongoing threat of feral ungulates (pigs, goats, cattle, sheep, and mouflon sheep). Additionally, if not adequately managed or controlled, impacts from axis deer may also become a threat to these ecosystems in the future. These negative impacts result in the destruction and degradation of habitat for these 15 native species on Hawaii Island. The effects of these nonnative animals include the destruction of vegetative cover; trampling of plants and seedlings; direct consumption of native vegetation; soil disturbance and sedimentation; dispersal of alien plant seeds on hooves and coats, and through the spread of seeds in feces; alteration of soil nitrogen availability; and creation of open, disturbed areas conducive to further invasion by nonnative pest plant species. All of these impacts lead to the subsequent conversion of a plant community dominated by native species to one dominated by nonnative species (see “Habitat Destruction and Modification by Nonnative Plants,” below). In addition, because these mammals inhabit terrain that is often steep and remote (Cuddihy and Stone 1990, p. 59), foraging and trampling contributes to severe erosion of watersheds and degradation of streams (Dunkell

et al.

2011, pp. 175-194). As early as 1900, there was increasing concern expressed about the integrity of island watersheds, due to effects of ungulates and other factors, leading to the establishment of a professional forestry program emphasizing soil and water conservation (Nelson 1989, p. 3).

Habitat Destruction and Modification by Nonnative Plants

Native vegetation on all of the main Hawaiian Islands has undergone extreme alteration because of past and present land management practices, including ranching, the deliberate introduction of nonnative plants and animals, and agricultural development (Cuddihy and Stone 1990, pp. 27, 58). The original native flora of Hawaii (species that were present before humans arrived) consisted of about 1,000 taxa, 89 percent of which were endemic (species that occur only in the Hawaiian Islands). Over 800 plant taxa have been introduced from elsewhere, and nearly 100 of these have become pests (e.g., injurious plants) in Hawaii (Smith 1985, p. 180; Cuddihy and Stone 1990, p. 73; Gagne and Cuddihy 1999, p. 45). Of these 100 nonnative pest plant species, over 35 species have altered the habitat of 14 of the 15 species in this final rule (only the anchialine pool shrimp is not directly impacted by nonnative plants (see Table 3)).

The most-often cited effects of nonnative plants on native plant species are competition and displacement. Competition may be for water, light, or nutrients, or it may involve allelopathy (chemical inhibition of other plants). Alien plants displace native species of plants by preventing their reproduction, usually by shading and taking up available sites for seedling establishment. Alien plant invasions alter entire ecosystems by forming monotypic stands, changing fire characteristics of native communities, altering soil-water regimes, changing nutrient cycling, or encouraging other nonnative organisms (Smith 1989, pp. 61-69; Vitousek

et al.

1987, pp. 224-227).

Nonnative plants pose serious and ongoing threats to 14 of the 15 species (not the anchialine pool shrimp) in this final rule throughout their ranges by destroying and modifying habitat. They can adversely impact microhabitat by modifying the availability of light and nutrient cycling processes, and by altering soil-water regimes. They can also alter fire regimes affecting native plant habitat, leading to incursions of fire-tolerant nonnative plant species into native habitat. Alteration of fire regimes clearly represents an ecosystem-level change caused by the invasion of nonnative grasses (D'Antonio and Vitousek 1992, p. 73). The grass lifeform supports standing dead material that burns readily, and grass tissues have large surface-to-volume ratios and can dry out quickly (D'Antonio and Vitousek 1992, p. 73). The flammability of biological materials is determined primarily by their surface-to-volume ratio and moisture content, and secondarily by mineral content and tissue chemistry (D'Antonio and Vitousek 1992, p. 73). The finest size classes of material (mainly grasses) ignite and spread fires under a broader range of conditions than do woody fuels or even surface litter (D'Antonio and Vitousek 1992, p. 73). The grass life form allows rapid recovery following fire; there is little above-ground structural tissue, so almost all new tissue fixes carbon and contributes to growth (D'Antonio and Vitousek 1992, p. 73). Grass canopies also support a microclimate in which surface temperatures are hotter, vapor pressure deficits are larger, and the drying of tissues more rapid than in forests or woodlands (D'Antonio and Vitousek 1992, p. 73). Thus, conditions that favor fire are much more frequent in grasslands (D'Antonio and Vitousek 1992, p. 73).

