Chapter 5. Overview of Native and Polynesian-Introduced Species of the Hawaiian Islands Life in the Islands, Evolutionary Processes, and Threats to Native Species
[Illustration: Native Bird, Fish, and Sea Turtle]
Part 1: Evolutionary Processes in the Hawaiian Islands Colonization—Native and Endemic Species Methods of Arrival—How Native Species Traveled to the Hawaiian Islands Endemism and Adaptive Radiation
Part 2: Native and Polynesian-Introduced Species of the Hawaiian Islands Past Extinctions—Bird Species Native Hawaiian Birds Bishop Museum’s Hawaii Biological Survey Arthropods Snails and Other Invertebrates Reptiles and Amphibians Fish Native Reef Fish Sharks, Rays and Eels Other Marine Species Land and Sea Mammals Native and Polynesian-Introduced Plants Other Polynesian-Introduced Species Intentional Polynesian Introductions Unintentional Polynesian Introductions Geckos Skinks Plants Snails Other Polynesian-Introduced Species
Part 3: Threats to Native Species Endangered Species and Extinctions Threats to Native Species—Before 1778 Threats to Native Species—Post-Contact A Summary of the Primary Threats to Native Species of the Hawaiian Islands Preventing Extinctions
Part 1: Evolutionary Processes in the Hawaiian Islands
[Illustration: Hawaiian Islands, Frigatebird]
Key to Species Classifications: Native—Arrived in the Hawaiian Islands without the aid of humans (indigenous), or evolved in the Hawaiian Islands (endemic). Indigenous—Native to the Hawaiian Islands and other places. Endemic—Evolved in the Hawaiian Islands from an indigenous species; native to the Hawaiian Islands and nowhere else. Polynesian Introduction—Brought to the Hawaiian Islands by the early Polynesian settlers. Post-Contact Introduction—Brought to the Hawaiian Islands after Western contact (1778). Naturalized—Not native to the Hawaiian Islands, but now growing wild in the Hawaiian Islands.
After the volcanoes of the Hawaiian Islands formed and cooled, the weathering of the sun, rain, and wind began to break down the lava rock into soil. Primitive plants known as lichens also helped to create soil, allowing more plant species to take hold. These plant species in turn created new habitats that allowed the Hawaiian Islands’ original colonizing species to evolve, over millions of years, into thousands of plant, insect, and bird species.
Relatively few species were able to reach the Hawaiian archipelago before humans arrived, but those species that did become established were able to take advantage of the Hawaiian Islands’ diversity of habitats and ecosystems. The original colonizing species evolved into an array of uniquely adapted species that gradually transformed the Hawaiian Islands barren and rocky volcanoes into lush tropical islands full of life.
Colonization—Native and Endemic Species
Located more than 2,400 miles (3,862 km) from the nearest continental land mass, the Hawaiian Islands were not easily colonized by new species prior to the arrival of humans. The extreme geographical isolation of the Hawaiian Islands also limited the types and the numbers of species that became established.
All of the Hawaiian Islands’ native species are divided into two categories: indigenous and endemic. Species that somehow made it to the Hawaiian Islands without the aid of humans, and then were able to establish native populations, are now classified as indigenous to the Hawaiian Islands. These indigenous species are native to the Hawaiian Islands, and are also native to other places (wherever the species originally came from).
The other category of native species is the endemic Hawaiian species, which evolved in the Hawaiian Islands from species that were already established in (and thus native to) the Hawaiian Islands. Endemic Hawaiian species may have evolved from an indigenous Hawaiian species or from another endemic Hawaiian species. In either case, all endemic Hawaiian species evolved in the Hawaiian Islands, and thus are unique to the Hawaiian Islands (found naturally nowhere else). Hawai‘i’s geographic isolation and diversity of habitats and food sources led to the evolution of many new (endemic) species from the relatively few colonizing species that were able to make it to the Hawaiian Islands and reproduce. This resulted in extremely high rates of endemism for various categories of species. For example, about 79% of the native flowering plant species of the Hawaiian Islands are endemic, along with about 70% of native ferns of the Hawaiian Islands. About 25% of native Hawaiian reef fish are endemic to the Hawaiian Islands.
Methods of Arrival—How Native Species Traveled to the Hawaiian Islands Indigenous species inhabiting the Hawaiian Islands arrived in the Islands by a variety of methods. For example, seeds floated to the Hawaiian Islands on the ocean, or were carried in the wing feathers or in the digestive tracts of birds, or in mud on the birds’ feet. The colonizing bird species may have come to the Hawaiian Islands on their natural migration tracks, or arrived after being blown off course by a storm. Snails and other species may have been transported to the Hawaiian Islands by hitching a ride on floating logs or on rafts of debris that floated over the Pacific Ocean and washed up on Hawaiian shores. Tiny snail eggs were also carried to the Hawaiian Islands embedded in mud in the feet of birds. Insects and spiders in far distant places were lifted by winds high into the jetstream, more than 8 miles (13 km) up, where they were carried out over the Pacific Ocean at more than 120 miles per hour (193 km/hr) and then set down on the Hawaiian Islands, there to remain and multiply.
For many species, the journey to the Hawaiian Islands was too long and the odds too great. Even over many millions of years the species were never able to reach the Hawaiian Islands, or if they did, there was not suitable habitat for them to survive. Some species may have arrived in the Hawaiian Islands in such small numbers that they were not able to establish breeding (reproducing) populations.
It is estimated that about 39% of plant species in the Hawaiian Islands came in the stomachs of birds; about 13% arrived in mud on birds’ feet; about 23% arrived after being stuck to the feathers of birds; about 9% arrived on rafts of seaweed or debris on the ocean; and about 1.4% came by floating through the air to the Hawaiian Islands.[1]
Many plants and animals that are very common in other places never became established in the Hawaiian Islands until humans brought the species to the Hawaiian Islands. For example, in the Hawaiian Islands there are no gymnosperms (e.g., pine trees), no native amphibians, no native terrestrial reptiles, and no native ants. However, all of these organisms are now found in the Hawaiian Islands, having been introduced to the Islands by humans.
Endemism and Adaptive Radiation
Species that somehow were able to traverse the vast distance over the Pacific Ocean to the Hawaiian Islands and become established in the Hawaiian Islands (without the aid of humans) were then able to adapt to a wide variety of environmental conditions (ecological niches) found in the Hawaiian Islands. Populations of species often became divided, with each separated sub-population adapting to different habitats and food sources, and eventually evolving into completely new species.
For example, one finch species that established itself in the Hawaiian Islands millions of years ago evolved into at least 50 species and subspecies of Hawaiian honeycreepers. The evolutionary process whereby one species evolves into many new species is known as adaptive radiation. The Hawaiian Islands are renowned for their historical record of adaptive radiation, particularly regarding bird and insect species. Over hundreds of generations, many plant, animal, and insect species evolved adaptations to the unique conditions found on each of the Hawaiian Islands. Many of these adaptations distinguished the separate populations, which eventually evolved into unique species or subspecies, endemic to each particular island. For example, the fragrant mokihana plant (Pelea anisata) is endemic to the uplands of Kaua‘i, as is the puaiohi (Myadestes palmeri), a tiny pink-legged forest bird. Some native Hawaiian species may be endemic to very small areas. For example, the endangered plant known as koki‘o ke‘oke‘o (Hibiscus waimeae subspecies hannerae, white hibiscus) is found only in a few northwest Kaua‘i valleys, including the valleys of Limahuli and Hanakāpī‘ai. Native Hawaiian species isolated within a particular valley for many thousands of years often developed variations that aided their survival in the particular habitat. This isolation often led to distinct subpopulations of species that eventually became distinguishable as endemic subspecies or completely new species. An ancient proverb states, “‘O‘opu peke o Hanakāpī‘ai.” (“The stunted ‘o‘opu fish of Hanakāpī‘ai,” referring to “...the ‘o‘opu of Hanakāpī‘ai, which are said to be plump and shorter in length than those elsewhere. Sometimes applied humorously to a short, plump person.”[2] The evolutionary process of adaptive radiation allowed the relatively few colonizers of the Hawaiian Islands to develop into a vast array of life forms that today constitutes the diverse native flora and fauna of the Hawaiian Islands. (For a more in-depth explanation of adaptive radiation, see this book’s Introduction as well as the Honeycreepers and Honeyeaters section in Native Birds of the Hawaiian Islands, Chapter 7.)
Part 2—Native and Polynesian-Introduced Species of the Hawaiian Islands
Past Extinctions—Bird Species
Worldwide there are about 9,700 bird species, but only about 2% of these bird species are native to the Hawaiian Islands. When Captain Cook arrived in the Hawaiian Islands in 1778, there were about 80 species of endemic Hawaiian landbirds, along with at least 24 species and subspecies of native Hawaiian seabirds (4 endemic) and 34 species and two subspecies of waterbirds (29 endemic). More than 35 endemic Hawaiian landbirds had already gone extinct by 1778 when Cook’s arrival established Western contact with the Hawaiian Islands.
Bird species that were extinct prior to Cook’s arrival met their demise from a variety of causes. The Polynesians settlers of the Hawaiian Islands hunted birds for food and for feathers, though sometimes the birds were released unharmed after some of the feathers were collected.(See Native Birds of the Hawaiian Islands, Chapter 7.)
Pua‘a (Sus scrofa, pigs) and ‘īlio (Canis familiaris, dogs) brought by the Polynesians also had an effect on island bird populations, as did ‘iole (Rattus exulans, Polynesian black rats) that hitched a ride on the voyaging canoes. These pigs, dogs, and rats were particularly detrimental to the Hawaiian Islands’ ground-nesting birds, which were also the most likely birds to be hunted by humans for food.
Aia a pohā ka leo o ka ‘a‘o, kāpule ke momona o ka ‘uwa‘u i ka puapua. When the ‘a‘o birds’ voices are distinctly heard, the ‘uwa‘u birds are fat even to the very tails. The ‘a‘o bird was not heard during the nesting season. When the fledglings emerged and their cries were heard, the season had come when young ‘uwa‘u were best for eating, and the people went to snare them. (Pukui: 32-6)
About 23 of the bird species that became extinct prior to Western contact were ground-nesting birds. Known only from subfossil discoveries, these species included flightless ducks (four species); flightless ibises (three species); flightless rails (at least 10 species); and flightless geese (six species). Some of the flightless geese species were more than 3 feet (.9 m) tall.[iii]
Other species that went extinct prior to Western contact included at least four long-legged owl species, one shearwater, one bald eagle, two crow species, one petrel, a harrier, and more than 21 honeycreeper species along with one honeyeater species. It is possible that some of these bird species may have become extinct prior to Polynesian settlement in the Hawaiian Islands, but most (if not all) of these bird species probably became extinct after the Polynesians arrived in the Hawaiian Islands, but before Cook arrived in 1778 establishing Western contact.
Since 1778, at least 24 more native Hawaiian land bird species or subspecies have become extinct.[iv] For example, the native kioea (Chaetoptila angustipluma) was last seen in 1859. In addition to the 59+ extinct endemic Hawaiian bird species and subspecies, more than 30 other endemic Hawaiian bird species are currently considered rare or endangered, and six of these species may already be extinct. (See Forest and Mountain Birds of the Hawaiian Islands, Chapter 7.)
Native Hawaiian Birds
Some bird species may have inhabited the Hawaiian Islands only after the Polynesian settlers arrived. This occurred because the original Hawaiian settlers, in the course of their food-producing activities, altered the native landscape by creating particular types of habitat (e.g., lo‘i kalo, or taro patches) that may have allowed certain wetland bird species to take hold.
The transformation of the environment by the Polynesian settlers of the Hawaiian Islands created substantial new areas of aquatic habitat. These new wetland habitats likely allowed some native species to expand their range, and may have allowed other species to become established. For example, the koloa (Anas wyvilliana, duck) and the ‘alae ula (Gallinula chloropus sandvicensis, gallinule) may have become established only after new wetland habitat was created by the Polynesian settlers. This is conjectured by researchers due to the absence of these birds from Pleistocene bird fossil and subfossil deposits excavated from various sites around the Hawaiian Islands.
The Polynesians’ introduction of the Polynesian black rat (Rattus exulans) to the Hawaiian Islands also provided a new food source for raptors, and may have allowed certain bird species (e.g., owls) to become established in the Hawaiian Islands. In a published study entitled Fossil evidence for a diverse biota from Kaua‘i and its transformation since human arrival(2001),[v]the authors detail the excavation of a Kaua‘i sinkhole and cave system on the south coast of Hawai‘i Island. Remains of pueo (Asio flammeus, Short-eared owls) did not show up in prehuman sediments but were found in sediments corresponding to the time after the Polynesians arrived.
The excavation results led the authors to note that the findings were “...consistent with Olson and James’ suggestion that this rodent-catching owl was able to colonize the archipelago only after the Pacific rat had been introduced by the Polynesians.”[vi] The authors also stated that, “...surprisingly...the only anseriform that has escaped extinction on the island and that one might expect to find in a wetland deposit, the Hawaiian Duck, or Koloa (Anas wyvilliana), is rare or absent...only a few bones from the site have been tentatively referred to as A. wyvilliana.”[vii] Thus the evidence remains inconclusive in regards to the whether koloa were found in the Hawaiian Islands before humans arrived or if they instead became established in the Hawaiian Islands only after the initial Polynesian settlement of the archipelago (and subsequent increase in wetland habitat). Information regarding the exact time sequence of particular bird species inhabiting the Hawaiian Islands is still being gathered, and a more accurate picture is gradually emerging of the role that Polynesians’ played in creating new habitats that led to new bird species becoming established in the Hawaiian Islands. Today, at least 22 species of marine birds are native to the Hawaiian Islands, with a total population of more than 12 million birds. More than 60% of these marine birds are noddies and terns. At least 33 species of geese and ducks (many migratory) are native to the Hawaiian Islands, along with five species of waterbirds, all endangered. (See Chapter 67) As part of Bernice Pauahi Bishop Museum’s ongoing Hawaii Biological Survey, updated in the Records of the Hawaii Biological Survey for 2000,[viii] editors Lucius G. Eldredge and Scott E. Miller documented 228 bird species seen in the Hawaiian Islands (including 168 breeding and non-breeding migratory birds, some seen only rarely) as well as 60 endemic resident species. If the 46 introduced bird species are included, the total rises to 274 birds species in the Hawaiian Islands. The study by Eldredge and Miller also noted that a total of 475 bird species are “known to occur or be sighted in Hawai‘i.”[ix] This number included extinct species (35+ becoming extinct before Cook arrived in 1778; 32 of these described from subfossils), and another 16 species extinct since 1778 (though a greater number are now presumed extinct, see above), and also included 150+ non-native birds that have been seen in the Hawaiian Islands but are not considered introduced species because they have not yet established breeding populations in the Hawaiian Islands. The current total count of bird species in the Hawaiian Islands is apparently now at least 495, as the Records of the Hawaii Biological Survey for 2000[x]documented 10 new indigenous species (3 endemic and 7 introduced species), increasing the total number of bird species now established in the Hawaiian Islands to 294 species, including 241 native species (178 indigenous, 63 endemic) and 53 introduced species. (See Native Birds of the Hawaiian Islands, Chapter 7.)
Bishop Museum’s Hawaii Biological Survey The Hawaii Biological Survey (with no ‘okina in “Hawaii”) is a Bishop Museum program that was established in 1992 by Hawai‘i’s State Legislature to maintain an ongoing inventory of the Hawaiian Islands’ natural history, including non-native as well as native species. In coordination with a variety of state, federal and private groups (e.g., U.S. Fish & Wildlife Service, Nature Conservancy, University of Hawai‘i, and others), the Hawaii Biological Survey’s mission is to “gather, analyze, and disseminate the biological information necessary for the wise stewardship of Hawai‘i’s biological resources,” and also to “conduct a coordinated inventory and monitoring program to assess the overall status and trends in the abundance, health, and distribution of plants and animals, as well as the ecosystems upon which they depend.”[xi] Bishop Museum scientists have done extensive studies to determine the total number of species found in the Hawaiian Islands, and have assembled a comprehensive tabulation of scientifically documented species, published by Bishop Museum and updated annually, including the Records of the Hawaii Biological Survey for 2000,[xii] published on March 25, 2002. This count of the Hawaiian Islands’ endemic, indigenous, and introduced species, edited by Neal L. Evenhuis and Lucius G. Eldredge, includes plants, trees, mollusks (e.g., snails), invertebrates (e.g., insects), fish, birds, reptiles (e.g., sea turtles), mammals (e.g., bats, monk seals, whales and dolphins), fungi, lichens, protists (including algae and human parasitic protists), and helminths. A total of 23,680 native Hawaiian species were documented, including 9,151 indigenous species, 9,456 endemic species, and 5,073 human-introduced species. Viruses and bacteria were not included. Of the 18,607 native Hawaiian species documented, about half (9,456 species) are endemic to the Hawaiian Islands. The documented native Hawaiian species include: 1,995 protist species (82 endemic); 2,088 fungi and lichen species (240 endemic); 1,163 flowering plants (918 endemic); 719 other native plants (241 endemic); 1,243 mollusks (962 endemic); 1,143 fish species (149 endemic); 4 reptiles (sea turtles, none endemic); 241 bird species (63 endemic); and 25 mammals (2 endemic) and 9,270 native invertebrates (7,239 endemic). These native Hawaiian invertebrates include 6,284 arthropods [which included 5,818 insect species (5,462 endemic)] and 466 other arthropods (366 endemic)], along with 1,743 other invertebrates (449 endemic). About 79% of native flowering plants in the Hawaiian Islands are endemic (918 of 1,163 species) along with approximately 26% of native birds, though nearly all native land birds of the Hawaiian Islands are endemic. About 94% of native insects of the Hawaiian Islands are endemic, as are 77% of native mollusks, 78% of native invertebrates, and 13% of native fish. Counts of indigenous and endemic Hawaiian species are constantly changing as researchers discover new species, analyze relationships between species (and subspecies) and further refine scientific classifications. F.R. Warshauer, in The Atlas of Hawai‘i: Third Edition (1998), and relying in part on the Bishop Museum’s Hawaii Biological Survey, reported 142 native bird species (including extinct species); 7,800 described arthropods (out of an estimated 10,000 to 12,000 native Hawaiian arthropods); 1,300 vascular plants; 1,600 lower plants; 788 mollusks, 166 other invertebrates; and one mammal (not counting marine mammals), totaling about 15,000 native species in those categories.[xiii]
Arthropods
Arthropods are invertebrates (animals without backbones). All arthropods have a rigid external skeleton as well as segmented body parts and segmented legs (arthropod means “jointed feet”). At least 6,284 native Hawaiian arthropods have been documented.
