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Threats to Native Aquatic Insect Biodiversity in Hawai'i and the Pacific

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Threats to Native Aquatic Insect Biodiversity in Hawai'i and the Pacific THREATS TO NATIVE AQUATIC INSECT BIODIVERSITY IN HAWAI'I AND THE PACIFIC, AND CHALLENGES IN THEIR CONSERVATION A DISSERTATION SUBMITTED TO THE GRADUATE DIVISION OF THE UNIVERSITY OF HAWAI 'I IN PARTIAL FULFILLMENT OF THE REQUIREMENTS FOR THE DEGREE OF DOCTOR OF PHILOSOPHY IN ENTOMOLOGY AUGUST 2005 By Ronald A. Englund Dissertation Committee: Mark Wright, Chairperson Dan Rubinoff Neal Evenhuis Dan Polhemus Andrew Taylor TABLE OF CONTENTS ACKNOWLEDGEMENTS , ii ABSTRACT iii LIST OF TABLES vi LIST OF FIGURES viii CHAPTER 1. THE IMPACTS OF INTRODUCED POECILIID FISH AND ODONATA ON THE ENDEMIC MEGALAGRION (ODONATA) DAMSELFLIES OF 0'AHU ISLAND, HAWAI'I 1 CHAPTER 2: EVALUATING THE EFFECTS OF INTRODUCED RAINBOW TROUT (Oncorhynchus mykiss) ON NATIVE STREAM INSECTS ON KAUA'I ISLAND, HAWAI'I 40 CHAPTER 3. LONG-TERM MONITORING OF ONE OF THE MOST RESTRICTED INSECT POPULATIONS IN THE UNITED STATES, Megalagrion xanthomelas Selys-Longchamps, 1876, AT TRIPLERARMY MEDICAL CENTER, O'AHU, HAWAI'I... 76 CHAPTER 4. THE LOSS OF NATIVE BIODIVERSITY AND CONTINUING NONINDIGENOUS SPECIES INTRODUCTIONS IN FRESHWATER, ESTUARINE, AND WETLAND COMMUNITIES OF PEARL HARBOR, O'AHU, HAWAIIAN ISLANDS 91 CHAPTER 5. FLOW RESTORATION AND PERSISTENCE OF INTRODUCED SPECIES IN WAlKELE STREAM, 0'AHU 125 CHAPTER 6: INVASIVE SPECIES THREATS TO NATIVE AQUATIC INSECT AND ARTHROPOD BIODIVERSITY IN HAWAI'I, THE PACIFIC AND OTHER RELEVANT AREAS WITH DISCUSSION OF CONSERVATION MEASURES 143 ACKNOWLEDGEMENTS I would like to thank the many people that have made this dissertation possible. I especially would like to extend my thanks and warmest gratitude to my advisor Mark Wright, whose sense of humor and keen intellect made this process as enjoyable as it can be. My committee, consisting of Dan Rubinoff, Neal Evenhuis, Dan Polhemus and Andrew Taylor provided valuable insights and advice throughout. I sincerely acknowledge and appreciate the efforts of my entire committee throughout my time at the University of Hawai'i at Manoa. I am also deeply appreciative of Neal Evenhuis and Allen Allison for their encouragement, and allowing me the flexibility to pursue a Doctorate while being employed at the Bishop Museum. I have greatly enjoyed the scientific and cultural 'ohana at the Bishop Museum that always provided an ideal research and working atmosphere. Rob Cowie and Frank Howarth of the Bishop Museum also provided valuable reviews and advice for many of these chapters. Bishop Museum librarians Patti Belcher and B.J. Short were always helpful in tracking down the many obscure references. Several key organizations provided the support that allowed my research to take place, and I thank the following organizations that funded this research: Bishop Museum, Hawaii Division of Aquatic Resources, Nature Conservancy, Smithsonian Institution, and the Delegation ala Recherche Polynesie fran<;aise. People too numerous to mention assisted me in various aspects of the fieldwork required for this wide­ ranging dissertation, and I greatly appreciate help from David Preston, Betsy Gagne, Dan Polhemus, Jean­ Yves Meyer, BenoIT Fontaine, Olivier Gargominy, Tina Lau, Stephanie Loo, Brian Naeole, Alison Sherwood, and Steve Jordan. Special thanks goes to the Hawaii Division of Aquatic Resources crew including Bob Nishimoto, Glenn Higashi, Darrell Kuamo'o, John Kahiapo, Skippy Hau, Bill Puleloa, and Mike Yamamoto. Bill Devick and Bob Nishimoto were also instrumental in encouraging and funding much of this research as well. La Vonne Furtado provided consistent moral support during the crucial final stages of this journey. I am also privileged to have incredibly supportive parents and a wonderful family, without whom I would have never attempted this work. I would like to dedicate this dissertation to my wonderful parents, Stanley and Marjorie Englund. 11 ABSTRACT Although the decline in numbers and diversity and threat to native insects in the Hawaiian Islands is widely recognized by field scientists there has been little progress in either documenting the real decline of native species, or in demonstrating specific causes of the overall decline of these species. Additionally, few conservation actions to either restore populations or mitigate actual threats to native arthropods have been mentioned in the literature. The following chapters examine several assessments of relevant aquatic systems and the native aquatic insects dwelling within, where there has either been a perceived or real decline of these native Hawaiian aquatic arthropods because of threats from invasive or introduced species. The large adaptive radiation of the endemic native damselflies (Coenagrionidae: Megalagrion) in Hawai'i has received considerable attention and study since at least the 1880s. Endemic Megalagrion are in many ways reflective of a great loss because they are largely now found in remote upper headwater areas of streams, yet they also represent the hope of preserving highly diverse freshwater ecosystems found throughout the Hawaiian archipelago. The first two chapters of this dissertation examine the impacts of two differing taxa of introduced fish on Hawaiian Megalagrion, Poeciliidae (livebearers or mosquitofish family) and Salmonidae (trout). The effects of each fish species on native aquatic insects depended mainly on the invasive status ofeach group; for example, Chapter 1 (Englund 1999) examines the impacts of introduced poeciliids on native damselflies. Damselflies were completely eliminated on the island of 0'ahu wherever species in the highly invasive mosquitofish family were found, and only remnant populations were found in high elevations lacking introduced fish. Chapter 2 (Englund and Polhemus 2001) examines the impacts of the non-invasive rainbow trout (Oncorhynchus clarkI) on Megalagrion damselflies. Damselflies and all other native aquatic insects were not found to be harmed by trout in the uppermost elevations of Kaua'i streams where trout reproduce naturally, and even had more robust populations than in some nearby non-trout containing streams. The lack of impacts on native damselflies by a large, generalist predator such as rainbow trout pointed out a seeming paradox. Whereas the small but ubiquitous mosquitofish appears to have completely devastated native aquatic fauna wherever it has been introduced outside of its natural range, iii trout, because of their restricted range and smaller population sizes have had minimal, if any impacts on native invertebrates in Hawai'i. Because introduced fish species have caused either the extinction or severe range contractions of Megalagrion damselflies in Hawai'i, long-term monitoring of the remnant populations has become necessary to preserve these remaining populations. Chapter 3 (Englund 2001) provides a case study in both the monitoring and preservation of a remnant O'ahu damselfly population now found in only 95 m of fishless stream at the TripIer Army Medical Center. Chapter 3 also provides several harrowing examples of how this species was nearly been eliminated in the past 10 years through accidents and mismanagement. Not only are the endemic Megalagrion now missing from all lowland areas of O'ahu (with the exception of the TripIer population), lowland aquatic insect diversity throughout O'ahu is at a remnant status, and biodiversity surveys for native aquatic insects in the Pearl Harbor watersheds in Chapter 4 (Englund 2002) indicated a near absence of native aquatic insects in these freshwater habitats. Lower Pearl Harbor watersheds were documented to have lost many native aquatic insect taxa such as all native Heteroptera, damselflies, Coleoptera, and many Diptera species, while introduced insect species were abundant. A variety of conservation measures have been suggested to either restore or maintain the current levels of freshwater biodiversity in Hawai'i. In Chapter 5 (Englund and Filbert 1999), the case of significantly increasing and restoring stream flow in a formerly diverted stream was examined to determine whether this factor alone would lead to a restoration of native aquatic species. It was found that merely increasing stream flow by itself was not enough to rid the stream of any alien aquatic species, in fact, several new nonindigenous aquatic species became established after stream flows were increased. The results ofChapter 5 confirm that an integrated, balanced and possibly drastic approach will be required to maintain and preserve Hawai'i's native aquatic insect fauna. A wide-variety of conservation measures in the Hawaiian archipelago will be needed to maintain current biodiversity levels, and also hopefully restore native freshwater biodiversity in selected areas. iv To put the Hawai'i problem into perspective, a brief review of the impacts of invasive species on native insects in other tropical areas is provided in Chapter 6. This review chapter also provides a synthesis of the problem facing Hawaiian freshwater insects and other terrestrial arthropods in Hawai'i and elsewhere due to invasive species, and how the Hawaiian case study of invasive species impacts has many parallels to other vulnerable biotas. Finally, drawing on a mixed record ofpast mistakes and successes in Hawai'i and elsewhere, some potential practical conservation measures intended to preserve and restore endemic island aquatic insects are provided in Chapter 6. v LIST OF TABLES Table 1.1. Extinctions of Megalagrion species in surveyed 0'ahu aquatic habitats since 1936 8 Table 1.2. Remnant native Megalagrion species found in O'ahu streams, tributaries in parentheses,
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