Nonnative plants outcompete native plants by growing faster, and some may release chemicals that inhibit the growth of other plants. Nonnative plants may also displace native species by preventing their reproduction, usually by shading and taking up available sites for seedling establishment (Vitousek

et al.

1987, pp. 224-227). These competitive advantages allow nonnative plants to convert native-dominated plant communities to nonnative plant communities (Cuddihy and Stone 1990, p. 74; Vitousek 1992, pp. 33-35).

In summary, nonnative plants adversely impact native habitat in Hawaii, including 9 of the described Hawaii Island ecosystems that support 14 of the 15 species (not the anchialine pool shrimp), and directly adversely impact the 13 plant species, by: (1) Modifying the availability of light through alterations of the canopy structure; (2) altering soil-water regimes; (3) modifying nutrient cycling; (4) altering the fire regime affecting native plant communities (e.g., successive fires that burn farther and farther into native habitat, destroying native plants and removing habitat for native species by altering microclimatic conditions to favor alien species); and (5) ultimately converting native-dominated plant communities to nonnative plant communities (Smith 1985, pp. 180-181; Cuddihy and Stone, 1990, p. 74; D'Antonio and Vitousek 1992, p. 73; Vitousek

et al.

1997, p. 6).

A summary of the specific impacts of nonnative plant species is included below. Please refer to the proposed rule (77 FR 63928; October 17, 2012) for a list of nonnative plants organized by their ecosystems, a detailed discussion of their specific negative effects on the 14 affected Hawaii Island species, and the literature cited for each nonnative plant species. In particular, we note that we provide discussions of nonnative plants in coastal, lowland wet, dry cliff, and wet cliff ecosystems in this rule (below), but the discussions for nonnative plants in the lowland dry, lowland mesic, montane dry, montane mesic, and montane wet ecosystems can be found in the October 17, 2012, proposed rule (77 FR 63928). Based on comments we received on the proposed rule, we have also added information below regarding the nonnative plants wedelia, strawberry guava, and skunk weed that pose threats to three plants,

Bidens hillebrandiana

ssp.

hillebrandiana

(threats from wedelia),

Cyanea tritomantha

(threats from strawberry guava), and

Cyrtandra nanawaleensis

(threats from skunk weed), in this final rule.

•

Andropogon virginicus

may release allelopathic substances that dramatically decrease native plant reestablishment, and has become dominant in areas subjected to natural or human-induced fires.

•

Anemone hupehensis

var.

japonica

has wind-distributed seeds, and resists grazing because of toxic chemicals that induce vomiting when ingested.

•

Angiopteris evecta

forms dense stands that displace and shade out native plants.

•

Axonopus fissifolius

can outcompete other grasses in wet forests and bogs and outcompetes native plants for moisture.

•

Buddleia asiatica

can tolerate a wide range of habitats, forms dense thickets, and is rapidly spreading into wet forest and lava and cinder substrate areas in Hawaii, displacing native vegetation.

•

Casuarina equisetifolia

forms monotypic stands under which little else grows. It is thought that the roots and needle litter exude a chemical that kills other plants.

•

Clidemia hirta

forms a dense understory, shades out native plants, and prevents their regeneration.

•

Delairea odorata

covers and suppresses growth and germination of native species by carpeting the ground and rooting down at leaf nodes. This species can also grow in the canopy, where it smothers native trees.

•

Digitaria setigera

propagates by seeds and runners; a single flowering stem produces hundreds of seeds.

•

Ehrharta stipoides

creates a thick mat in which other species cannot regenerate; its seeds are easily dispersed by awns (slender, terminal bristle-like process found at the spikelette in many grasses) that attach to fur or clothing.

•

Erigeron karvinskianus

spreads rapidly by stem layering and regrowth of broken roots to form dense mats, crowding out and displacing ground-level plants.

•

Falcataria moluccana

can quickly establish in disturbed and nondisturbed mesic to wet areas. Its rapid growth habit enables it to outcompete slow-growing native trees by reducing light availability, and its abundant, high-quality litter al

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