About 95% of Hawaiian arthropods are insects, while the other 5% are closely related species that are not true insects, including centipedes, millipedes, spiders, and many others.
More than 94% of the Hawaiian Islands’ 5,818 known native insect species are endemic to the Hawaiian Islands. These 5,462 endemic insect species evolved over a period of at least 30 million years, inhabiting the ancient islands of the Hawaiian archipelago. At least 23 species and subspecies of damselflies are endemic to the Hawaiian Islands along with five endemic dragonfly species. (See Damselflies and Dragonflies, Chapter 10.) Many more introduced damselfly and dragonfly species are now established in the Hawaiian Islands. Two butterfly species are native to the Hawaiian Islands, along with about 13 more introduced butterfly species. (See HawaiianButterflies, Chapter 10.) More than 60 cave species (living in caves), are native to the Hawaiian Islands, and many of these are endemic to particular islands. (See Hawaiian Cave Species, Chapter 10.)
Pomace flies, (Drosophilidae and Scaptomyza), are a renowned example of adaptive radiation among the Hawaiian Islands’ insect fauna, with at least 860 documented species of Drosophilidae (in five genera), and perhaps many more yet to be described. All of these flies apparently evolved from a common pair of ancestors that lived in the Hawaiian Islands about 10 million years ago. For thousands of generations (over millions of years) the flies reproduced, and island-hopped from one volcano to the next as the islands eroded away at the northwest end of the chain and emerged at the southeast end of the chain.
Currently there are at least 2,161 known beetle species native to the Hawaiian Islands, and at least 1,416 of these beetle species are endemic to the Hawaiian Islands. There are more species of beetles than any other organism in the Hawaiian Islands, and the Hawaiian Islands also have the world’s highest proportion of beetles to other living things.
Thousands of species of native Hawaiian insects have yet to be scientifically described. It is estimated that the total number of native Hawaiian insects could exceed 10,000 species, which evolved from less than 500original colonizing species.[xiv] Also now established in the Hawaiian Islands are more than 550 introduced terrestrial arthropods and more than 2,700 non-native (introduced) insect species.
Snails and other Invertebrates
The Bishop Museum’s Hawai‘i Biological Survey has documented at least 781 terrestrial gastropods (759 endemic), and 7 Hawaiian freshwater gastropods (all endemic).
Endemic Hawaiian tree snails (terrestrial gastropods) provide another astounding example of adaptive radiation in the Hawaiian Islands, with more than 750 endemic species having evolved from just a few pioneers that reached the Islands several million years ago, probably in the feathers of migratory birds, or in mud on the birds’ feet. The snails are known by the Hawaiian name pūpū kani oe (“shell that sounds long”[xv]), referring to the belief that the snails sing.
Endemic Hawaiian land and freshwater snails evolved from about 25 colonizing species. These extremely adaptable snails had no natural predators, and were once found in a wide variety of habitats where they fed on the fungi that grow on leaves and forest debris (making the snails an important component of native ecosystems). The snails were known to vary considerably from valley to valley, and exhibited many shapes and colors, from gold tints to cream-colored with brown stripes.
Unfortunately, these native terrestrial snails have a very slow reproductive cycle as well as a low birthrate, and populations were significantly diminished by collectors and other causes. The snails are extremely vulnerable to non-native predators, particularly rats (Rattus exulans; R.. rattus;R. norvegicus); introduced snails (e.g., Euglandina rosea, the “cannibal snail,”); and the New Guinea flatworm (Platydemis manokwari), which was introduced to the Hawaiian Islands in the 1990s and now threatens the last, high-elevation habitats of the native snails.[xvi]
There were once more than 300 native Hawaiian snail species in the family Amastridae, but only about 12 or fewer of these species survive today. Recent research studying genetic mutation rates has shown that dozens of tree snail species in the subfamily achatinellinae originated from a single ancestor that likely lived in O‘ahu’s Ko‘olau mountain range about 3 million years ago. The last remaining habitat of native Hawaiian terrestrial snails is at high elevations on Moloka‘i, Lāna‘i, and O‘ahu, as well as some middle elevation areas on Hawai‘i Island and Maui.
Three brackish/freshwater snail species are endemic to the Hawaiian Islands. Two of these species, hapawai (Neritina vespertina) and pīpīwai (Theodoxus cariosus), prefer brackish waters, while a third species, hīhīwai (Neritina granosa) lives in freshwater stream areas. There are four native species of lymnaeid snails (pond snails) that live in streams and ponds as well as lo‘i kalo (taro patches).[xvii]
Overall, at least 1,243 native Hawaiian mollusks have been documented, and about 80% of these mollusk species are snails, sea slugs, and related species. Native Hawaiian mollusks are invertebrates that evolved from about 25 original immigrant species. There are also thousands of other native Hawaiian species of invertebrates (animals without backbones), including crabs, shrimp, and sea urchins.
Two native Hawaiian shrimp species, ‘ōpaekala‘ole (Atyoida bisulcata) and ‘ōpae ‘oeha‘a (Macrobrachium grandimanus), live in streams, while a third, ‘ōpae ‘ula (Halocaridina rubra) prefers anchialine ponds that are connected to the sea only by the percolations of saltwater and freshwater through lava rock. Heteromyenia baileyi is a native (indigenous) freshwater sponge that sometimes attains a bright green color.
Reptiles and Amphibians No terrestrial reptiles or amphibians are considered native to the Hawaiian Islands, though there remains a possibility that some lizard species arrived in the Hawaiian Islands on their own (see below). Marine reptiles native to the Hawaiian Islands include five sea turtle species, some seen only rarely in Hawaiian waters. The two sea turtle species that come onto Hawaiian shores to lay their eggs are honu (Chelonia mydas, green sea turtle) and honu‘ea (Eretmochelys imbricata, hawksbill sea turtle). On Monday, October 7, 2002, a small, female Olive ridley sea turtle (Lepidochelys olivacea) came ashore at Bayfront Beach on the island of Hawai‘i’s Hilo Bay. If confirmed, this is only the second reported sighting of a nesting Olive ridley sea turtle in the Hawaiian Islands. (See Hawaiian Sea Turtles, Chapter 6.) Another marine reptile occasionally seen in Hawaiian waters (and thus considered native) is the Yellow-bellied sea snake (Pelamis platurus), a venomous yellow and black reptile that is about 29 inches (74 cm) long. A distant relative of the cobra, the Yellow-bellied sea snake is the only truly pelagic (open-ocean) sea snake of 55 known sea snake species. [Illustration: Pelamis platurus (see above).]
Three skink species and four gecko species were established in the Hawaiian Islands before Western contact. These species likely came as stowaways aboard the voyaging canoes of the Polynesian settlers of the Hawaiian Islands, though it is possible some of these lizards may have reached the Hawaiian Islands on their own (e.g., by rafting on a log or on other ocean debris).
As of 2004 at least 21 lizard species have been found in the wild in the Hawaiian Islands (and thus are assumed to have established breeding populations). These lizards include geckos, skinks, anolis lizards, chameleons, horned lizards, and iguanas. Some of these lizard species are found throughout the Hawaiian Islands, while others are only found in very small areas on particular islands. Three of the lizard species are (presumably) no longer found in the wild in the Hawaiian Islands. (See Geckos and Other Reptiles, Chapter 10.) No terrestrial snakes are native to the Hawaiian Islands, but there is one introduced snake species, called the Island blind snake (Ramphotyphlops braminus), which arrived in the Hawaiian Islands sometime around the 1930s in the soil in potted palm trees imported from the Philippines for landscaping. Now well established in the Hawaiian Islands, the Island blind snake is a burrowing snake that is about 6½ inches (16.5 cm) long, feeds on insects and small invertebrates, and spends most of its life underground.
Fish Of the 24,000 known species of fish in the world, about 1,143 are native to the Hawaiian Islands, including 149 endemic fish species. About 536 of these fish species are inshore fishes found near reefs and other nearshore areas to a depth of about 200 feet (61 m). About 25% of these 536 inshore species are endemic to the Hawaiian Islands. The overall rate of endemism for native Hawaiian fish is about 13% (149 endemic species and 994 indigenous species). (See Hawaiian Reef Fish, Chapter 6.) Fish native to the Hawaiian Islands also include five freshwater species, known as ‘o‘opu, which spend part of their lives in the ocean. These five ‘o‘opu fish species are endemic to the Hawaiian Islands, and include four species in the family Gobiidae (commonly called gobies) and one species in the family Eleotridae(‘o‘opu ‘akupa, Eleotris sandwicensis). All five Hawaiian ‘o‘opu species are born as larvae in freshwater streams. The tiny larvae are then washed down into the ocean where they develop into fish before returning to the stream where they were born and swimming back up through the current. ‘O‘opu have specially adapted pelvic fins, which are fused to form a sucking disc, allowing the fish to climb up rocks and waterfalls and reach the upper levels of the streams to lay eggs and complete their life cycle. An ancient Hawaiian proverb refers to the ‘o‘opu as “Ka i‘a a ka wai nui i lawe mai ai,” (“The fish borne along by the flood.”[xviii]
Native Reef Fish Native reef fish species of the Hawaiian Islands include:
Ø5 native species of angelfishes (Pomacanthidae).
Ø24 native species of butterflyfishes (Chaetodontidae).
Ø7 native species of parrotfishes (Scaridae).
ØAt least 24 native species of surgeonfishes (Acanthuridae).
Ø43 native species of wrasses (Labridae).
Ø8 native species of filefishes (Monacanthidae).
Ø25 native species of scorpionfishes (Scorpaenidae).
Ø11 native species of triggerfishes (Balistidae).
Ø1 native species of trumpetfish (Aulostonus chinensis).
Ø2 native species of cornetfishes (Fistulariidae).
Ø17 species of damselfishes (Pomacentridae)
(See Hawaiian Reef Fish, Chapter 6, for more information about the fish listed above.)
The endemic āholehole fish (Kuhlia sandvicensis, young stage of āhole, Hawaiian flagtail) sometimes enters lower stream areas, as do pua ‘ama‘ama (Mugil cephalus, young stage of ‘ama‘ama, mullet). ‘Ama‘ama were the main fish raised in ancient Hawaiian loko i‘a (fishponds). Also raised in fishponds were awa(Chanos chanos, milkfish), which reach a length of 6 feet (1.8 m).
Sharks, Rays and Eels At least 41[xix] known species of sharks may be seen in Hawaiian waters (and are thus considered native to the Hawaiian Islands), and 20 of these shark species inhabit deep waters. Six shark species are relatively common around the Hawaiian Islands. (See Sharks, Chapter 6.) There are five native ray species, including a manta ray, an eagle ray, and three stingray species. (See Rays, Chapter 6.) There are at least 38 native Hawaiian species of moray eel (Muraenidae), along with 16 native species of snake eels (Ophichthidae) and 3 native species of conger and garden eels (Congridae). (See Eels, Chapter 6.)
Other Marine Species More than 860 marine and freshwater algae species are native to the Hawaiian Islands, including at least 80 endemic algae species. Of the more than 340 known native Hawaiian species of red limu, at least 67 are endemic and most are edible. (See Limu in Native Plants and Ferns of the Hawaiian Islands, Chapter 8.) There are at least 150 different native Hawaiian coral species (approximately 20 to 30 percent are endemic), including 47 hard stony corals and more than 100 species of sea fan and other soft corals. (See Coral Reefs, Chapter 6.)
Land and Sea Mammals There are 78 species of cetaceans worldwide, including all whales, dolphins, and porpoises. At least 22 cetacean species are considered native to the waters surrounding the Hawaiian Islands, though some are only occasionally seen near the Hawaiian Islands. The 22 native Hawaiian cetacean species include 17 toothed cetaceans, known as Odontocetes (e.g., dolphins), and five baleen whales, known as Mysticetes (e.g., humpback whales). (See Humpback Whales, and Spinner and Bottlenose Dolphins, Chapter 6.) Aside from cetaceans, the only sea mammal native to the Hawaiian Islands is the Hawaiian monk seal, which often feeds near shore and then rests during the day on Island beaches. (See Hawaiian Monk Seals, Chapter 6.) There are about 4,600 mammal species in the world, but just one native Hawaiian land mammal: ‘ōpe‘ape‘a (Lasiurus cinereus semotus, Hawaiian hoary bat). This furry insect-eating bat weighs in at about 6 ounces (170 g) with a wingspan up to 14 inches (36 cm). (See Hawaiian Hoary Bat, Chapter 10.)
Native and Polynesian-Introduced Plants
Before Western contact began to drastically change the landscape of the Hawaiian Islands, there were more than 1,700 known native Hawaiian plant species, including more than 1,000 native flowering plant species that had evolved from less than 300 original immigrants. At least 178 fern species are native to the Hawaiian Islands (124 endemic species). These native ferns evolved from about 135 colonizing species. At least 22 species of fern allies are native to the Hawaiian Islands, and seven of these fern ally species are endemic. The 1999 Manual of the Flowering Plants of Hawai‘i: Revised Edition[xx]formally recognized 956 species of native Hawaiian flowering plants, including 850 species that are endemic to the Hawaiian Islands. Many more native Hawaiian plant species have since been discovered (see Introduction to Native Plants and Ferns of the Hawaiian Islands, Chapter 8), and are documented in the 2002 Electronic Supplement to the Manual of the Flowering Plants of Hawai‘i[xxi]as well as the Records of theHawaii Biological Survey (see above), with a current total of at least 1,882 documented native Hawaiian plant species (1,159 endemic), including 1,163 native flowering plant species (918 endemic). The extremely high rate of flowering plant endemism (79% endemic) in the Hawaiian Islands is primarily attributed to the geographical isolation of the Islands in the middle of the Pacific Ocean, thousands of miles from any continent. For more information about endemism and the evolution of native plant species, see the introduction to Chapter 8, Native Plants and Ferns of the Hawaiian Islands. The Polynesian settlers of the Hawaiian Islands also brought at least 24 species of plants to the Islands, and may have brought as many as 30 plant species or more. (See Introduction to Chapter 9, Polynesian-Introduced Plants.)
Other Polynesian-Introduced Species The early Polynesian settlers also brought many other species to Hawai’i besides plants. Some of these other species were brought to the Hawaiian Islands intentionally for use as food sources, but other species were brought unintentionally, coming as stowaways on the Polynesians’ voyaging canoes, and then establishing breeding populations in the Hawaiian Islands. The geckos and the skinks listed below are presumed to have arrived in the Hawaiian Islands in this manner (as stowaways on the voyaging canoes), but there remains a possibility that some of these lizard species may instead have arrived in the Hawaiian Islands on their own (e.g., floating to the Hawaiian Islands on rafts of debris), and thus are native to the Hawaiian Islands. Following is a summary of all species known to be introduced to the Hawaiian Islands by the early Polynesian settlers, as well as other species that may have been brought to the Hawaiian Islands, intentionally or unintentionally, by the Polynesians previous to Western contact in 1778.
Intentional Polynesian Introductions
Pua‘a (Sus scrofa, pigs)
Moa(Gallus gallus gallus, chickens)
‘Īlio(Canis familiaris, dogs)
At least 24 (and probably more than 26) plant species
§Neke(Cyclosorus interruptus, formerly Thelypteris interrupta, maiden fern)
§Paspalum scrobiculatum (ricegrass, no known Hawaiian name)
*Some of these plant species may be indigenous Hawaiian species, but have been noted as potential Polynesian introductions based on research. See Introduction to Polynesian-Introduced Plants, Chapter 9 for more information about the status of each of these species.
Other (Unintentional) Polynesian-Introduced Species
§Polynesian black rat (Rattus exulans)
§Ectoparasites
oLaelaps hawaiiensis, and others (carried on rats)[xxiii]
§Freshwater clams*
oPisidium casertanum
oMusculium partumeium
*Found in ancient taro pond, thought to have been transported on taro stock brought to Hawai‘i by early Polynesian settlers.[xxiv]
Endangered Species and Extinctions Currently at least 30 species and subspecies of Hawaiian birds are federally listed as endangered, along with one subspecies listed as threatened.[xxv] About half of the Hawaiian Islands’ endemic birds are already extinct. More than 40 Hawaiian insect species have gone extinct in the last century, along with about 10% of native Hawaiian flowering plants (more than 100 extinct species). Just as the Hawaiian Islands are known as the world’s capital of endemic species, it may be considered the capital of extinct species. At least 24 native Hawaiian bird taxa (species and subspecies) have gone extinct since 1778, from the Laysan rail (Porzana palmeri), which was last seen in 1944, to the Greater koa finch (Rhodacanthis palmeri), last seen in 1896. The Hawai‘i ‘ō‘ō (Moho nobilis), a bird long sought after for its beautiful feathers (see Native Birds of the Hawaiian Islands, Chapter 7) for use in ‘ahu ‘ula (royal cloaks), hasn’t been seen since 1934.[xxvi] [Illustration: Hawai‘i ‘ō‘ō (Moho nobilis)]
Also extinct or likely extinct are 97 taxa (species and subspecies) of plants, 72 taxa of snails, and 74 taxa of insects.[xxvii] The presumably extinct insects include: 15 species of Coleoptera (beetles); five species of Diptera (flies, gnats, and mosquitoes); 15 species of Homoptera (leafhoppers, planthoppers, mealybugs, and scales); 38 species of Lepidoptera (moths and butterflies); one species of Odonata (damselflies and dragonflies); two species of Orthoptera (crickets and grasshoppers); and one species of Hemiptera (true bugs).[xxviii] More than one third of the plant and animal species that are federally listed as endangered are Hawaiian. In all, more than 360 species are listed as either threatened or endangered under the federal and state Endangered Species Acts. These species include more than 275 endangered taxa (species and subspecies) of plants and more than 35 endangered vertebrate species, with at least another 40 invertebrate species listed as threatened. The Manual of the Flowering Plants of Hawai‘i(1999) notes that, of the 1,094 taxa (including species and subspecies) of native Hawaiian plants included in the text, “...423 (38%) are considered to be extinct or threatened to some degree; of these taxa, 107 are presumed extinct (10% of the native taxa), 139 endangered (12%), 39 vulnerable (4%), and 138 rare (12%).”[xxix] The U.S. Fish & Wildlife Service endangered species program (in regards to Hawaiian species) does not count subspecies separately, resulting in different totals and percents in the categories listed above.
Though the Hawaiian Islands are just two tenths of one percent of the size of the United States, the Hawaiian Islands account for nearly 70% of the United States’ historically documented plant and animal extinctions. More than 30% of federally listed endangered species are Hawaiian, along with more than 40% of all birds listed as endangered. More than half of the Hawaiian Islands’ endemic plants are now threatened, endangered, or extinct, along with about half of the Hawaiian Islands’ endemic birds.
Threats to Native Species—Before 1778
The first threats to native species of the Hawaiian Islands came from the ancient Polynesian settlers of the Hawaiian Islands, who brought pua‘a (Sus scrofa, pigs), moa (Gallus gallus gallus, chickens), and ‘īlio (Canis familiaris, dogs), as well unintentional species introductions that came as stowaways on the voyaging canoes, including geckos (Gekkonidae), skinks (Scincidae), rats (Rattus exulans), and several other species (see above).
Early Hawaiian settlers also hunted birds for food (e.g., flightless geese, shearwaters and petrels) as well as for feathers (e.g., ‘ō‘ō (Moho species); mamo (Drepanis pacifica); and numerous other species). The bird feathers were prized for use in ‘ahu ‘ula (royal capes and cloaks), mahiole (feather-crested helmets), kāhili (royal feather standards) and other items of Hawaiian featherwork. (See Native Birds of the Hawaiian Islands, Chapter 7.) After 1778, hunting by foreigners brought new pressures on many native birds, such as various species of albatross whose eggs and feathers were extensively collected, and the ‘alalā (Corvus hawaiiensis, Hawaiian crows), which were systematically shot by farmers protecting their poultry yards and feed pens.
In the decades after Captain Cook’s first visit to the Hawaiian Islands, the landscape of the Islands began to be drastically altered by increasing numbers of cattle, sheep, goats, and pigs. The pigs brought by Westerners were significantly larger than those brought to the Hawaiian Islands by the first Polynesian settlers.
The Chinese market for sandalwood (Santalum species) also led to deforestation, particularly from 1810 to 1840, the peak years of the sandalwood trade. Increasing demands for wood and pasture eventually cleared huge areas of the Hawaiian landscape.
The loss of native forest habitat decimated many of the endemic Hawaiian bird species, particularly populations at lower elevations (where most of the alteration of the landscape took place). Due to this habitat loss as well as mosquito-borne diseases (e.g., avian malaria), the last surviving species of these small and endangered, endemic Hawaiian birds are now restricted to the higher elevations.
Threats to Native Species—Post-Contact
Since Captain Cook established Western contact with the Hawaiian Islands in 1778, thousands of species have been introduced to the Hawaiian Islands accidentally or intentionally, and for various purposes, including pest control and ornamental landscaping. Many escaped pets, such as cats (Felis domesticus), have also established significant wild populations in some Hawaiian forests. Various lizards have become established in some areas, including skinks, chameleons, geckos, anoles, and iguanas. (See Geckos and other Reptiles, Chapter 10.)
After Western contact with the Hawaiian Islands was established in 1778, the influx of new species was rapid, and had devastating effects on native species. These introduced species (also known as alien, exotic, adventive, or introduced species) included birds, lizards, plants, and animals that detrimentally affected native species of the Hawaiian Islands. Some introduced species destroyed native habitats while other preyed on native species or competed with native species for food. Sheep, goats, pigs, deer (axis and black-tailed), donkeys, and wallabies run wild in certain areas of the Hawaiian Islands. The wild donkeys are found on the island of Hawai‘i in the region north of the Kona airport near the Kona Village Resort and Four Seasons Hotel. The wild donkeys, known as the Kona Nightingales, are the remnants of a herd that was used to carry coffee from the uplands to the ocean until the end of World War II, when military jeeps began to be used to carry the coffee, and many of the donkeys were released on Hualālai Volcano. Axis deer are found on Moloka‘i, Maui, and Lāna‘i. Black-tailed deer are found in northwestern Kaua‘i, and feral donkeys are found on the island of Hawai‘i. Goats and pigs are now found on all of the Hawaiian Islands with the exception of Lāna‘i.
Ka i‘a holehole iwi o ka ‘āina. The fish of the land that strips the flesh from the bones. Goats. When one pursues them for meat, many a limb suffers skinning and bruises. (Pukui: 1337-145)
Two species of sheep may be found on the upper elevations of Mauna Kea Volcano on the island of Hawai‘i. One of these species, the Mouflon sheep, is also found on Lāna‘i. Recently, both sheep species have been nearly eliminated in an effort to protect native species.
Thousands of axis deer roam wild on the island of Maui, mostly from Hāna to Kapalua. These deer are the descendants of five deer that were introduced to Maui by State of Hawai‘i officials in 1959, and four more deer brought to Maui in 1960. The deer were initially brought to create new hunting opportunities. Maui’s axis deer population has increased rapidly over the last several decades (approximately doubling about every four years), with significant negative effects on native species and ecosystems of the Hawaiian Islands.
In 2002, the State of Hawai‘i signed a ten-year agreement to continue the availability of Mouflon sheep and axis deer hunting programs on Lāna‘i, where hunters take about 600 sheep and 700 deer each year. All of the above mentioned non-native feral ungulate species have had significant negative effects on native Hawaiian ecosystems as well as the native Hawaiian species they support.
An estimated 40 to 250 rock wallabies live in O‘ahu’s Kalihi Valley. Three of the miniature, kangaroo-like marsupials were brought to the Hawaiian Islands from Australia in 1916 to add to the collection of a private zoo. Two of the wallabies escaped, and today’s wild population in Kalihi Valley are their descendants.
A Summary of the Primary Threats to Native Species of the Hawaiian Islands
Historically, the primary threats to native species have included:
Ø The destruction of native habitat by hoofed animals (e.g., cattle, sheep, pigs, and goats).
Ø Predation on native species by non-native (introduced) rats, including the Polynesian black rat (Rattus exulans) that first came to the Hawaiian Islands as a stowaway on the Hawaiian voyaging canoes, and then two other rat species (Rattus rattus and Rattus norvegicus) brought by early European ships.
Ø Predation by the small Indian mongoose (Herpestesauropunctatus), which feeds on the eggs of ground-nesting native birds. Mongoose are now found on all of the main Hawaiian Islands except Kaua‘i.
Ø Predation on young, native waterbirds by various introduced species, including cattle egrets (Bubulcus ibis), and bullfrogs (Rana catesbeiana).
Ø Predation by non-native domestic (and feral) animals such as cats (Felis domesticus) and dogs (Canis familiaris).
Ø Destruction of native habitat by other non-native species with established breeding populations in the Hawaiian Islands, including axis deer, black-tailed deer, donkeys, wallabies, and Mouflon sheep (see above).
Ø Hunting of birds (by humans) for food, feathers, and for other reasons.
Ø Competition from aggressive non-native species, such as the Banana Poka vine (Passiflora mollissima) that blankets large areas of native forest and strangles native trees. Native Hawaiian plants now compete with more than 1,000 introduced plant species. Native bird species also must compete for habitat with introduced species, such as the barn owl (Tyto alba).
Ø Parasites and foreign diseases spread by non-native species (e.g., mosquitoes spreading avian malaria and avian pox to native forest birds). Many of the Hawaiian Islands’ native species are particularly vulnerable to foreign diseases.
Ø Introduced insects harmful to native species. For example, native kou trees (Cordia subcordata) were devastated by an introduced moth species.
Ø Loss of native habitat to agricultural enterprises, particularly sugarcane (Saccharum officinarum) and pineapple (Ananas comosus).
Ø The demand for fine woods, such as sandalwood (Santalum species) andkoa (Acacia koa), leading to the harvesting of native trees.
Ø Loss of habitat due to construction of homes and commercial developments.
Preventing Extinctions
The biological diversity of the Hawaiian Islands is astonishing: lush rainforests, underwater coral reefs, coastal dunes, subterranean lava tubes, and many other ecosystems support a variety of life forms. Unfortunately, many Hawaiian species cling tenuously to survival in remote and diminishing native habitat.
The combined efforts of scientists and concerned local citizens are helping to preserve native ecosystems and prevent extinctions. For example, there has been and continues to be a concerted community effort to rescue stranded wedge-tailed shearwater birds(Puffinus pacificus chlororhynchus). (See Newell’s and Wedge-Tailed Shearwaters section in Native Birds of the Hawaiian Islands, Chapter 7.) Other community volunteer efforts attempt to eradicate introduced plant species that threaten native ecosystems. One such event is the Banana Poka Festival in Kaua‘i’s Kōke‘e State Park.
Many native Hawaiian species are little known and even less seen. Scientists estimate that there are thousands of native Hawaiian species still unknown to science. [Graphic: Endangered Species Distribution]
Chapter 5 Source Notes
All Hawaiian words in this text, the Hawaiian Encyclopedia, include proper diacritical marks, including the ‘okina (hamzah, or glottal stop) and kahakō (macron), except when the word is cited directly as part of a title or in a direct quote, in which case the words are spelled as originally published. See Appendix 1: Note On Sources for more information about diacritical marks; objectivity; Hawaiian word spellings and meanings; scientific, cultural, and historic information; new research and discoveries; verifying dates and details of historic events; and Hawaiian sovereignty and other native issues.
[Note: This list is still in progress and being updated.] Abbott, Isabella Aiona. Lā‘au Hawai‘i: Traditional Hawaiian Uses of Plants. Honolulu: Bishop Museum Press, 1992. Abbott, Isabella Aiona. Limu: An Ethnobotanical Study of Some Hawaiian Seaweeds. Lawai: National Tropical Botanical Garden, 1984. All About Hawaii (1948-1949): Thrum’s Hawaiian Annual and Standard Guide. Publications Division: Honolulu Star-Bulletin, Honolulu, Hawaii, 1949. Ariyoshi, Rita. The National Geographic Traveler: Hawaii. Washington, D.C.: National Geographic Society, 2000. Atlas of Hawai‘i: Second Edition. Honolulu: University of Hawai‘i Press, 1983. Atlas of Hawai‘i: Third Edition. Honolulu: University of Hawai‘i Press, 1998. Balcomb, Kenneth C. III. The Whales of Hawaii: Including All Species of Marine Mammals in Hawaiian and Adjacent Waters. San Francisco, California: Marine Mammal Fund, 1987. Beletsky, Les. Hawaii: The Ecotravellers’ Wildlife Guide. London: Academic Press, 2000. Berger, Andrew J. Bird Life in Hawaii. Honolulu: Island Heritage Limited, 1983. Bishop Museum: Hawaii’s Extinct Species. Internet site: http://hbs.bishopmuseum.org/endangered/extinct.html, 7/28/2002. Bishop Museum—Hawaii’s Extinct Species: Insects. Internet site: http://hbs.bishopmuseum.org/endangered/ext-insects.html, 7/28/2002. Bishop Museum—Hawaii’s Extinct Species: Plants. Internet site: http://hbs.bishopmuseum.org/endangered/ext-plants.html, 7/28/2002. Bishop Museum—Hawaii’s Extinct Species: Snails. Internet site: http://hbs.bishopmuseum.org/endangered/ext-snails.html, 7/28/2002. Bishop Museum: Native plant species in Hawai‘i. Internet site: http://www.bishopmuseum.org/greenwell/gwplants.html, 8/07/2002. Bishop Museum: The How Many Species Are there in Hawaii Web Page. Includes: How many species are there in Hawaii?, from the “Records of the Hawaii Biological Survey for 1994” as published in the Bishop Museum Occasional Paper volume 41: 3-18; and Number of Hawaiian Species: Supplement 1, originally published in Bishop Museum Occasional Papers 45: 8-17 (1996). Internet site: http://hbs.bishopmuseum.org/hispp.html, 10/17/2002. Bishop Museum—The role of the Hawaii Biological Survey. Internet site: http://hbs.bishopmuseum.org/hbs3.html. Boynton, David. Boar hunting: Hunting the feral pigs helps to protect the forest ecology. Kauai Magazine, Fall, 1992. Cabrera, Theresa. The Hawaiian hoary bat: Hawaii’s only native land mammal. Honolulu: Nature Conservancy of Hawaii Newsletter, Spring, 1995. Cave critters added to endangered species list. The Garden Island, 1/16/2000. Chang, Lester. Visit to “sinkhole” offers a look at thousands of years of Kaua‘i’s natural history. The Garden Island, 3/25/2002. Chang, Lester. $2.8 million helping bats, other species: Federal protection money pouring into state. The Garden Island, 9/29/2001. Clark, Hugh. Turtle nest in Hilo great rarity in itself. The Honolulu Advertiser, 10/12/2002. Culliney, John L. Islands in a Far Sea. San Francisco: Sierra Club Books, 1988. Daws, Gavan. Shoal of Time: A History of the Hawaiian Islands. Honolulu: University of Hawaii Press, 1968. Degener, Otto. Plants of Hawaii National Parks: Illustrative of Plants and Customs of the South Seas. Braun-Brumfield, Inc., 1973. Denny, Jim. The Birds of Kaua‘i. Honolulu: University of Hawai‘i Press, 1999. Endangered animals in the Hawaiian Islands: Hawaiian Bat. Internet site produced by the US Fish and Wildlife Service: http://www.r1.fws.gov/pacific/wesa/hrybatindex.html. Eyre, David L. By Wind, By Wave. Honolulu: The Bess Press, 2000. Federal Register, 2002, Vol. 67, U.S. Fish & Wildlife Service, Threatened and Endangered Species System (TESS), Listings by State and Territory as of 7/20/2002. Internet site: http://ecos.fws.gov/servlet/TESSWebpageUsaLists?state=HI, 7/28/2002. Fish breakthrough good news for Hawai‘i. The Honolulu Advertiser, 1/24/2002. Gulko, David. Hawaiian Coral Reef Ecology. Honolulu, Hawaii: Mutual Publishing, 1998. Handy, Emory, Bryan, Buck, Wise and Others. Ancient Hawaiian Civilization: A Series of Lectures Delivered at the Kamehameha Schools. Mutual Publishing: Honolulu, Hawaii, 1999. Handy, E.S. Craighill, and Handy, Elizabeth Green, with the collaboration of Mary Kawena Pukui. Native Planters in Old Hawaii: Their Life, Lore, and Environment. Honolulu: Bishop Museum Press, 1972. Hargreaves, Dorothy and Bob. Tropical Blossoms of the Pacific. Hargreaves Company, Inc., 1970. Hargreaves, Dorothy and Bob. Tropical Trees of the Pacific. Hargreaves Company, Inc., 1970. Hawaiian and Pacific Islands National Wildlife Refuges. United States Department of the Interior Fish and Wildlife Service, September, 1985. Hawaiian Cave Animals: Threatened and Endangered Species; Endangered Animals in the Hawaiian Islands. U.S. Fish and Wildlife Service. Internet site: http://www.r1.fws.gov/pacific/wesa/caveanimals.html, 6/17/2000. Hawaiian Native Plant Genera. University of Hawai‘i Botany Department. Internet site: http://www.botany.hawaii.edu/faculty/carr/natives.htm, 5/03/2002. Hawai‘i Data Book: A Statistical Reference to Hawai‘i’s Social, Economic and Political Trends. Edited by Robert C. Schmitt. Honolulu: Mutual Publishing, 2002. Hawaii’s Birds. Honolulu: Hawaiian Audubon Society, 1993. Hayworth, Philip. Hawai‘i’s Marine Zoos: The Waikīkī Aquarium, Sea Life Park and Maui Ocean Center showcase the incredible diversity of marine life flourishing in Hawaiian waters. Spirit of Aloha Magazine, May/June, 2002. Hobson, Edmund, and Chave, E.H. Hawaiian Reef Animals. University of Hawaii Press, 1972. Hoover, John P. A Pocket Guide to Hawai‘i’s Underwater Paradise. Honolulu: Mutual Publishing, 1997. Hoover, John P. Hawaii’s Fishes: A Guide for Snorkelers, Divers and Aquarists. Honolulu: Mutual Publishing, 1993. Hoover, John P. Hawai‘i’s Sea Creatures: A Guide to Hawai‘i’s Marine Invertebrates. Honolulu: Mutual Publishing, 1998. Howarth, F.G. and Mull W.P. Hawaiian Insects and their Kin. Honolulu: University of Hawai‘i Press, 1992. Hurley, Timothy. Another