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The Taxonomy and Biogeography of the Thick-Billed Flowerpecker Complex in Borneo

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The Taxonomy and Biogeography of the Thick-Billed Flowerpecker Complex in Borneo THE TAXONOMY AND BIOGEOGRAPHY OF THE THICK-BILLED FLOWERPECKER COMPLEX IN BORNEO FREDERICK H. SHELDON • WesternFoundation of VertebrateZoology, 1100 Glendon Avenue, LosAngeles, California 90024 USA ASSTRACT.--TheWestern Foundationof VertebrateZoology expedition to Sabah,East Ma- laysia(North Borneo)collected specimens and field data on Brown-backedFlowerpeckers (Dicaeumeveretti) and Thick-billed Flowerpeckers(D. agile)in 1982 and 1983. Dicaeumagile had not been known from Borneopreviously, and its presencethere changesthe biogeo- graphic assumptionsSalomonsen (1960) used to classifythe "thick-billed" flowerpeckers. Salomonsen'sD. everettiwas found to be a valid species,but his splitting of the Philippine StripedFlowerpecker (D. aeruginosum)from D. agilewas not justified.Dicaeum agile of Borneo is here placed in the race remotumwith the Malayan form of the species.The Borneanand Malayan races of D. everettiare grouped in the subspecieseveretti. Received 2 October1984, accepted12 February1985. THE thick-billed flowerpecker complex of In- D. everetti specimenswere thought to be the dia, SoutheastAsia, and the Philippines is list- first from Borneoin 27 years (Pfeifer 1961). In ed by Salomonsen (1967) as comprising three fact, no specimensof D. everettihave been col- species, the Thick-billed Flowerpecker (Di- lected from Borneo since 1901, even though caeumagile), the Brown-backedFlowerpecker (D. several museums have specimens labeled as everetti),and the Striped Flowerpecker (D. aeru- such. The two specieshave been confusedfor ginosum).Of thesespecies, only D. agileof India 85 years, largely becausethe scarcityand dis- and Sri Lanka is common and well documented persionof specimenshave led to oversightsin (Ali and Ripley 1974). In other parts of South- the taxonomic literature. east Asia, the three speciesrarely have been The Western Foundation field data and the recorded, and their statuses are uncertain. This new and old specimenmaterial permit a criti- is especiallytrue in Sundaland,the areasof land cal evaluation of Salomonsen's (1960) classifi- such as Malaya, Borneo, and much of the In- cation of the thick-billed flowerpecker com- donesianarchipelago associated with the Sun- plex. His controversial separation of D. everetti da shelf. Dicaeumeveretti in Malaya is known from D. agile(sensu Mayr and Areadon 1947) is from only four specimens.Dicaeum agile in Java strongly supported, but questions are raised is known from only two specimens.Dicaeum about some of his racial taxonomy and about everettiin the Natuna Islands and D. agile in the schemeproposed to explain the radiation Sumatra are known from only one specimen and speciationof the thick-billed flowerpeck- each. ers in the Sunda and Philippine islands. In 1982 and 1983, the Western Foundation of VertebrateZoology expeditionto Sabah,East TAXONOMIC HISTORY Malaysiacollected both D. agileand D. everetti. Salomonsen (1967) divided the thick-billed The D. agilespecimens were thought to be the first from Borneo (Smythies 1981). However, flowerpeckercomplex into three speciesbased museumstudy subsequent to the expeditionhas on his 1960 review: D. agile,encompassing the shown that specimensof D. agilehave existed racesagile, zeylonicum, modestum, remotum, atje- from Borneo, in small numbers, since 1898. The hense,finschi, tinctum, and obsoletum;D. everetti, comprising everetti, bungurense,and sordidum; and D. aeruginosum,composed of affine,aerugi- *Present address:Department of Biology, Yale nosum, and striatissimum. He considered D. ev- University, P.O. Box6666, New Haven, Connecticut eretti and D. aeruginosumto constitute a super- 06511 USA. species. The only sympatric forms in the 606 The Auk 102: 606-612. July 1985 July 1985] BorneanThick-billed Flowerpeckers 6O7 T^BLE1. Rangesand characteristicsof the subspeciesof the thick-billed flowerpeckercomplex. Wingb Bill Subspecies Locality Dorsum' Underparts Tail (mm) shape agile India, Pakistan Olive-gray Faint streaks Bright spots 55-65' Thick zeylonicum Sri Lanka Dark olive-gray Moderatestreaks Bright spots 55-57' Thick modestum Assam-Indochina Olive-green Moderatestreaks Bright spots 57-61 (13) Thin remotum Malay Peninsula, Olive Moderate streaks Faint spots 57-63 (24) Thin Borneo atjehense Sumatra Olive Moderate streaks Faint spots -- Thin a finschi Java Olive Moderate streaks No spots 62 (1) Thin tinctum Sumba, Lesser Olive Faint streaks Faint spots 54-59 (9) Thick Sundas obsoletum Timor Grayish olive No streaks Faint spots 53-62 (11) Thick everetti Borneo Olive-brown No streaks No spots 54-60 (14) Thin bungurense Great Natuna Olive-brown No streaks No spots 63• Thin Island sordidum Malaya, Riau Olive-brown No streaks No spots 59-61 (3) Thin Archipelago affine Palawan Grayisholive Heavy streaks Bright spots 61-66 (7) Thick aeruginosum Mindoro, Negros, