Reef fishes of the Bird’s Head Peninsula, West Papua, Indonesia

A checklist of shallow (to 60 m depth) reef fishes is provided for the Bird’s Head Peninsula region of West Papua, Indonesia. The area, which occupies the extreme western end of New Guinea, contains the world’s most diverse assemblage of coral reef fishes. The current checklist, which includes both historical records and recent survey results, includes 1,511 species in 451 genera and 111 families. Respective species totals for the three main coral reef areas – Raja Ampat Islands, Fakfak-Kaimana coast, and Cenderawasih Bay – are 1320, 995, and 877. In addition to its extraordinary species diversity, the region exhibits a remarkable level of endemism considering its relatively small area. A total of 26 species in 14 families are currently considered to be confined to the region.


Introduction
The region consisting of eastern Indonesia, East Timor, Sabah, Philippines, Papua New Guinea, and the Solomon Islands is the global centre of reef fish diversity (Allen 2008).Approximately 2,460 species or 60 percent of the entire reef fish fauna of the Indo-West Pacific inhabits this region, which is commonly referred to as the Coral Triangle (CT).The first author has worked extensively throughout the CT for the past 36 years, with special emphasis on the ultimate identification of the "bullseye" of reef fish diversity.Data gathered by the authors over the past decade reveals that the Bird's Head Peninsula (Figure 1) of West Papua, Indonesia harbours the most diverse reef fish community in the CT (Allen, in preparation).
The present paper provides an annotated checklist of species and brief historical account of fish collections from this globally important location.
There are numerous complex factors, past and present, which are responsible for the amazing wealth of Bird's Head species.Perhaps the most important include an amazing level of habitat diversity, a strategic location lying at a biogeographical "crossroad" or "melting pot" that captures faunal elements from Indonesia, the Melanesian Archipelago, and Timor/Arafura Sea, and finally a complex geologic past highlighted by shifting island arcs, oceanic plate collisions, and widely fluctuating sea levels (Polhemus 2007).The Bird's Head Peninsula and surrounding waters has attracted the attention of naturalists and scientists ever since it was first visited by European explorers.The Raja Ampat Group and Waigeo Island, in particular, was the focus of early French visits by several vessels including L'Uranie (1818-1819), La Coquille (1823), and L'Astrolabe (1826).Consequently, approximately 70 fish species were recorded from the region primarily by Quoy andGaimard (1824 and1834), Lesson (1828Lesson ( -1830)), and Cuvier and Valenciennes .Waigeo is an important type locality for a variety of widespread species including the Blacktip Reef Shark (Carcharhinus melanopterus), Bluefin Trevally (Caranx melampygus), Bigeye Trevally (Caranx sexfasciatus), Semicircular Angelfish (Pomacanthus semicirculatus), and Sergeant Major (Abudefduf vaigiensis).
Following the early French explorations, most ichthyological activity was provided by Dutch researchers.The surgeon-naturalist Pieter Bleeker periodically received specimens from government agents and in 1868 published on a collection of Waigeo fishes that included 88 species.He added a further 12 species in subsequent papers.Albert Günther, the Curator of Fishes at the British Museum, recorded 28 species from the island of Misool, during the cruise of the "Curacao" in 1865 (Günther 1873).The Dutch ichthyologists Weber and de Beaufort were keenly interested in New Guinean freshwater and marine fishes and contributed to our knowledge of Raja Ampat fishes during the first half of the past century.The work of Weber (1913), in particular, was the most extensive effort on Raja Ampat fishes until recent times, and includes accounts of 117 species based on 748 specimens.These were obtained by de Beaufort during a visit to the East Indies in 1909-1910, and were mainly collected at Waigeo in the vicinity of Saonek Island and Mayalibit Bay.
Weber and de Beaufort and various co-authors including Koumans, Chapman, and Briggs reported an additional 67 records from Waigeo and Misool in the Fishes of the Indo-Australian Archipelago (E.J. Brill, Leiden;11 volumes published between 1921-1962).The Denison-Crockett South Pacific Expedition made small collections at Batanta and Salawati consisting of 29 species that were reported by Fowler (1939).The only other fish collection of note was that by Collette (1983) who recorded 37 species from mangrove habitats on Misool and Batanta.The first author made the first comprehensive underwater observations of Raja Ampat fishes during two brief visits in 1998-1999.Although freshwater fish research was the main focus, approximately 20 hours of scuba and snorkel diving yielded observations of more than 500 coral reef fishes.The first major survey of the islands was conducted in 2001, when the first author participated in a marine rapid assessment survey (RAP) organized by Conservation International (CI).A total of 45 sites were assessed during a 15-day period (Allen 2002).An additional 50 sites were visited in 2002 during a rapid ecological assessment (REA) conducted by The Nature Conservancy (TNC).The first author's combined CI-TNC effort raised the number of the known reef fishes of the Raja Ampats from about 236 to 1,102 (Allen, unpublished data).
Compared to the Raja Ampat Islands, the remainder of the Bird's Head Seascape has been largely ignored by previous researchers.In order to fill this gap in our knowledge CI organized two expeditions in 2006 to Cenderawasih Bay and the Fakfak-Kaimana Coast.Several additional trips were made by the authors to these two locations in 2007-2009.