nastier lizard found. The Honolulu Advertiser, 3/19/2002. Hurley, Timothy. Axis deer a growing nuisance on Maui: Report urges containment. The Honolulu Advertiser, 8/20/2002. Hurley, Timothy. Death revives donkey debate. The Honolulu Advertiser, 10/06/2002. Hurley, Timothy. Poison frog found on Maui. The Honolulu Advertiser, 3/29/2002. Hurley, Timothy. State extends Lāna‘i hunting. The Honolulu Advertiser, 7/05/2002. Ii, John Papa. Fragments of Hawaiian History. Translated by Mary Kawena Pukui; Edited by Dorothy B. Barrère. Honolulu: Bishop Museum Press, 1959. Joesting, Edward. Kauai: The Separate Kingdom. Hawaii: University of Hawaii Press, 1984. Kamakau, Samuel Manaiakalani. Ka Po‘e Kahiko: The People of Old. Translated from the Newspaper Ke Au ‘Oko‘a by Mary Kawena Pukui. Arranged and edited by Dorothy B. Barrère. Bishop Museum Special Publication 51. Honolulu: Bishop Museum Press, 1991. Kāne, Herb Kawainui. Ancient Hawai‘i. Captain Cook, Hawai‘i: The Kawainui Press, 1997. Kaufman, Gregory Dean, and Forestell, Paul Henry. Hawaii’s Humpback Whales. Honolulu: Island Heritage Publishing, 1986. Kawaharada, Dennis. The Settlement of Polynesia, Part 1. Polynesian Voyaging Society. Internet Site: http://leahi.kcc.hawaii.edu/org/pvs/mirgrationspart1.html. Kepler, Angela Kay. Hawai‘i’s Floral Splendor: A Friendly and Color Identification Guide to Native and Introduced Flowers of All the Hawaiian Islands. Honolulu: Mutual Publishing, 1997. Kilauea Point Natural History Association. Hawaii’s Wetlands. Kauai: Hawai‘i Nature Focus: Nature Studies for Children, # 11. Kirch, Patrick Vinton. Feathered Gods and Fishhooks. Honolulu: University of Hawai‘i Press, 1985. Kirch, Patrick V., and Sahlins, Marshall. Anahulu: The Anthropology of History in the Kingdom of Hawaii. Volume One: Historical Ethnography (Marshall Sahlins, with the Assistance of Dorothy B. Barrère). Chicago and London: The University of Chicago Press, 1992. Kirch, Patrick V., and Sahlins, Marshall. Anahulu: The Anthropology of History in the Kingdom of Hawaii. Volume Two: The Archaeology of History (Patrick V. Kirch). Chicago and London: The University of Chicago Press, 1992. Krauss, Beatrice H. Plants in Hawaiian Culture. Honolulu: University of Hawaii Press, 1993. Krauss, Beatrice H. Plants in Hawaiian Medicine. Honolulu: The Bess Press, 2001. Life in Early Hawai‘i: The Ahupua‘a—Third Edition. Honolulu: Kamehameha Schools Press, 1994. Lum, Curtis. Iguanas making themselves at home in O‘ahu communities. The Honolulu Advertiser, 4/09/2002. Malo, David. Hawaiian Antiquities: Mo‘olelo Hawai‘i. Translated from the Hawaiian by Dr. Nathaniel B. Emerson, 1898. Bernice P. Bishop Museum Special Publication 2: Second Edition. Honolulu: Bishop Museum, 1951. Manguchei, Kendyce. Poor man’s time machine uncovers Kaua‘i’s past at Maha‘ulepu. The Garden Island, 6/23/2002. Matisoo-Smith, E., Roberts, R.M., Irwin, G.J., Allen, J.S., Penny, D. and Lambert, D.M. Patterns of prehistoric human mobility in Polynesia indicated by mtDNA from the Pacific rat. Anthropology, Vol. 95, pp. 15145-15150, December, 1998. McKeown, Sean. A Field Guide to Reptiles and Amphibians in the Hawaiian Islands. California: Diamond Head Publishing, Inc., 1996. Mee, Jonathan. Living lava lamps. Oregon Coast, July/August, 1995. Meinkoth, Norman A. The Audubon Society Field Guide to North American Seashore Creatures. New York: Alfred A. Knopf, 1981. Memories of Duke Kahanamoku. Internet site: http://planet-hawaii.com/duke/memories.htm Merlin, Mark. Hawaiian Coastal Plants: An Illustrated Field Guide. Honolulu: Pacific Guide Books, 1999. Merlin, Mark David. Hawaiian Forest Plants: A Hiker’s Guide. Honolulu: The Oriental Publishing Company, 1977. Merriam Webster’s Collegiate Dictionary: Tenth Edition. Springfield, Massachusetts: Merriam-Webster, Incorporated, 1996. Mitchell, Donald D. Kilolani. Resource Units in Hawaiian Culture: Revised Edition. Honolulu: Kamehameha Schools Press, 1992. Miyano, Leland, and Peebles, Douglas. A Pocket Guide to Hawaii’s Flowers. Honolulu Hawaii: Mutual Publishing, 1997. Miyano, Leland. Hawai‘i’s Beautiful Trees. Honolulu: Mutual Publishing, 1997. Moriarty, Linda Paik. Pūpū ‘o Ni‘ihau: Shell Lei of Ni‘ihau. Honolulu: Honolulu Academy of Arts, 2001. Mullins, Joseph G. Hawaiian Journey. Honolulu: Mutual Publishing, 1978. New lizard species established on Oahu. The Garden Island, 2/12/2001. Oliver, Anthony Michael. Hawaii Fact and Reference Book. Mutual Publishing, 1995. Oliver, Douglas. Polynesia in Early Historic Times. Honolulu, Hawai‘i: The Bess Press, 2002. Payne, Roger. Among Whales. New York: Charles Scribner’s Sons: Simon and Schuster Inc., 1995. Polhemus, Dan, and Asquith, Adam. Hawaiian Damselflies: A Field Identification Guide. Honolulu: Bishop Museum Press, 1996. Pratt, H. Douglas. A Pocket Guide to Hawai‘i’s Birds. Honolulu: Mutual Publishing, 1996. Pratt, H. Douglas. Enjoying Birds in Hawaii: A Birdfinding Guide to the Fiftieth State. Honolulu: Mutual Publishing, 1993. Pukui, Mary Kawena, and Elbert, Samuel H. Hawaiian Dictionary: Revised and Enlarged Edition. Honolulu: University of Hawaii Press, 1986. Pukui, Mary Kawena, and Elbert, Samuel H, with Mookini, Esther T. and Nishizawa, Yu Mapuana. New Pocket Hawaiian Dictionary. Honolulu: University of Hawaii Press, 1992. Pukui, Mary Kawena. ‘Ōlelo No‘eau: Hawaiian Proverbs & Poetical Sayings. Honolulu: Bishop Museum Press, 1983. Pukui, Mary Kawena, and Elbert, Samuel H. & Mookini, Esther T. Place Names of Hawaii: Revised & Expanded Edition. Honolulu: University of Hawaii Press, 1974. Randall, John E. Shore Fishes of Hawai‘i. Honolulu: University of Hawai‘i Press, 1996. Randall, John E. Underwater Guide to Hawaiian Reef Fishes. Pennsylvania: Harrowood Books, 1981. Rare turtle lays eggs on Hilo beach. The Garden Island, 10/12/2002. Rock Wallaby released in Kalihi Valley. The Garden Island, 6/17/2002. Rogers, Michael. Little Birds Lost. National Wildlife, October/November, 1993. Ronck, Ronn. Hawaii Almanac. Honolulu: University of Hawaii Press, 1984. Roth, Susan. Hawai‘i expert testifies for House bill on invasive species: Administration says effort not a priority. The Honolulu Advertiser, 3/15/2002. Royte, Elizabeth. On the brink: Hawaii’s vanishing species. National Geographic Magazine, September, 1995. Russo, Ron. Hawaiian Reefs: A Natural History Guide. Wavecrest Publications, 1994. Sabelli, Bruno. Simon & Schuster’s Guide to Shells. Edited by Harold S. Feinberg. New York: Simon & Schuster Inc., 1979. Sahlins, Marshall. Anahulu: The Anthropology of History in the Kingdom of Hawai‘i. Chicago: The University of Chicago Press, 1992. Schmid, Randolph E. Pacific states vulnerable to giant sea waves. The Garden Island, 7/29/1998. Scientists unstick the gecko mystery: Foot hairs pave way for ‘glue’. The Honolulu Advertiser (from Los Angeles Times), 6/19/2000. Scott, Susan. Plants and Animals of Hawai‘i. Honolulu: Bess Press, 1991. Scott, Susan and Thomas, Craig, M.D. Poisonous Plants of Paradise: First Aid and Medical Treatment of Injuries from Hawai‘i’s Plants. Honolulu: University of Hawai‘i Press, 2000. Seven ‘alala chicks released into wild. ‘Elepaio: Journal of the Hawaiian Audubon Society, Volume 55, #1, February, 1995. Silverstein, Duane. Our tribe must make all islands survivors. The Honolulu Advertiser, 3/24/2002. Sprague, Brandon. In pursuit of Kauai’s dwindling native bird population. The Garden Island. (no date) TenBruggencate, Jan. Aliens overrun Island environs. The Honolulu Advertiser, 7/09/2001. TenBruggencate, Jan. Beetles live here in abundance. The Honolulu Advertiser, 6/02/2003. TenBruggencate, Jan. Botanical secrets of sea await discovery. The Honolulu Advertiser, 7/3/2000. TenBruggencate, Jan. Kaua‘i limestone 315,000 years old. The Honolulu Advertiser, 2/19/2002. TenBruggencate, Jan. Last habitat for crow endangered. The Honolulu Advertiser, 2/15/1996. TenBruggencate, Jan. Loss of variability left nēnē vulnerable. The Honolulu Advertiser, 2002. TenBruggencate, Jan. Nene flourish on Kauai. The Honolulu Advertiser, 7/19/1995. TenBruggencate, Jan. Science mission finds new species: Discoveries rife on outlying isles. The Honolulu Advertiser, 10/19/2000. TenBruggencate, Jan. Sinkhole yields treasure trove of bird fossils. The Honolulu Advertiser, 2/25/2002. TenBruggencate, Jan. Variety fills island ecosystems. The Honolulu Advertiser, 9/16/2002. The Audubon Society Field Guide to North American Fishes, Whales, and Dolphins.Knopf, Alfred A., 1983. The Plant Book: The World of Plants in a Single Volume. Australia: James Mills-Hicks, Random House Australia Pty Ltd., 2001. Thompson, Rod. Endangered olive ridley turtle lays eggs at Hilo Bay: Officials are asking the public to not disturb the rare nest, which has 124 eggs. The Honolulu Star-Bulletin, 10/12/2002. United States Fish & Wildlife Service: Pacific Islands—Endangered Species; Threatened and Endangered Animals in the Hawaiian Islands. Hawaiian Cave Animals: Kaua‘i Cave Wolf Spider and Kaua‘i Cave Amphipod. Internet site: http://pacificislands.fws.gov/wesa/caveanimals.html, 7/24/2002. U.S. Fish & Wildlife Service, Threatened and Endangered Species System (TESS): Listings by State and Territory, as of 7/28/2002. Internet site: http://ecosfws.gov/webpage/webpage_usa_lists.html?state=all&module=undefined, 7/28/2002. U.S. Fish & Wildlife Service, Threatened and Endangered Species System (TESS): U.S. Listed Vertebrate Animal Species Report by Taxonomic Group as of 7/28/2002. Internet site: http://ecos.fws.gov/webpage/webpage_vip_listed.html?module=undefined&code=V&listings=0, 7/28/2002. Valier, Kathy. Ferns of Hawai‘i. Honolulu, University of Hawai‘i Press, 1995. Viotti, Vicki. The lei of the land. The Honolulu Advertiser, 4/16/1995. Wagner, Warren L., and Herbst, Derral R. Electronic Supplement to the Manual of the Flowering Plants of Hawai‘i. Internet site:http://rathbun.si.edu/botany/pacificislandbiodiversity/hawaiianflora/supplement.htm, 3/05/2002. Wagner, Warren L., Herbst, Derral R., and Sohmer, S.H. Manual of the Flowering Plants of Hawai‘i: Revised Edition, Volume 1. Honolulu: University of Hawai‘i Press; Bishop Museum Press, 1999. Wagner, Warren L., Herbst, Derral R., and Sohmer, S.H. Manual of the Flowering Plants of Hawai‘i: Revised Edition, Volume 2. Honolulu: University of Hawai‘i Press; Bishop Museum Press, 1999. Witte, Astrid and Mahaney, Casey. Hawaiian Reef Fish. Waipahu, Hawai‘i: Island Heritage Publishing, 1998. Yamamoto, Mike N., and Tagawa, Annette W. Hawai‘i’s Native & Exotic Freshwater Animals. Honolulu: Mutual Publishing, 2000.
[1]Juvik, James O. Biogeography in Atlas of Hawai‘i: Third Edition. Edited by Sonia P. Juvik and James O. Juvik. Honolulu: University of Hawai‘i Press, 1998. [2] p. 276, Pukui, Mary Kawena. ‘Ōlelo No‘eau: Hawaiian Proverbs & Poetical Sayings. Honolulu: Bishop Museum Press, 1983. Proverb 2529. [iii] Conant, Sheila. Extinct Birds, in Atlas of Hawai‘i: Third Edition. Edited by Sonia P. Juvik and James O. Juvik. Honolulu: University of Hawai‘i Press, 1998. [iv]BishopMuseum: Hawai‘i’s Extinct Species—Birds. Internet site: http://hbs.bishopmuseum.org/endangered/ext-birds.html, 7/28/2002. [v] Burney, David A., James, Helen F., Burney, Lida Pigott, Olson, Storrs L., Kikuchi, William, Wagner, Warren L., Burney, Mara, McCloskey, Deirdre, Kikuchi, Delores, Grady, Frederick V., Gage II, Reginald, and Nishek, Robert. Fossil evidence for a diverse biota from Kaua‘i and its transformation since human arrival. Ecological Monographs, 71 (4), 2001, pp. 615-641. [vi] Burney, David A., James, Helen F., Burney, Lida Pigott, Olson, Storrs L., Kikuchi, William, Wagner, Warren L., Burney, Mara, McCloskey, Deirdre, Kikuchi, Delores, Grady, Frederick V., Gage II, Reginald, and Nishek, Robert. Fossil evidence for a diverse biota from Kaua‘i and its transformation since human arrival. Ecological Monographs, 71 (4), 2001, pp. 615-641. [vii] Burney, David A., James, Helen F., Burney, Lida Pigott, Olson, Storrs L., Kikuchi, William, Wagner, Warren L., Burney, Mara, McCloskey, Deirdre, Kikuchi, Delores, Grady, Frederick V., Gage II, Reginald, and Nishek, Robert. Fossil evidence for a diverse biota from Kaua‘i and its transformation since human arrival. Ecological Monographs, 71 (4), 2001, pp. 615-641. [viii] Evenhuis, Neal L., and Eldredge, Lucius G., Editors. Records of the Hawaii Biological Survey for 2000. Bishop Museum Occasional Papers, Number 68, 69. Honolulu: Bishop Museum, 3/25/2002. [ix]The How Many Species Are there in Hawaii Web Page. Includes: How many species are there in Hawaii?, from the “Records of the Hawaii Biological Survey for 1994” as published in the Bishop Museum Occasional Paper volume 41: 3-18; and Number of Hawaiian Species: Supplement 1, originally published in Bishop Museum Occasional Papers 45: 8-17 (1996). Internet site: http://hbs.bishopmuseum.org/hispp.html, 10/17/2002. [x] Evenhuis, Neal L., and Eldredge, Lucius G., Editors. Records of the Hawaii Biological Survey for 2000. Bishop Museum Occasional Papers, Number 68, 69. Honolulu: Bishop Museum, 3/25/2002. [xi]The Role of the Hawaii Biological Survey. Bishop Museum Internet site: http://hbs.bishopmuseum.org/hbs3.html, 8/072002. [xii] Evenhuis, Neal L., and Eldredge, Lucius G., Editors. Records of the Hawaii Biological Survey for 2000. Bishop Museum Occasional Papers, Number 68, 69. Honolulu: Bishop Museum, 3/25/2002. [xiii] Warshauer, F.R. Alien Species and Threats to Native Ecology, in the Atlas of Hawai‘i: Third Edition. Edited by Sonia P. Juvik and James O. Juvik. Honolulu: University of Hawai‘i Press, 1998. [xiv] Howarth, F.G., and Mull, W.P. Hawaiian Insects and their Kin. Honolulu: University of Hawai‘i Press, 1992. [xv] Pukui, Mary Kawena, and Elbert, Samuel H. Hawaiian Dictionary: Revised and Enlarged Edition. Honolulu: University of Hawaii Press, 1986. [xvi] Hadfield, Michael G. Hawaiian Tree Snails, in the Atlas of Hawai‘i: Third Edition. Edited by Sonia P. Juvik and James O. Juvik. Honolulu: University of Hawai‘i Press, 1998. [xvii] Yamamoto, Mike N., and Tagawa, Annette W. Hawai‘i’s Native & Exotic Freshwater Animals. Honolulu: Mutual Publishing, 2000. [xviii] p. 144, Pukui, Mary Kawena. ‘Ōlelo No‘eau: Hawaiian Proverbs & Poetical Sayings. Honolulu: Bishop Museum Press, 1983. Proverb 1323. [xix] Includes viper shark (Trigonognathus kabeyai) reported in: Evenhuis, Neal L., and Eldredge, Lucius G., Editors. Records of the Hawaii Biological Survey for 2000. Bishop Museum Occasional Papers, Number 68, 69. Honolulu: Bishop Museum, 3/25/2002. [xx] Wagner, Warren L., Herbst, Derral R., and Sohmer, S.H. Manual of the Flowering Plants of Hawai‘i: Revised Edition, Volumes 1 and 2. Honolulu: University of Hawai‘i Press; Bishop Museum Press, 1999. [xxi] Wagner, Warren L., and Herbst, Derral R. Electronic Supplement to the Manual of the Flowering Plants of Hawai‘i. Internet site:http://rathbun.si.edu/botany/pacificislandbiodiversity/hawaiianflora/supplement.htm, 3/05/2002. [xxii] Kirch, Patrick V. (Citing Cooke 1926; Cooke and Kondo 1960; Pilsbry 1916-1918; Solem 1959.) The Impact of the Prehistoric Polynesians on the Hawaiian Ecosystem. Pacific Science, Vol.36, No.1, January, 1982. [xxiii] Kirch, Patrick V. The Impact of the Prehistoric Polynesians on the Hawaiian Ecosystem. Pacific Science, Vol.36, No.1, January, 1982. [xxiv] Evenhuis, Neal L., and Eldredge, Lucius G., Editors. Records of the Hawaii Biological Survey for 2000. Bishop Museum Occasional Papers, Number 68, 69. Honolulu: Bishop Museum, 3/25/2002. [xxv]Federal Register, 2002, Vol. 67, U.S. Fish & Wildlife Service, Threatened and Endangered Species System (TESS), Listings by State and Territory as of 7/20/2002. Internet site: http://ecos.fws.gov/servlet/TESSWebpageUsaLists?state=HI, 7/28/2002. [xxvi]BishopMuseum: Hawai‘i’s Extinct Species—Birds. Internet site: http://hbs.bishopmuseum.org/endangered/ext-birds.html, 7/28/2002. [xxvii]BishopMuseum - Hawaii’s Extinct Species. Internet site: http://hbs.bishopmuseum.org/endangered/extinct.html, 7/28/2002. [xxviii]BishopMuseum - Hawaii’s Extinct Species: Insects. Internet site: http://hbs.bishopmuseum.org/endangered/ext-insects.html, 7/28/2002. [xxix] Wagner, Warren L., Herbst, Derral R., and Sohmer, S.H. Manual of the Flowering Plants of Hawai‘i: Revised Edition, Volumes 1 and 2. Honolulu: University of Hawai‘i Press; Bishop Museum Press, 1999.
Life in the Islands, Evolutionary Processes, and Threats to Native Species
[Illustration: Native Bird, Fish, and Sea Turtle]
Part 1: Evolutionary Processes in the Hawaiian Islands
Colonization—Native and Endemic Species
Methods of Arrival—How Native Species Traveled to the Hawaiian Islands
Endemism and Adaptive Radiation
Part 2: Native and Polynesian-Introduced Species of the Hawaiian Islands
Past Extinctions—Bird Species
Native Hawaiian Birds
Bishop Museum’s Hawaii Biological Survey
Arthropods
Snails and Other Invertebrates
Reptiles and Amphibians
Fish
Native Reef Fish
Sharks, Rays and Eels
Other Marine Species
Land and Sea Mammals
Native and Polynesian-Introduced Plants
Other Polynesian-Introduced Species
Intentional Polynesian Introductions
Unintentional Polynesian Introductions
Geckos
Skinks
Plants
Snails
Other Polynesian-Introduced Species
Part 3: Threats to Native Species
Endangered Species and Extinctions
Threats to Native Species—Before 1778
Threats to Native Species—Post-Contact
A Summary of the Primary Threats to Native Species of the Hawaiian Islands
Preventing Extinctions
Part 1: Evolutionary Processes in the Hawaiian Islands
[Illustration: Hawaiian Islands, Frigatebird]
Key to Species Classifications:
Native—Arrived in the Hawaiian Islands without the aid of humans (indigenous), or evolved in the Hawaiian Islands (endemic).
Indigenous—Native to the Hawaiian Islands and other places.
Endemic—Evolved in the Hawaiian Islands from an indigenous species; native to the Hawaiian Islands and nowhere else.
Polynesian Introduction—Brought to the Hawaiian Islands by the early Polynesian settlers.
Post-Contact Introduction—Brought to the Hawaiian Islands after Western contact (1778).
Naturalized—Not native to the Hawaiian Islands, but now growing wild in the Hawaiian Islands.
After the volcanoes of the Hawaiian Islands formed and cooled, the weathering of the sun, rain, and wind began to break down the lava rock into soil. Primitive plants known as lichens also helped to create soil, allowing more plant species to take hold. These plant species in turn created new habitats that allowed the Hawaiian Islands’ original colonizing species to evolve, over millions of years, into thousands of plant, insect, and bird species.
Relatively few species were able to reach the Hawaiian archipelago before humans arrived, but those species that did become established were able to take advantage of the Hawaiian Islands’ diversity of habitats and ecosystems. The original colonizing species evolved into an array of uniquely adapted species that gradually transformed the Hawaiian Islands barren and rocky volcanoes into lush tropical islands full of life.
Colonization—Native and Endemic Species
Located more than 2,400 miles (3,862 km) from the nearest continental land mass, the Hawaiian Islands were not easily colonized by new species prior to the arrival of humans. The extreme geographical isolation of the Hawaiian Islands also limited the types and the numbers of species that became established.
All of the Hawaiian Islands’ native species are divided into two categories: indigenous and endemic. Species that somehow made it to the Hawaiian Islands without the aid of humans, and then were able to establish native populations, are now classified as indigenous to the Hawaiian Islands. These indigenous species are native to the Hawaiian Islands, and are also native to other places (wherever the species originally came from).
The other category of native species is the endemic Hawaiian species, which evolved in the Hawaiian Islands from species that were already established in (and thus native to) the Hawaiian Islands. Endemic Hawaiian species may have evolved from an indigenous Hawaiian species or from another endemic Hawaiian species. In either case, all endemic Hawaiian species evolved in the Hawaiian Islands, and thus are unique to the Hawaiian Islands (found naturally nowhere else).
Hawai‘i’s geographic isolation and diversity of habitats and food sources led to the evolution of many new (endemic) species from the relatively few colonizing species that were able to make it to the Hawaiian Islands and reproduce. This resulted in extremely high rates of endemism for various categories of species. For example, about 79% of the native flowering plant species of the Hawaiian Islands are endemic, along with about 70% of native ferns of the Hawaiian Islands. About 25% of native Hawaiian reef fish are endemic to the Hawaiian Islands.
Methods of Arrival—How Native Species Traveled to the Hawaiian Islands
Indigenous species inhabiting the Hawaiian Islands arrived in the Islands by a variety of methods. For example, seeds floated to the Hawaiian Islands on the ocean, or were carried in the wing feathers or in the digestive tracts of birds, or in mud on the birds’ feet. The colonizing bird species may have come to the Hawaiian Islands on their natural migration tracks, or arrived after being blown off course by a storm.
Snails and other species may have been transported to the Hawaiian Islands by hitching a ride on floating logs or on rafts of debris that floated over the Pacific Ocean and washed up on Hawaiian shores. Tiny snail eggs were also carried to the Hawaiian Islands embedded in mud in the feet of birds.
Insects and spiders in far distant places were lifted by winds high into the jetstream, more than 8 miles (13 km) up, where they were carried out over the Pacific Ocean at more than 120 miles per hour (193 km/hr) and then set down on the Hawaiian Islands, there to remain and multiply.
For many species, the journey to the Hawaiian Islands was too long and the odds too great. Even over many millions of years the species were never able to reach the Hawaiian Islands, or if they did, there was not suitable habitat for them to survive. Some species may have arrived in the Hawaiian Islands in such small numbers that they were not able to establish breeding (reproducing) populations.
It is estimated that about 39% of plant species in the Hawaiian Islands came in the stomachs of birds; about 13% arrived in mud on birds’ feet; about 23% arrived after being stuck to the feathers of birds; about 9% arrived on rafts of seaweed or debris on the ocean; and about 1.4% came by floating through the air to the Hawaiian Islands.[1]
Many plants and animals that are very common in other places never became established in the Hawaiian Islands until humans brought the species to the Hawaiian Islands. For example, in the Hawaiian Islands there are no gymnosperms (e.g., pine trees), no native amphibians, no native terrestrial reptiles, and no native ants. However, all of these organisms are now found in the Hawaiian Islands, having been introduced to the Islands by humans.
Endemism and Adaptive Radiation
Species that somehow were able to traverse the vast distance over the Pacific Ocean to the Hawaiian Islands and become established in the Hawaiian Islands (without the aid of humans) were then able to adapt to a wide variety of environmental conditions (ecological niches) found in the Hawaiian Islands. Populations of species often became divided, with each separated sub-population adapting to different habitats and food sources, and eventually evolving into completely new species.
For example, one finch species that established itself in the Hawaiian Islands millions of years ago evolved into at least 50 species and subspecies of Hawaiian honeycreepers. The evolutionary process whereby one species evolves into many new species is known as adaptive radiation. The Hawaiian Islands are renowned for their historical record of adaptive radiation, particularly regarding bird and insect species.
Over hundreds of generations, many plant, animal, and insect species evolved adaptations to the unique conditions found on each of the Hawaiian Islands. Many of these adaptations distinguished the separate populations, which eventually evolved into unique species or subspecies, endemic to each particular island. For example, the fragrant mokihana plant (Pelea anisata) is endemic to the uplands of Kaua‘i, as is the puaiohi (Myadestes palmeri), a tiny pink-legged forest bird.
Some native Hawaiian species may be endemic to very small areas. For example, the endangered plant known as koki‘o ke‘oke‘o (Hibiscus waimeae subspecies hannerae, white hibiscus) is found only in a few northwest Kaua‘i valleys, including the valleys of Limahuli and Hanakāpī‘ai.
Native Hawaiian species isolated within a particular valley for many thousands of years often developed variations that aided their survival in the particular habitat. This isolation often led to distinct subpopulations of species that eventually became distinguishable as endemic subspecies or completely new species. An ancient proverb states, “‘O‘opu peke o Hanakāpī‘ai.” (“The stunted ‘o‘opu fish of Hanakāpī‘ai,” referring to “...the ‘o‘opu of Hanakāpī‘ai, which are said to be plump and shorter in length than those elsewhere. Sometimes applied humorously to a short, plump person.”[2]
The evolutionary process of adaptive radiation allowed the relatively few colonizers of the Hawaiian Islands to develop into a vast array of life forms that today constitutes the diverse native flora and fauna of the Hawaiian Islands. (For a more in-depth explanation of adaptive radiation, see this book’s Introduction as well as the Honeycreepers and Honeyeaters section in Native Birds of the Hawaiian Islands, Chapter 7.)
Part 2—Native and Polynesian-Introduced Species of the Hawaiian Islands
Past Extinctions—Bird Species
Worldwide there are about 9,700 bird species, but only about 2% of these bird species are native to the Hawaiian Islands. When Captain Cook arrived in the Hawaiian Islands in 1778, there were about 80 species of endemic Hawaiian landbirds, along with at least 24 species and subspecies of native Hawaiian seabirds (4 endemic) and 34 species and two subspecies of waterbirds (29 endemic). More than 35 endemic Hawaiian landbirds had already gone extinct by 1778 when Cook’s arrival established Western contact with the Hawaiian Islands.
Bird species that were extinct prior to Cook’s arrival met their demise from a variety of causes. The Polynesians settlers of the Hawaiian Islands hunted birds for food and for feathers, though sometimes the birds were released unharmed after some of the feathers were collected.(See Native Birds of the Hawaiian Islands, Chapter 7.)
Pua‘a (Sus scrofa, pigs) and ‘īlio (Canis familiaris, dogs) brought by the Polynesians also had an effect on island bird populations, as did ‘iole (Rattus exulans, Polynesian black rats) that hitched a ride on the voyaging canoes. These pigs, dogs, and rats were particularly detrimental to the Hawaiian Islands’ ground-nesting birds, which were also the most likely birds to be hunted by humans for food.
Aia a pohā ka leo o ka ‘a‘o, kāpule ke momona o ka ‘uwa‘u i ka puapua.
When the ‘a‘o birds’ voices are distinctly heard, the ‘uwa‘u birds
are fat even to the very tails.
The ‘a‘o bird was not heard during the nesting season. When the fledglings emerged and their cries were heard, the season had come when young ‘uwa‘u were best for eating, and the people went to snare them.
(Pukui: 32-6)
About 23 of the bird species that became extinct prior to Western contact were ground-nesting birds. Known only from subfossil discoveries, these species included flightless ducks (four species); flightless ibises (three species); flightless rails (at least 10 species); and flightless geese (six species). Some of the flightless geese species were more than 3 feet (.9 m) tall.[iii]
Other species that went extinct prior to Western contact included at least four long-legged owl species, one shearwater, one bald eagle, two crow species, one petrel, a harrier, and more than 21 honeycreeper species along with one honeyeater species. It is possible that some of these bird species may have become extinct prior to Polynesian settlement in the Hawaiian Islands, but most (if not all) of these bird species probably became extinct after the Polynesians arrived in the Hawaiian Islands, but before Cook arrived in 1778 establishing Western contact.
Since 1778, at least 24 more native Hawaiian land bird species or subspecies have become extinct.[iv] For example, the native kioea (Chaetoptila angustipluma) was last seen in 1859. In addition to the 59+ extinct endemic Hawaiian bird species and subspecies, more than 30 other endemic Hawaiian bird species are currently considered rare or endangered, and six of these species may already be extinct. (See Forest and Mountain Birds of the Hawaiian Islands, Chapter 7.)
Native Hawaiian Birds
Some bird species may have inhabited the Hawaiian Islands only after the Polynesian settlers arrived. This occurred because the original Hawaiian settlers, in the course of their food-producing activities, altered the native landscape by creating particular types of habitat (e.g., lo‘i kalo, or taro patches) that may have allowed certain wetland bird species to take hold.
The transformation of the environment by the Polynesian settlers of the Hawaiian Islands created substantial new areas of aquatic habitat. These new wetland habitats likely allowed some native species to expand their range, and may have allowed other species to become established. For example, the koloa (Anas wyvilliana, duck) and the ‘alae ula (Gallinula chloropus sandvicensis, gallinule) may have become established only after new wetland habitat was created by the Polynesian settlers. This is conjectured by researchers due to the absence of these birds from Pleistocene bird fossil and subfossil deposits excavated from various sites around the Hawaiian Islands.
The Polynesians’ introduction of the Polynesian black rat (Rattus exulans) to the Hawaiian Islands also provided a new food source for raptors, and may have allowed certain bird species (e.g., owls) to become established in the Hawaiian Islands. In a published study entitled Fossil evidence for a diverse biota from Kaua‘i and its transformation since human arrival (2001),[v]the authors detail the excavation of a Kaua‘i sinkhole and cave system on the south coast of Hawai‘i Island. Remains of pueo (Asio flammeus, Short-eared owls) did not show up in prehuman sediments but were found in sediments corresponding to the time after the Polynesians arrived.
The excavation results led the authors to note that the findings were “...consistent with Olson and James’ suggestion that this rodent-catching owl was able to colonize the archipelago only after the Pacific rat had been introduced by the Polynesians.”[vi] The authors also stated that, “...surprisingly...the only anseriform that has escaped extinction on the island and that one might expect to find in a wetland deposit, the Hawaiian Duck, or Koloa (Anas wyvilliana), is rare or absent...only a few bones from the site have been tentatively referred to as A. wyvilliana.”[vii] Thus the evidence remains inconclusive in regards to the whether koloa were found in the Hawaiian Islands before humans arrived or if they instead became established in the Hawaiian Islands only after the initial Polynesian settlement of the archipelago (and subsequent increase in wetland habitat).
Information regarding the exact time sequence of particular bird species inhabiting the Hawaiian Islands is still being gathered, and a more accurate picture is gradually emerging of the role that Polynesians’ played in creating new habitats that led to new bird species becoming established in the Hawaiian Islands.
Today, at least 22 species of marine birds are native to the Hawaiian Islands, with a total population of more than 12 million birds. More than 60% of these marine birds are noddies and terns. At least 33 species of geese and ducks (many migratory) are native to the Hawaiian Islands, along with five species of waterbirds, all endangered. (See Chapter 67)
As part of Bernice Pauahi Bishop Museum’s ongoing Hawaii Biological Survey, updated in the Records of the Hawaii Biological Survey for 2000,[viii] editors Lucius G. Eldredge and Scott E. Miller documented 228 bird species seen in the Hawaiian Islands (including 168 breeding and non-breeding migratory birds, some seen only rarely) as well as 60 endemic resident species. If the 46 introduced bird species are included, the total rises to 274 birds species in the Hawaiian Islands.
The study by Eldredge and Miller also noted that a total of 475 bird species are “known to occur or be sighted in Hawai‘i.”[ix] This number included extinct species (35+ becoming extinct before Cook arrived in 1778; 32 of these described from subfossils), and another 16 species extinct since 1778 (though a greater number are now presumed extinct, see above), and also included 150+ non-native birds that have been seen in the Hawaiian Islands but are not considered introduced species because they have not yet established breeding populations in the Hawaiian Islands.
The current total count of bird species in the Hawaiian Islands is apparently now at least 495, as the Records of the Hawaii Biological Survey for 2000[x]documented 10 new indigenous species (3 endemic and 7 introduced species), increasing the total number of bird species now established in the Hawaiian Islands to 294 species, including 241 native species (178 indigenous, 63 endemic) and 53 introduced species. (See Native Birds of the Hawaiian Islands, Chapter 7.)
Bishop Museum’s Hawaii Biological Survey
The Hawaii Biological Survey (with no ‘okina in “Hawaii”) is a Bishop Museum program that was established in 1992 by Hawai‘i’s State Legislature to maintain an ongoing inventory of the Hawaiian Islands’ natural history, including non-native as well as native species.
In coordination with a variety of state, federal and private groups (e.g., U.S. Fish & Wildlife Service, Nature Conservancy, University of Hawai‘i, and others), the Hawaii Biological Survey’s mission is to “gather, analyze, and disseminate the biological information necessary for the wise stewardship of Hawai‘i’s biological resources,” and also to “conduct a coordinated inventory and monitoring program to assess the overall status and trends in the abundance, health, and distribution of plants and animals, as well as the ecosystems upon which they depend.”[xi]
Bishop Museum scientists have done extensive studies to determine the total number of species found in the Hawaiian Islands, and have assembled a comprehensive tabulation of scientifically documented species, published by Bishop Museum and updated annually, including the Records of the Hawaii Biological Survey for 2000,[xii] published on March 25, 2002. This count of the Hawaiian Islands’ endemic, indigenous, and introduced species, edited by Neal L. Evenhuis and Lucius G. Eldredge, includes plants, trees, mollusks (e.g., snails), invertebrates (e.g., insects), fish, birds, reptiles (e.g., sea turtles), mammals (e.g., bats, monk seals, whales and dolphins), fungi, lichens, protists (including algae and human parasitic protists), and helminths. A total of 23,680 native Hawaiian species were documented, including 9,151 indigenous species, 9,456 endemic species, and 5,073 human-introduced species. Viruses and bacteria were not included. Of the 18,607 native Hawaiian species documented, about half (9,456 species) are endemic to the Hawaiian Islands.
The documented native Hawaiian species include: 1,995 protist species (82 endemic); 2,088 fungi and lichen species (240 endemic); 1,163 flowering plants (918 endemic); 719 other native plants (241 endemic); 1,243 mollusks (962 endemic); 1,143 fish species (149 endemic); 4 reptiles (sea turtles, none endemic); 241 bird species (63 endemic); and 25 mammals (2 endemic) and 9,270 native invertebrates (7,239 endemic). These native Hawaiian invertebrates include 6,284 arthropods [which included 5,818 insect species (5,462 endemic)] and 466 other arthropods (366 endemic)], along with 1,743 other invertebrates (449 endemic).
About 79% of native flowering plants in the Hawaiian Islands are endemic (918 of 1,163 species) along with approximately 26% of native birds, though nearly all native land birds of the Hawaiian Islands are endemic. About 94% of native insects of the Hawaiian Islands are endemic, as are 77% of native mollusks, 78% of native invertebrates, and 13% of native fish. Counts of indigenous and endemic Hawaiian species are constantly changing as researchers discover new species, analyze relationships between species (and subspecies) and further refine scientific classifications.