Materials and methods
The current study is restricted to shallow reef fishes, which are herein defined as species that are entirely or mainly confined to coral reefs and intermingled substrata (sand/rubble patches, seagrass beds, etc.) less than 60 m deep.The 60metre depth limit is near the lower limit of routine scuba diving, and also approximates the limit of most reef-building scleractinian corals (Veron 2000).The list also includes several goby species that normally occur in mangrove estuarine habitats, but due to the closely intertwined mangrove and reef habitats that are characteristic of the Raja Ampat Islands, they are frequently seen during reef surveys.
The annotated checklist of species presented below is based on several sources: 1. Historical records prior to 1950 that were summarized by Weber and de Beaufort (1921de Beaufort ( -1962) )

Results
The currently known fauna of the Bird's Head Peninsula as reported in our checklist consists of 1,511 species in 451 genera and 111 families.Respective species totals for the three main coral reef areas -Raja Ampat Islands, Fakfak-Kaimana coast, and Cenderawasih Bay -are 1320, 995, and 877.The figure for the Raja Ampats is particularly impressive and is the most species ever recorded for an area of this relatively small size (estimated at 50,000 km 2 including the islands and surrounding seas).

Endemism
In addition to its extraordinary species diversity, the Bird's Head Peninsula exhibits a remarkable level of endemism considering the size of the area.A total of 26 species in 14 families are currently considered to be confined to the region (Table 1).We are confident of the endemic status of the majority of these species as they are conspicuous fishes that we have not seen elsewhere, despite our intensive collecting efforts in adjacent regions.DNA tissue samples of several additional species that are potential endemics are currently being analyzed by the authors.
The factors responsible for local endemism are no doubt complex and intimately associated with the geological history of the region.Polhemus (2006) provided a detailed summary of the tectonic history of the New Guinea region.Geologists are in general agreement that the core of the Vogelkop Peninsula is a section of the Australian craton that became detached from the main continental mass sometime in the Mesozoic (Hamilton 1979) and was sutured to the main mass of New Guinea approximately 15 million years ago.Although details of the biogeographical history remain poorly understood for most of the Vogelkop, there is compelling evidence that the inner half of Cenderawasih Bay was essentially isolated for a substantial period over the past five million years due to periodic sea-level lowering and effective closure of much of the bay by westward drifting island-arc fragments (Allen et al, in preparation).

Checklist of Fishes
The phylogenetic sequence of families appearing in the checklist follows Eschmeyer (1998) with slight modification that reflects recent taxonomic studies.The recent update of the Catalog of Fishes by Eschmeyer and Fricke (2009) was used extensively to check spellings, authorships, and dates of publication.References are cited for previous records from the Bird's Head region and include the page number and geographic location.Symbols and abbreviations used in the checklist are explained as follows: an asterisk (*) indicates a new record (i.e., previously unpublished in the peer-reviewed literature) for the Bird's Head

Figure 1 .
Figure 1.Map of the Bird's Head Peninsula region of West Papua, Indonesia.
, Museum National d'Histoire Naturelle (Paris), and Natural History Museum (London).The largest recent (1975-2008) collections from the area are deposited at the Western Australian Museum (Perth), National Museum of Natural History (Washington, D.C.) and the Research Center for Oceanography (Jakarta).The authors in conjunction with faculty and students from the University of Papua (Manokwari, West Papua) have also established a local research collection in the University.Species that are represented by voucher specimens at Jakarta, Perth, Washington, D.C., and Manokwari are indicated in the following checklist.
region; CB, FK, and RA indicate the species presence at Cenderawasih Bay, Fakfak/Kaimana region, and Raja Ampat Islands respectively; institutional acronyms (NCIP, UNIPA, USNM and WAM) indicate that voucher specimens are lodged at Pusat Penelitian Oseanografi (Research Center for Oceanography), Jakarta, University of Papua (Manokwari, Indonesia), United States National Museum (Washington, D.C.) and Western Australian Museum (Perth).New records that do not have a voucher specimen indicated are based on either underwater observations or photographs taken by the first author.Unidentifed species indicated as "sp." are primarily new taxa awaiting description.