F.R. Warshauer, in The Atlas of Hawai‘i: Third Edition (1998), and relying in part on the Bishop Museum’s Hawaii Biological Survey, reported 142 native bird species (including extinct species); 7,800 described arthropods (out of an estimated 10,000 to 12,000 native Hawaiian arthropods); 1,300 vascular plants; 1,600 lower plants; 788 mollusks, 166 other invertebrates; and one mammal (not counting marine mammals), totaling about 15,000 native species in those categories.[xiii]
Arthropods
Arthropods are invertebrates (animals without backbones). All arthropods have a rigid external skeleton as well as segmented body parts and segmented legs (arthropod means “jointed feet”). At least 6,284 native Hawaiian arthropods have been documented.
About 95% of Hawaiian arthropods are insects, while the other 5% are closely related species that are not true insects, including centipedes, millipedes, spiders, and many others.
More than 94% of the Hawaiian Islands’ 5,818 known native insect species are endemic to the Hawaiian Islands. These 5,462 endemic insect species evolved over a period of at least 30 million years, inhabiting the ancient islands of the Hawaiian archipelago.
At least 23 species and subspecies of damselflies are endemic to the Hawaiian Islands along with five endemic dragonfly species. (See Damselflies and Dragonflies, Chapter 10.) Many more introduced damselfly and dragonfly species are now established in the Hawaiian Islands.
Two butterfly species are native to the Hawaiian Islands, along with about 13 more introduced butterfly species. (See Hawaiian Butterflies, Chapter 10.) More than 60 cave species (living in caves), are native to the Hawaiian Islands, and many of these are endemic to particular islands. (See Hawaiian Cave Species, Chapter 10.)
Pomace flies, (Drosophilidae and Scaptomyza), are a renowned example of adaptive radiation among the Hawaiian Islands’ insect fauna, with at least 860 documented species of Drosophilidae (in five genera), and perhaps many more yet to be described. All of these flies apparently evolved from a common pair of ancestors that lived in the Hawaiian Islands about 10 million years ago. For thousands of generations (over millions of years) the flies reproduced, and island-hopped from one volcano to the next as the islands eroded away at the northwest end of the chain and emerged at the southeast end of the chain.
Currently there are at least 2,161 known beetle species native to the Hawaiian Islands, and at least 1,416 of these beetle species are endemic to the Hawaiian Islands. There are more species of beetles than any other organism in the Hawaiian Islands, and the Hawaiian Islands also have the world’s highest proportion of beetles to other living things.
Thousands of species of native Hawaiian insects have yet to be scientifically described. It is estimated that the total number of native Hawaiian insects could exceed 10,000 species, which evolved from less than 500original colonizing species.[xiv] Also now established in the Hawaiian Islands are more than 550 introduced terrestrial arthropods and more than 2,700 non-native (introduced) insect species.
Snails and other Invertebrates
The Bishop Museum’s Hawai‘i Biological Survey has documented at least 781 terrestrial gastropods (759 endemic), and 7 Hawaiian freshwater gastropods (all endemic).
Endemic Hawaiian tree snails (terrestrial gastropods) provide another astounding example of adaptive radiation in the Hawaiian Islands, with more than 750 endemic species having evolved from just a few pioneers that reached the Islands several million years ago, probably in the feathers of migratory birds, or in mud on the birds’ feet. The snails are known by the Hawaiian name pūpū kani oe (“shell that sounds long”[xv]), referring to the belief that the snails sing.
Endemic Hawaiian land and freshwater snails evolved from about 25 colonizing species. These extremely adaptable snails had no natural predators, and were once found in a wide variety of habitats where they fed on the fungi that grow on leaves and forest debris (making the snails an important component of native ecosystems). The snails were known to vary considerably from valley to valley, and exhibited many shapes and colors, from gold tints to cream-colored with brown stripes.
Unfortunately, these native terrestrial snails have a very slow reproductive cycle as well as a low birthrate, and populations were significantly diminished by collectors and other causes. The snails are extremely vulnerable to non-native predators, particularly rats (Rattus exulans; R.. rattus; R. norvegicus); introduced snails (e.g., Euglandina rosea, the “cannibal snail,”); and the New Guinea flatworm (Platydemis manokwari), which was introduced to the Hawaiian Islands in the 1990s and now threatens the last, high-elevation habitats of the native snails.[xvi]
There were once more than 300 native Hawaiian snail species in the family Amastridae, but only about 12 or fewer of these species survive today. Recent research studying genetic mutation rates has shown that dozens of tree snail species in the subfamily achatinellinae originated from a single ancestor that likely lived in O‘ahu’s Ko‘olau mountain range about 3 million years ago. The last remaining habitat of native Hawaiian terrestrial snails is at high elevations on Moloka‘i, Lāna‘i, and O‘ahu, as well as some middle elevation areas on Hawai‘i Island and Maui.
Three brackish/freshwater snail species are endemic to the Hawaiian Islands. Two of these species, hapawai (Neritina vespertina) and pīpīwai (Theodoxus cariosus), prefer brackish waters, while a third species, hīhīwai (Neritina granosa) lives in freshwater stream areas. There are four native species of lymnaeid snails (pond snails) that live in streams and ponds as well as lo‘i kalo (taro patches).[xvii]
Overall, at least 1,243 native Hawaiian mollusks have been documented, and about 80% of these mollusk species are snails, sea slugs, and related species. Native Hawaiian mollusks are invertebrates that evolved from about 25 original immigrant species. There are also thousands of other native Hawaiian species of invertebrates (animals without backbones), including crabs, shrimp, and sea urchins.
Two native Hawaiian shrimp species, ‘ōpaekala‘ole (Atyoida bisulcata) and ‘ōpae ‘oeha‘a (Macrobrachium grandimanus), live in streams, while a third, ‘ōpae ‘ula (Halocaridina rubra) prefers anchialine ponds that are connected to the sea only by the percolations of saltwater and freshwater through lava rock. Heteromyenia baileyi is a native (indigenous) freshwater sponge that sometimes attains a bright green color.
Reptiles and Amphibians
No terrestrial reptiles or amphibians are considered native to the Hawaiian Islands, though there remains a possibility that some lizard species arrived in the Hawaiian Islands on their own (see below). Marine reptiles native to the Hawaiian Islands include five sea turtle species, some seen only rarely in Hawaiian waters. The two sea turtle species that come onto Hawaiian shores to lay their eggs are honu (Chelonia mydas, green sea turtle) and honu‘ea (Eretmochelys imbricata, hawksbill sea turtle).
On Monday, October 7, 2002, a small, female Olive ridley sea turtle (Lepidochelys olivacea) came ashore at Bayfront Beach on the island of Hawai‘i’s Hilo Bay. If confirmed, this is only the second reported sighting of a nesting Olive ridley sea turtle in the Hawaiian Islands. (See Hawaiian Sea Turtles, Chapter 6.)
Another marine reptile occasionally seen in Hawaiian waters (and thus considered native) is the Yellow-bellied sea snake (Pelamis platurus), a venomous yellow and black reptile that is about 29 inches (74 cm) long. A distant relative of the cobra, the Yellow-bellied sea snake is the only truly pelagic (open-ocean) sea snake of 55 known sea snake species.
[Illustration: Pelamis platurus (see above).]
Three skink species and four gecko species were established in the Hawaiian Islands before Western contact. These species likely came as stowaways aboard the voyaging canoes of the Polynesian settlers of the Hawaiian Islands, though it is possible some of these lizards may have reached the Hawaiian Islands on their own (e.g., by rafting on a log or on other ocean debris).
As of 2004 at least 21 lizard species have been found in the wild in the Hawaiian Islands (and thus are assumed to have established breeding populations). These lizards include geckos, skinks, anolis lizards, chameleons, horned lizards, and iguanas. Some of these lizard species are found throughout the Hawaiian Islands, while others are only found in very small areas on particular islands. Three of the lizard species are (presumably) no longer found in the wild in the Hawaiian Islands. (See Geckos and Other Reptiles, Chapter 10.)
No terrestrial snakes are native to the Hawaiian Islands, but there is one introduced snake species, called the Island blind snake (Ramphotyphlops braminus), which arrived in the Hawaiian Islands sometime around the 1930s in the soil in potted palm trees imported from the Philippines for landscaping. Now well established in the Hawaiian Islands, the Island blind snake is a burrowing snake that is about 6½ inches (16.5 cm) long, feeds on insects and small invertebrates, and spends most of its life underground.
Fish
Of the 24,000 known species of fish in the world, about 1,143 are native to the Hawaiian Islands, including 149 endemic fish species. About 536 of these fish species are inshore fishes found near reefs and other nearshore areas to a depth of about 200 feet (61 m). About 25% of these 536 inshore species are endemic to the Hawaiian Islands. The overall rate of endemism for native Hawaiian fish is about 13% (149 endemic species and 994 indigenous species). (See Hawaiian Reef Fish, Chapter 6.)
Fish native to the Hawaiian Islands also include five freshwater species, known as ‘o‘opu, which spend part of their lives in the ocean. These five ‘o‘opu fish species are endemic to the Hawaiian Islands, and include four species in the family Gobiidae (commonly called gobies) and one species in the family Eleotridae (‘o‘opu ‘akupa, Eleotris sandwicensis).
All five Hawaiian ‘o‘opu species are born as larvae in freshwater streams. The tiny larvae are then washed down into the ocean where they develop into fish before returning to the stream where they were born and swimming back up through the current.
‘O‘opu have specially adapted pelvic fins, which are fused to form a sucking disc, allowing the fish to climb up rocks and waterfalls and reach the upper levels of the streams to lay eggs and complete their life cycle. An ancient Hawaiian proverb refers to the ‘o‘opu as “Ka i‘a a ka wai nui i lawe mai ai,” (“The fish borne along by the flood.”[xviii]
Native Reef Fish
Native reef fish species of the Hawaiian Islands include:
- Ø 5 native species of angelfishes (Pomacanthidae).
- Ø 24 native species of butterflyfishes (Chaetodontidae).
- Ø 7 native species of parrotfishes (Scaridae).
- Ø At least 24 native species of surgeonfishes (Acanthuridae).
- Ø 43 native species of wrasses (Labridae).
- Ø 8 native species of filefishes (Monacanthidae).
- Ø 25 native species of scorpionfishes (Scorpaenidae).
- Ø 11 native species of triggerfishes (Balistidae).
- Ø 1 native species of trumpetfish (Aulostonus chinensis).
- Ø 2 native species of cornetfishes (Fistulariidae).
- Ø 17 species of damselfishes (Pomacentridae)
(See Hawaiian Reef Fish, Chapter 6, for more information about the fish listed above.)The endemic āholehole fish (Kuhlia sandvicensis, young stage of āhole, Hawaiian flagtail) sometimes enters lower stream areas, as do pua ‘ama‘ama (Mugil cephalus, young stage of ‘ama‘ama, mullet). ‘Ama‘ama were the main fish raised in ancient Hawaiian loko i‘a (fishponds). Also raised in fishponds were awa (Chanos chanos, milkfish), which reach a length of 6 feet (1.8 m).
Sharks, Rays and Eels
At least 41[xix] known species of sharks may be seen in Hawaiian waters (and are thus considered native to the Hawaiian Islands), and 20 of these shark species inhabit deep waters. Six shark species are relatively common around the Hawaiian Islands. (See Sharks, Chapter 6.) There are five native ray species, including a manta ray, an eagle ray, and three stingray species. (See Rays, Chapter 6.)
There are at least 38 native Hawaiian species of moray eel (Muraenidae), along with 16 native species of snake eels (Ophichthidae) and 3 native species of conger and garden eels (Congridae). (See Eels, Chapter 6.)
Other Marine Species
More than 860 marine and freshwater algae species are native to the Hawaiian Islands, including at least 80 endemic algae species. Of the more than 340 known native Hawaiian species of red limu, at least 67 are endemic and most are edible. (See Limu in Native Plants and Ferns of the Hawaiian Islands, Chapter 8.)
There are at least 150 different native Hawaiian coral species (approximately 20 to 30 percent are endemic), including 47 hard stony corals and more than 100 species of sea fan and other soft corals. (See Coral Reefs, Chapter 6.)
Land and Sea Mammals
There are 78 species of cetaceans worldwide, including all whales, dolphins, and porpoises. At least 22 cetacean species are considered native to the waters surrounding the Hawaiian Islands, though some are only occasionally seen near the Hawaiian Islands. The 22 native Hawaiian cetacean species include 17 toothed cetaceans, known as Odontocetes (e.g., dolphins), and five baleen whales, known as Mysticetes (e.g., humpback whales). (See Humpback Whales, and Spinner and Bottlenose Dolphins, Chapter 6.)
Aside from cetaceans, the only sea mammal native to the Hawaiian Islands is the Hawaiian monk seal, which often feeds near shore and then rests during the day on Island beaches. (See Hawaiian Monk Seals, Chapter 6.)
There are about 4,600 mammal species in the world, but just one native Hawaiian land mammal: ‘ōpe‘ape‘a (Lasiurus cinereus semotus, Hawaiian hoary bat). This furry insect-eating bat weighs in at about 6 ounces (170 g) with a wingspan up to 14 inches (36 cm). (See Hawaiian Hoary Bat, Chapter 10.)
Native and Polynesian-Introduced Plants
Before Western contact began to drastically change the landscape of the Hawaiian Islands, there were more than 1,700 known native Hawaiian plant species, including more than 1,000 native flowering plant species that had evolved from less than 300 original immigrants. At least 178 fern species are native to the Hawaiian Islands (124 endemic species). These native ferns evolved from about 135 colonizing species. At least 22 species of fern allies are native to the Hawaiian Islands, and seven of these fern ally species are endemic.
The 1999 Manual of the Flowering Plants of Hawai‘i: Revised Edition[xx]formally recognized 956 species of native Hawaiian flowering plants, including 850 species that are endemic to the Hawaiian Islands. Many more native Hawaiian plant species have since been discovered (see Introduction to Native Plants and Ferns of the Hawaiian Islands, Chapter 8), and are documented in the 2002 Electronic Supplement to the Manual of the Flowering Plants of Hawai‘i[xxi]as well as the Records of the Hawaii Biological Survey (see above), with a current total of at least 1,882 documented native Hawaiian plant species (1,159 endemic), including 1,163 native flowering plant species (918 endemic).
The extremely high rate of flowering plant endemism (79% endemic) in the Hawaiian Islands is primarily attributed to the geographical isolation of the Islands in the middle of the Pacific Ocean, thousands of miles from any continent. For more information about endemism and the evolution of native plant species, see the introduction to Chapter 8, Native Plants and Ferns of the Hawaiian Islands.
The Polynesian settlers of the Hawaiian Islands also brought at least 24 species of plants to the Islands, and may have brought as many as 30 plant species or more. (See Introduction to Chapter 9, Polynesian-Introduced Plants.)
Other Polynesian-Introduced Species
The early Polynesian settlers also brought many other species to Hawai’i besides plants. Some of these other species were brought to the Hawaiian Islands intentionally for use as food sources, but other species were brought unintentionally, coming as stowaways on the Polynesians’ voyaging canoes, and then establishing breeding populations in the Hawaiian Islands.
The geckos and the skinks listed below are presumed to have arrived in the Hawaiian Islands in this manner (as stowaways on the voyaging canoes), but there remains a possibility that some of these lizard species may instead have arrived in the Hawaiian Islands on their own (e.g., floating to the Hawaiian Islands on rafts of debris), and thus are native to the Hawaiian Islands.
Following is a summary of all species known to be introduced to the Hawaiian Islands by the early Polynesian settlers, as well as other species that may have been brought to the Hawaiian Islands, intentionally or unintentionally, by the Polynesians previous to Western contact in 1778.
Intentional Polynesian Introductions
- Pua‘a (Sus scrofa, pigs)
- Moa(Gallus gallus gallus, chickens)
- ‘Īlio(Canis familiaris, dogs)
- At least 24 (and probably more than 26) plant species
(See Polynesian-Introduced Plants, Chapter 9.)Unintentional Polynesian Introductions
- Geckos (Gekkonidae)
- § Indo-Pacific Gecko(Hemidactylus garnotii)
- § Mourning Gecko (Lepidodactylus lugubris)
- § Stump-Toed Gecko (Gehyra mutilata)
- § Tree Gecko (Hemiphyllodactylus typus)
(See Geckos and other Reptiles, Chapter 10.)- Skinks (Scincidae)
- § Azure-Tailed Skink (Emoia impar)
- § Moth Skink (Lipinia noctua noctua)
- § Snake-Eyed Skink (Cryptoblepharus poecilopleurus)
(See Geckos and other Reptiles, Chapter 10.)- Plants (possibly Polynesian-introduced)*
- § Kāmole(Ludwigia octivalvis, primrose willow)
- § ‘Ihi ‘Ai(Oxalis corniculata, yellow wood sorrel)
- § Kūkaepua‘a(Digitaria setigera)
- § Koali Kua Hulu(Merremia aegyptia, Hairy merremia)
- § Neke(Cyclosorus interruptus, formerly Thelypteris interrupta, maiden fern)
- § Paspalum scrobiculatum (ricegrass, no known Hawaiian name)
*Some of these plant species may be indigenous Hawaiian species, but have been noted as potential Polynesian introductions based on research. See Introduction to Polynesian-Introduced Plants, Chapter 9 for more information about the status of each of these species.- Other (Unintentional) Polynesian-Introduced Species
- § Polynesian black rat (Rattus exulans)
- § Ectoparasites
- o Laelaps hawaiiensis, and others (carried on rats)[xxiii]
- § Freshwater clams*
- o Pisidium casertanum
- o Musculium partumeium
*Found in ancient taro pond, thought to have been transported on taro stock brought to Hawai‘i by early Polynesian settlers.[xxiv]Endangered Species and Extinctions
Currently at least 30 species and subspecies of Hawaiian birds are federally listed as endangered, along with one subspecies listed as threatened.[xxv] About half of the Hawaiian Islands’ endemic birds are already extinct. More than 40 Hawaiian insect species have gone extinct in the last century, along with about 10% of native Hawaiian flowering plants (more than 100 extinct species). Just as the Hawaiian Islands are known as the world’s capital of endemic species, it may be considered the capital of extinct species.
At least 24 native Hawaiian bird taxa (species and subspecies) have gone extinct since 1778, from the Laysan rail (Porzana palmeri), which was last seen in 1944, to the Greater koa finch (Rhodacanthis palmeri), last seen in 1896. The Hawai‘i ‘ō‘ō (Moho nobilis), a bird long sought after for its beautiful feathers (see Native Birds of the Hawaiian Islands, Chapter 7) for use in ‘ahu ‘ula (royal cloaks), hasn’t been seen since 1934.[xxvi]
[Illustration: Hawai‘i ‘ō‘ō (Moho nobilis)]
Also extinct or likely extinct are 97 taxa (species and subspecies) of plants, 72 taxa of snails, and 74 taxa of insects.[xxvii] The presumably extinct insects include: 15 species of Coleoptera (beetles); five species of Diptera (flies, gnats, and mosquitoes); 15 species of Homoptera (leafhoppers, planthoppers, mealybugs, and scales); 38 species of Lepidoptera (moths and butterflies); one species of Odonata (damselflies and dragonflies); two species of Orthoptera (crickets and grasshoppers); and one species of Hemiptera (true bugs).[xxviii]
More than one third of the plant and animal species that are federally listed as endangered are Hawaiian. In all, more than 360 species are listed as either threatened or endangered under the federal and state Endangered Species Acts. These species include more than 275 endangered taxa (species and subspecies) of plants and more than 35 endangered vertebrate species, with at least another 40 invertebrate species listed as threatened.
The Manual of the Flowering Plants of Hawai‘i(1999) notes that, of the 1,094 taxa (including species and subspecies) of native Hawaiian plants included in the text, “...423 (38%) are considered to be extinct or threatened to some degree; of these taxa, 107 are presumed extinct (10% of the native taxa), 139 endangered (12%), 39 vulnerable (4%), and 138 rare (12%).”[xxix] The U.S. Fish & Wildlife Service endangered species program (in regards to Hawaiian species) does not count subspecies separately, resulting in different totals and percents in the categories listed above.
Though the Hawaiian Islands are just two tenths of one percent of the size of the United States, the Hawaiian Islands account for nearly 70% of the United States’ historically documented plant and animal extinctions. More than 30% of federally listed endangered species are Hawaiian, along with more than 40% of all birds listed as endangered. More than half of the Hawaiian Islands’ endemic plants are now threatened, endangered, or extinct, along with about half of the Hawaiian Islands’ endemic birds.
Threats to Native Species—Before 1778
The first threats to native species of the Hawaiian Islands came from the ancient Polynesian settlers of the Hawaiian Islands, who brought pua‘a (Sus scrofa, pigs), moa (Gallus gallus gallus, chickens), and ‘īlio (Canis familiaris, dogs), as well unintentional species introductions that came as stowaways on the voyaging canoes, including geckos (Gekkonidae), skinks (Scincidae), rats (Rattus exulans), and several other species (see above).
Early Hawaiian settlers also hunted birds for food (e.g., flightless geese, shearwaters and petrels) as well as for feathers (e.g., ‘ō‘ō (Moho species); mamo (Drepanis pacifica); and numerous other species). The bird feathers were prized for use in ‘ahu ‘ula (royal capes and cloaks), mahiole (feather-crested helmets), kāhili (royal feather standards) and other items of Hawaiian featherwork. (See Native Birds of the Hawaiian Islands, Chapter 7.)
After 1778, hunting by foreigners brought new pressures on many native birds, such as various species of albatross whose eggs and feathers were extensively collected, and the ‘alalā (Corvus hawaiiensis, Hawaiian crows), which were systematically shot by farmers protecting their poultry yards and feed pens.
In the decades after Captain Cook’s first visit to the Hawaiian Islands, the landscape of the Islands began to be drastically altered by increasing numbers of cattle, sheep, goats, and pigs. The pigs brought by Westerners were significantly larger than those brought to the Hawaiian Islands by the first Polynesian settlers.
The Chinese market for sandalwood (Santalum species) also led to deforestation, particularly from 1810 to 1840, the peak years of the sandalwood trade. Increasing demands for wood and pasture eventually cleared huge areas of the Hawaiian landscape.
The loss of native forest habitat decimated many of the endemic Hawaiian bird species, particularly populations at lower elevations (where most of the alteration of the landscape took place). Due to this habitat loss as well as mosquito-borne diseases (e.g., avian malaria), the last surviving species of these small and endangered, endemic Hawaiian birds are now restricted to the higher elevations.
Threats to Native Species—Post-Contact
Since Captain Cook established Western contact with the Hawaiian Islands in 1778, thousands of species have been introduced to the Hawaiian Islands accidentally or intentionally, and for various purposes, including pest control and ornamental landscaping. Many escaped pets, such as cats (Felis domesticus), have also established significant wild populations in some Hawaiian forests. Various lizards have become established in some areas, including skinks, chameleons, geckos, anoles, and iguanas. (See Geckos and other Reptiles, Chapter 10.)
After Western contact with the Hawaiian Islands was established in 1778, the influx of new species was rapid, and had devastating effects on native species. These introduced species (also known as alien, exotic, adventive, or introduced species) included birds, lizards, plants, and animals that detrimentally affected native species of the Hawaiian Islands. Some introduced species destroyed native habitats while other preyed on native species or competed with native species for food.
Sheep, goats, pigs, deer (axis and black-tailed), donkeys, and wallabies run wild in certain areas of the Hawaiian Islands. The wild donkeys are found on the island of Hawai‘i in the region north of the Kona airport near the Kona Village Resort and Four Seasons Hotel. The wild donkeys, known as the Kona Nightingales, are the remnants of a herd that was used to carry coffee from the uplands to the ocean until the end of World War II, when military jeeps began to be used to carry the coffee, and many of the donkeys were released on Hualālai Volcano.
Axis deer are found on Moloka‘i, Maui, and Lāna‘i. Black-tailed deer are found in northwestern Kaua‘i, and feral donkeys are found on the island of Hawai‘i. Goats and pigs are now found on all of the Hawaiian Islands with the exception of Lāna‘i.
Ka i‘a holehole iwi o ka ‘āina.
The fish of the land that strips the flesh from the bones.
Goats. When one pursues them for meat, many a limb suffers skinning and bruises.
(Pukui: 1337-145)
Two species of sheep may be found on the upper elevations of Mauna Kea Volcano on the island of Hawai‘i. One of these species, the Mouflon sheep, is also found on Lāna‘i. Recently, both sheep species have been nearly eliminated in an effort to protect native species.
Thousands of axis deer roam wild on the island of Maui, mostly from Hāna to Kapalua. These deer are the descendants of five deer that were introduced to Maui by State of Hawai‘i officials in 1959, and four more deer brought to Maui in 1960. The deer were initially brought to create new hunting opportunities. Maui’s axis deer population has increased rapidly over the last several decades (approximately doubling about every four years), with significant negative effects on native species and ecosystems of the Hawaiian Islands.
In 2002, the State of Hawai‘i signed a ten-year agreement to continue the availability of Mouflon sheep and axis deer hunting programs on Lāna‘i, where hunters take about 600 sheep and 700 deer each year. All of the above mentioned non-native feral ungulate species have had significant negative effects on native Hawaiian ecosystems as well as the native Hawaiian species they support.
An estimated 40 to 250 rock wallabies live in O‘ahu’s Kalihi Valley. Three of the miniature, kangaroo-like marsupials were brought to the Hawaiian Islands from Australia in 1916 to add to the collection of a private zoo. Two of the wallabies escaped, and today’s wild population in Kalihi Valley are their descendants.
A Summary of the Primary Threats to Native Species of the Hawaiian Islands
Historically, the primary threats to native species have included:Preventing Extinctions
The biological diversity of the Hawaiian Islands is astonishing: lush rainforests, underwater coral reefs, coastal dunes, subterranean lava tubes, and many other ecosystems support a variety of life forms. Unfortunately, many Hawaiian species cling tenuously to survival in remote and diminishing native habitat.
The combined efforts of scientists and concerned local citizens are helping to preserve native ecosystems and prevent extinctions. For example, there has been and continues to be a concerted community effort to rescue stranded wedge-tailed shearwater birds(Puffinus pacificus chlororhynchus). (See Newell’s and Wedge-Tailed Shearwaters section in Native Birds of the Hawaiian Islands, Chapter 7.) Other community volunteer efforts attempt to eradicate introduced plant species that threaten native ecosystems. One such event is the Banana Poka Festival in Kaua‘i’s Kōke‘e State Park.
Many native Hawaiian species are little known and even less seen. Scientists estimate that there are thousands of native Hawaiian species still unknown to science.
[Graphic: Endangered Species Distribution]
Chapter 5 Source Notes
All Hawaiian words in this text, the Hawaiian Encyclopedia, include proper diacritical marks, including the ‘okina (hamzah, or glottal stop) and kahakō (macron), except when the word is cited directly as part of a title or in a direct quote, in which case the words are spelled as originally published.
See Appendix 1: Note On Sources for more information about diacritical marks; objectivity; Hawaiian word spellings and meanings; scientific, cultural, and historic information; new research and discoveries; verifying dates and details of historic events; and Hawaiian sovereignty and other native issues.
[Note: This list is still in progress and being updated.]
Abbott, Isabella Aiona. Lā‘au Hawai‘i: Traditional Hawaiian Uses of Plants. Honolulu: Bishop Museum Press, 1992.
Abbott, Isabella Aiona. Limu: An Ethnobotanical Study of Some Hawaiian Seaweeds. Lawai: National Tropical Botanical Garden, 1984.
All About Hawaii (1948-1949): Thrum’s Hawaiian Annual and Standard Guide. Publications Division: Honolulu Star-Bulletin, Honolulu, Hawaii, 1949.
Ariyoshi, Rita. The National Geographic Traveler: Hawaii. Washington, D.C.: National Geographic Society, 2000.
Atlas of Hawai‘i: Second Edition. Honolulu: University of Hawai‘i Press, 1983.
Atlas of Hawai‘i: Third Edition. Honolulu: University of Hawai‘i Press, 1998.
Balcomb, Kenneth C. III. The Whales of Hawaii: Including All Species of Marine Mammals in Hawaiian and Adjacent Waters. San Francisco, California: Marine Mammal Fund, 1987.
Beletsky, Les. Hawaii: The Ecotravellers’ Wildlife Guide. London: Academic Press, 2000.
Berger, Andrew J. Bird Life in Hawaii. Honolulu: Island Heritage Limited, 1983.
Bishop Museum: Hawaii’s Extinct Species. Internet site: http://hbs.bishopmuseum.org/endangered/extinct.html, 7/28/2002.
Bishop Museum—Hawaii’s Extinct Species: Insects. Internet site: http://hbs.bishopmuseum.org/endangered/ext-insects.html, 7/28/2002.
Bishop Museum—Hawaii’s Extinct Species: Plants. Internet site: http://hbs.bishopmuseum.org/endangered/ext-plants.html, 7/28/2002.
Bishop Museum—Hawaii’s Extinct Species: Snails. Internet site: http://hbs.bishopmuseum.org/endangered/ext-snails.html, 7/28/2002.
Bishop Museum: Native plant species in Hawai‘i. Internet site: http://www.bishopmuseum.org/greenwell/gwplants.html, 8/07/2002.
Bishop Museum: The How Many Species Are there in Hawaii Web Page. Includes: How many species are there in Hawaii?, from the “Records of the Hawaii Biological Survey for 1994” as published in the Bishop Museum Occasional Paper volume 41: 3-18; and Number of Hawaiian Species: Supplement 1, originally published in Bishop Museum Occasional Papers 45: 8-17 (1996). Internet site: http://hbs.bishopmuseum.org/hispp.html, 10/17/2002.
Bishop Museum—The role of the Hawaii Biological Survey. Internet site: http://hbs.bishopmuseum.org/hbs3.html.
Boynton, David. Boar hunting: Hunting the feral pigs helps to protect the forest ecology. Kauai Magazine, Fall, 1992.
Cabrera, Theresa. The Hawaiian hoary bat: Hawaii’s only native land mammal. Honolulu: Nature Conservancy of Hawaii Newsletter, Spring, 1995.
Cave critters added to endangered species list. The Garden Island, 1/16/2000.
Chang, Lester. Visit to “sinkhole” offers a look at thousands of years of Kaua‘i’s natural history. The Garden Island, 3/25/2002.
Chang, Lester. $2.8 million helping bats, other species: Federal protection money pouring into state. The Garden Island, 9/29/2001.
Clark, Hugh. Turtle nest in Hilo great rarity in itself. The Honolulu Advertiser, 10/12/2002.
Culliney, John L. Islands in a Far Sea. San Francisco: Sierra Club Books, 1988.
Daws, Gavan. Shoal of Time: A History of the Hawaiian Islands. Honolulu: University of Hawaii Press, 1968.
Degener, Otto. Plants of Hawaii National Parks: Illustrative of Plants and Customs of the South Seas. Braun-Brumfield, Inc., 1973.
Denny, Jim. The Birds of Kaua‘i. Honolulu: University of Hawai‘i Press, 1999.
Endangered animals in the Hawaiian Islands: Hawaiian Bat. Internet site produced by the US Fish and Wildlife Service: http://www.r1.fws.gov/pacific/wesa/hrybatindex.html.
Eyre, David L. By Wind, By Wave. Honolulu: The Bess Press, 2000.
Federal Register, 2002, Vol. 67, U.S. Fish & Wildlife Service, Threatened and Endangered Species System (TESS), Listings by State and Territory as of 7/20/2002. Internet site: http://ecos.fws.gov/servlet/TESSWebpageUsaLists?state=HI, 7/28/2002.
Fish breakthrough good news for Hawai‘i. The Honolulu Advertiser, 1/24/2002.
Gulko, David. Hawaiian Coral Reef Ecology. Honolulu, Hawaii: Mutual Publishing, 1998.
Handy, Emory, Bryan, Buck, Wise and Others. Ancient Hawaiian Civilization: A Series of Lectures Delivered at the Kamehameha Schools. Mutual Publishing: Honolulu, Hawaii, 1999.
Handy, E.S. Craighill, and Handy, Elizabeth Green, with the collaboration of Mary Kawena Pukui. Native Planters in Old Hawaii: Their Life, Lore, and Environment. Honolulu: Bishop Museum Press, 1972.
Hargreaves, Dorothy and Bob. Tropical Blossoms of the Pacific. Hargreaves Company, Inc., 1970.
Hargreaves, Dorothy and Bob. Tropical Trees of the Pacific. Hargreaves Company, Inc., 1970.
Hawaiian and Pacific Islands National Wildlife Refuges. United States Department of the Interior Fish and Wildlife Service, September, 1985.
Hawaiian Cave Animals: Threatened and Endangered Species; Endangered Animals in the Hawaiian Islands. U.S. Fish and Wildlife Service. Internet site: http://www.r1.fws.gov/pacific/wesa/caveanimals.html, 6/17/2000.
Hawaiian Native Plant Genera. University of Hawai‘i Botany Department. Internet site: http://www.botany.hawaii.edu/faculty/carr/natives.htm, 5/03/2002.
Hawai‘i Data Book: A Statistical Reference to Hawai‘i’s Social, Economic and Political Trends. Edited by Robert C. Schmitt. Honolulu: Mutual Publishing, 2002.
Hawaii’s Birds. Honolulu: Hawaiian Audubon Society, 1993.
Hayworth, Philip. Hawai‘i’s Marine Zoos: The Waikīkī Aquarium, Sea Life Park and Maui Ocean Center showcase the incredible diversity of marine life flourishing in Hawaiian waters. Spirit of Aloha Magazine, May/June, 2002.
Hobson, Edmund, and Chave, E.H. Hawaiian Reef Animals. University of Hawaii Press, 1972.
Hoover, John P. A Pocket Guide to Hawai‘i’s Underwater Paradise. Honolulu: Mutual Publishing, 1997.
Hoover, John P. Hawaii’s Fishes: A Guide for Snorkelers, Divers and Aquarists. Honolulu: Mutual Publishing, 1993.
Hoover, John P. Hawai‘i’s Sea Creatures: A Guide to Hawai‘i’s Marine Invertebrates. Honolulu: Mutual Publishing, 1998.
Howarth, F.G. and Mull W.P. Hawaiian Insects and their Kin. Honolulu: University of Hawai‘i Press, 1992.
Hurley, Timothy. Another nastier lizard found. The Honolulu Advertiser, 3/19/2002.
Hurley, Timothy. Axis deer a growing nuisance on Maui: Report urges containment. The Honolulu Advertiser, 8/20/2002.
Hurley, Timothy. Death revives donkey debate. The Honolulu Advertiser, 10/06/2002.
Hurley, Timothy. Poison frog found on Maui. The Honolulu Advertiser, 3/29/2002.
Hurley, Timothy. State extends Lāna‘i hunting. The Honolulu Advertiser, 7/05/2002.
Ii, John Papa. Fragments of Hawaiian History. Translated by Mary Kawena Pukui; Edited by Dorothy B. Barrère. Honolulu: Bishop Museum Press, 1959.
Joesting, Edward. Kauai: The Separate Kingdom. Hawaii: University of Hawaii Press, 1984.
Kamakau, Samuel Manaiakalani. Ka Po‘e Kahiko: The People of Old. Translated from the Newspaper Ke Au ‘Oko‘a by Mary Kawena Pukui. Arranged and edited by Dorothy B. Barrère. Bishop Museum Special Publication 51. Honolulu: Bishop Museum Press, 1991.
Kāne, Herb Kawainui. Ancient Hawai‘i. Captain Cook, Hawai‘i: The Kawainui Press, 1997.
Kaufman, Gregory Dean, and Forestell, Paul Henry. Hawaii’s Humpback Whales. Honolulu: Island Heritage Publishing, 1986.
Kawaharada, Dennis. The Settlement of Polynesia, Part 1. Polynesian Voyaging Society. Internet Site: http://leahi.kcc.hawaii.edu/org/pvs/mirgrationspart1.html.
Kepler, Angela Kay. Hawai‘i’s Floral Splendor: A Friendly and Color Identification Guide to Native and Introduced Flowers of All the Hawaiian Islands. Honolulu: Mutual Publishing, 1997.
Kilauea Point Natural History Association. Hawaii’s Wetlands. Kauai: Hawai‘i Nature Focus: Nature Studies for Children, # 11.
Kirch, Patrick Vinton. Feathered Gods and Fishhooks. Honolulu: University of Hawai‘i Press, 1985.
Kirch, Patrick V., and Sahlins, Marshall. Anahulu: The Anthropology of History in the Kingdom of Hawaii. Volume One: Historical Ethnography (Marshall Sahlins, with the Assistance of Dorothy B. Barrère). Chicago and London: The University of Chicago Press, 1992.
Kirch, Patrick V., and Sahlins, Marshall. Anahulu: The Anthropology of History in the Kingdom of Hawaii. Volume Two: The Archaeology of History (Patrick V. Kirch). Chicago and London: The University of Chicago Press, 1992.
Krauss, Beatrice H. Plants in Hawaiian Culture. Honolulu: University of Hawaii Press, 1993.
Krauss, Beatrice H. Plants in Hawaiian Medicine. Honolulu: The Bess Press, 2001.
Life in Early Hawai‘i: The Ahupua‘a—Third Edition. Honolulu: Kamehameha Schools Press, 1994.
Lum, Curtis. Iguanas making themselves at home in O‘ahu communities. The Honolulu Advertiser, 4/09/2002.
Malo, David. Hawaiian Antiquities: Mo‘olelo Hawai‘i. Translated from the Hawaiian by Dr. Nathaniel B. Emerson, 1898. Bernice P. Bishop Museum Special Publication 2: Second Edition. Honolulu: Bishop Museum, 1951.
Manguchei, Kendyce. Poor man’s time machine uncovers Kaua‘i’s past at Maha‘ulepu. The Garden Island, 6/23/2002.
Matisoo-Smith, E., Roberts, R.M., Irwin, G.J., Allen, J.S., Penny, D. and Lambert, D.M. Patterns of prehistoric human mobility in Polynesia indicated by mtDNA from the Pacific rat. Anthropology, Vol. 95, pp. 15145-15150, December, 1998.
McKeown, Sean. A Field Guide to Reptiles and Amphibians in the Hawaiian Islands. California: Diamond Head Publishing, Inc., 1996.
Mee, Jonathan. Living lava lamps. Oregon Coast, July/August, 1995.
Meinkoth, Norman A. The Audubon Society Field Guide to North American Seashore Creatures. New York: Alfred A. Knopf, 1981.
Memories of Duke Kahanamoku. Internet site: http://planet-hawaii.com/duke/memories.htm
Merlin, Mark. Hawaiian Coastal Plants: An Illustrated Field Guide. Honolulu: Pacific Guide Books, 1999.
Merlin, Mark David. Hawaiian Forest Plants: A Hiker’s Guide. Honolulu: The Oriental Publishing Company, 1977.
Merriam Webster’s Collegiate Dictionary: Tenth Edition. Springfield, Massachusetts: Merriam-Webster, Incorporated, 1996.
Mitchell, Donald D. Kilolani. Resource Units in Hawaiian Culture: Revised Edition. Honolulu: Kamehameha Schools Press, 1992.
Miyano, Leland, and Peebles, Douglas. A Pocket Guide to Hawaii’s Flowers. Honolulu Hawaii: Mutual Publishing, 1997.
Miyano, Leland. Hawai‘i’s Beautiful Trees. Honolulu: Mutual Publishing, 1997.
Moriarty, Linda Paik. Pūpū ‘o Ni‘ihau: Shell Lei of Ni‘ihau. Honolulu: Honolulu Academy of Arts, 2001.
Mullins, Joseph G. Hawaiian Journey. Honolulu: Mutual Publishing, 1978.
New lizard species established on Oahu. The Garden Island, 2/12/2001.
Oliver, Anthony Michael. Hawaii Fact and Reference Book. Mutual Publishing, 1995.
Oliver, Douglas. Polynesia in Early Historic Times. Honolulu, Hawai‘i: The Bess Press, 2002.
Payne, Roger. Among Whales. New York: Charles Scribner’s Sons: Simon and Schuster Inc., 1995.
Polhemus, Dan, and Asquith, Adam. Hawaiian Damselflies: A Field Identification Guide. Honolulu: Bishop Museum Press, 1996.
Pratt, H. Douglas. A Pocket Guide to Hawai‘i’s Birds. Honolulu: Mutual Publishing, 1996.
Pratt, H. Douglas. Enjoying Birds in Hawaii: A Birdfinding Guide to the Fiftieth State. Honolulu: Mutual Publishing, 1993.
Pukui, Mary Kawena, and Elbert, Samuel H. Hawaiian Dictionary: Revised and Enlarged Edition. Honolulu: University of Hawaii Press, 1986.
Pukui, Mary Kawena, and Elbert, Samuel H, with Mookini, Esther T. and Nishizawa, Yu Mapuana. New Pocket Hawaiian Dictionary. Honolulu: University of Hawaii Press, 1992.
Pukui, Mary Kawena. ‘Ōlelo No‘eau: Hawaiian Proverbs & Poetical Sayings. Honolulu: Bishop Museum Press, 1983.
Pukui, Mary Kawena, and Elbert, Samuel H. & Mookini, Esther T. Place Names of Hawaii: Revised & Expanded Edition. Honolulu: University of Hawaii Press, 1974.
Randall, John E. Shore Fishes of Hawai‘i. Honolulu: University of Hawai‘i Press, 1996.
Randall, John E. Underwater Guide to Hawaiian Reef Fishes. Pennsylvania: Harrowood Books, 1981.
Rare turtle lays eggs on Hilo beach. The Garden Island, 10/12/2002.
Rock Wallaby released in Kalihi Valley. The Garden Island, 6/17/2002.
Rogers, Michael. Little Birds Lost. National Wildlife, October/November, 1993.
Ronck, Ronn. Hawaii Almanac. Honolulu: University of Hawaii Press, 1984.
Roth, Susan. Hawai‘i expert testifies for House bill on invasive species: Administration says effort not a priority. The Honolulu Advertiser, 3/15/2002.
Royte, Elizabeth. On the brink: Hawaii’s vanishing species. National Geographic Magazine, September, 1995.
Russo, Ron. Hawaiian Reefs: A Natural History Guide. Wavecrest Publications, 1994.
Sabelli, Bruno. Simon & Schuster’s Guide to Shells. Edited by Harold S. Feinberg. New York: Simon & Schuster Inc., 1979.
Sahlins, Marshall. Anahulu: The Anthropology of History in the Kingdom of Hawai‘i. Chicago: The University of Chicago Press, 1992.
Schmid, Randolph E. Pacific states vulnerable to giant sea waves. The Garden Island, 7/29/1998.
Scientists unstick the gecko mystery: Foot hairs pave way for ‘glue’. The Honolulu Advertiser (from Los Angeles Times), 6/19/2000.
Scott, Susan. Plants and Animals of Hawai‘i. Honolulu: Bess Press, 1991.
Scott, Susan and Thomas, Craig, M.D. Poisonous Plants of Paradise: First Aid and Medical Treatment of Injuries from Hawai‘i’s Plants. Honolulu: University of Hawai‘i Press, 2000.
Seven ‘alala chicks released into wild. ‘Elepaio: Journal of the Hawaiian Audubon Society, Volume 55, #1, February, 1995.
Silverstein, Duane. Our tribe must make all islands survivors. The Honolulu Advertiser, 3/24/2002.
Sprague, Brandon. In pursuit of Kauai’s dwindling native bird population. The Garden Island. (no date)
TenBruggencate, Jan. Aliens overrun Island environs. The Honolulu Advertiser, 7/09/2001.
TenBruggencate, Jan. Beetles live here in abundance. The Honolulu Advertiser, 6/02/2003.
TenBruggencate, Jan. Botanical secrets of sea await discovery. The Honolulu Advertiser, 7/3/2000.
TenBruggencate, Jan. Kaua‘i limestone 315,000 years old. The Honolulu Advertiser, 2/19/2002.
TenBruggencate, Jan. Last habitat for crow endangered. The Honolulu Advertiser, 2/15/1996.
TenBruggencate, Jan. Loss of variability left nēnē vulnerable. The Honolulu Advertiser, 2002.
TenBruggencate, Jan. Nene flourish on Kauai. The Honolulu Advertiser, 7/19/1995.
TenBruggencate, Jan. Science mission finds new species: Discoveries rife on outlying isles. The Honolulu Advertiser, 10/19/2000.
TenBruggencate, Jan. Sinkhole yields treasure trove of bird fossils. The Honolulu Advertiser, 2/25/2002.
TenBruggencate, Jan. Variety fills island ecosystems. The Honolulu Advertiser, 9/16/2002.
The Audubon Society Field Guide to North American Fishes, Whales, and Dolphins. Knopf, Alfred A., 1983.
The Plant Book: The World of Plants in a Single Volume. Australia: James Mills-Hicks, Random House Australia Pty Ltd., 2001.
Thompson, Rod. Endangered olive ridley turtle lays eggs at Hilo Bay: Officials are asking the public to not disturb the rare nest, which has 124 eggs. The Honolulu Star-Bulletin, 10/12/2002.
United States Fish & Wildlife Service: Pacific Islands—Endangered Species; Threatened and Endangered Animals in the Hawaiian Islands. Hawaiian Cave Animals: Kaua‘i Cave Wolf Spider and Kaua‘i Cave Amphipod. Internet site: http://pacificislands.fws.gov/wesa/caveanimals.html, 7/24/2002.
U.S. Fish & Wildlife Service, Threatened and Endangered Species System (TESS): Listings by State and Territory, as of 7/28/2002. Internet site: http://ecosfws.gov/webpage/webpage_usa_lists.html?state=all&module=undefined, 7/28/2002.
U.S. Fish & Wildlife Service, Threatened and Endangered Species System (TESS): U.S. Listed Vertebrate Animal Species Report by Taxonomic Group as of 7/28/2002. Internet site: http://ecos.fws.gov/webpage/webpage_vip_listed.html?module=undefined&code=V&listings=0, 7/28/2002.
Valier, Kathy. Ferns of Hawai‘i. Honolulu, University of Hawai‘i Press, 1995.
Viotti, Vicki. The lei of the land. The Honolulu Advertiser, 4/16/1995.
Wagner, Warren L., and Herbst, Derral R. Electronic Supplement to the Manual of the Flowering Plants of Hawai‘i. Internet site:http://rathbun.si.edu/botany/pacificislandbiodiversity/hawaiianflora/supplement.htm, 3/05/2002.
Wagner, Warren L., Herbst, Derral R., and Sohmer, S.H. Manual of the Flowering Plants of Hawai‘i: Revised Edition, Volume 1. Honolulu: University of Hawai‘i Press; Bishop Museum Press, 1999.
Wagner, Warren L., Herbst, Derral R., and Sohmer, S.H. Manual of the Flowering Plants of Hawai‘i: Revised Edition, Volume 2. Honolulu: University of Hawai‘i Press; Bishop Museum Press, 1999.
Witte, Astrid and Mahaney, Casey. Hawaiian Reef Fish. Waipahu, Hawai‘i: Island Heritage Publishing, 1998.
Yamamoto, Mike N., and Tagawa, Annette W. Hawai‘i’s Native & Exotic Freshwater Animals. Honolulu: Mutual Publishing, 2000.
[1]Juvik, James O. Biogeography in Atlas of Hawai‘i: Third Edition. Edited by Sonia P. Juvik and James O. Juvik. Honolulu: University of Hawai‘i Press, 1998.
[2] p. 276, Pukui, Mary Kawena. ‘Ōlelo No‘eau: Hawaiian Proverbs & Poetical Sayings. Honolulu: Bishop Museum Press, 1983. Proverb 2529.
[iii] Conant, Sheila. Extinct Birds, in Atlas of Hawai‘i: Third Edition. Edited by Sonia P. Juvik and James O. Juvik. Honolulu: University of Hawai‘i Press, 1998.
[iv] Bishop Museum: Hawai‘i’s Extinct Species—Birds. Internet site: http://hbs.bishopmuseum.org/endangered/ext-birds.html, 7/28/2002.
[v] Burney, David A., James, Helen F., Burney, Lida Pigott, Olson, Storrs L., Kikuchi, William, Wagner, Warren L., Burney, Mara, McCloskey, Deirdre, Kikuchi, Delores, Grady, Frederick V., Gage II, Reginald, and Nishek, Robert. Fossil evidence for a diverse biota from Kaua‘i and its transformation since human arrival. Ecological Monographs, 71 (4), 2001, pp. 615-641.
[vi] Burney, David A., James, Helen F., Burney, Lida Pigott, Olson, Storrs L., Kikuchi, William, Wagner, Warren L., Burney, Mara, McCloskey, Deirdre, Kikuchi, Delores, Grady, Frederick V., Gage II, Reginald, and Nishek, Robert. Fossil evidence for a diverse biota from Kaua‘i and its transformation since human arrival. Ecological Monographs, 71 (4), 2001, pp. 615-641.
[vii] Burney, David A., James, Helen F., Burney, Lida Pigott, Olson, Storrs L., Kikuchi, William, Wagner, Warren L., Burney, Mara, McCloskey, Deirdre, Kikuchi, Delores, Grady, Frederick V., Gage II, Reginald, and Nishek, Robert. Fossil evidence for a diverse biota from Kaua‘i and its transformation since human arrival. Ecological Monographs, 71 (4), 2001, pp. 615-641.
[viii] Evenhuis, Neal L., and Eldredge, Lucius G., Editors. Records of the Hawaii Biological Survey for 2000. Bishop Museum Occasional Papers, Number 68, 69. Honolulu: Bishop Museum, 3/25/2002.
[ix] The How Many Species Are there in Hawaii Web Page. Includes: How many species are there in Hawaii?, from the “Records of the Hawaii Biological Survey for 1994” as published in the Bishop Museum Occasional Paper volume 41: 3-18; and Number of Hawaiian Species: Supplement 1, originally published in Bishop Museum Occasional Papers 45: 8-17 (1996). Internet site: http://hbs.bishopmuseum.org/hispp.html, 10/17/2002.
[x] Evenhuis, Neal L., and Eldredge, Lucius G., Editors. Records of the Hawaii Biological Survey for 2000. Bishop Museum Occasional Papers, Number 68, 69. Honolulu: Bishop Museum, 3/25/2002.
[xi] The Role of the Hawaii Biological Survey. Bishop Museum Internet site: http://hbs.bishopmuseum.org/hbs3.html, 8/072002.
[xii] Evenhuis, Neal L., and Eldredge, Lucius G., Editors. Records of the Hawaii Biological Survey for 2000. Bishop Museum Occasional Papers, Number 68, 69. Honolulu: Bishop Museum, 3/25/2002.
[xiii] Warshauer, F.R. Alien Species and Threats to Native Ecology, in the Atlas of Hawai‘i: Third Edition. Edited by Sonia P. Juvik and James O. Juvik. Honolulu: University of Hawai‘i Press, 1998.
[xiv] Howarth, F.G., and Mull, W.P. Hawaiian Insects and their Kin. Honolulu: University of Hawai‘i Press, 1992.
[xv] Pukui, Mary Kawena, and Elbert, Samuel H. Hawaiian Dictionary: Revised and Enlarged Edition. Honolulu: University of Hawaii Press, 1986.
[xvi] Hadfield, Michael G. Hawaiian Tree Snails, in the Atlas of Hawai‘i: Third Edition. Edited by Sonia P. Juvik and James O. Juvik. Honolulu: University of Hawai‘i Press, 1998.
[xvii] Yamamoto, Mike N., and Tagawa, Annette W. Hawai‘i’s Native & Exotic Freshwater Animals. Honolulu: Mutual Publishing, 2000.
[xviii] p. 144, Pukui, Mary Kawena. ‘Ōlelo No‘eau: Hawaiian Proverbs & Poetical Sayings. Honolulu: Bishop Museum Press, 1983. Proverb 1323.
[xix] Includes viper shark (Trigonognathus kabeyai) reported in: Evenhuis, Neal L., and Eldredge, Lucius G., Editors. Records of the Hawaii Biological Survey for 2000. Bishop Museum Occasional Papers, Number 68, 69. Honolulu: Bishop Museum, 3/25/2002.
[xx] Wagner, Warren L., Herbst, Derral R., and Sohmer, S.H. Manual of the Flowering Plants of Hawai‘i: Revised Edition, Volumes 1 and 2. Honolulu: University of Hawai‘i Press; Bishop Museum Press, 1999.
[xxi] Wagner, Warren L., and Herbst, Derral R. Electronic Supplement to the Manual of the Flowering Plants of Hawai‘i. Internet site:http://rathbun.si.edu/botany/pacificislandbiodiversity/hawaiianflora/supplement.htm, 3/05/2002.
[xxii] Kirch, Patrick V. (Citing Cooke 1926; Cooke and Kondo 1960; Pilsbry 1916-1918; Solem 1959.) The Impact of the Prehistoric Polynesians on the Hawaiian Ecosystem. Pacific Science, Vol.36, No.1, January, 1982.
[xxiii] Kirch, Patrick V. The Impact of the Prehistoric Polynesians on the Hawaiian Ecosystem. Pacific Science, Vol.36, No.1, January, 1982.
[xxiv] Evenhuis, Neal L., and Eldredge, Lucius G., Editors. Records of the Hawaii Biological Survey for 2000. Bishop Museum Occasional Papers, Number 68, 69. Honolulu: Bishop Museum, 3/25/2002.
[xxv] Federal Register, 2002, Vol. 67, U.S. Fish & Wildlife Service, Threatened and Endangered Species System (TESS), Listings by State and Territory as of 7/20/2002. Internet site: http://ecos.fws.gov/servlet/TESSWebpageUsaLists?state=HI, 7/28/2002.
[xxvi] Bishop Museum: Hawai‘i’s Extinct Species—Birds. Internet site: http://hbs.bishopmuseum.org/endangered/ext-birds.html, 7/28/2002.
[xxvii] Bishop Museum - Hawaii’s Extinct Species. Internet site: http://hbs.bishopmuseum.org/endangered/extinct.html, 7/28/2002.
[xxviii] Bishop Museum - Hawaii’s Extinct Species: Insects. Internet site: http://hbs.bishopmuseum.org/endangered/ext-insects.html, 7/28/2002.
[xxix] Wagner, Warren L., Herbst, Derral R., and Sohmer, S.H. Manual of the Flowering Plants of Hawai‘i: Revised Edition, Volumes 1 and 2. Honolulu: University of Hawai‘i Press; Bishop Museum Press, 1999.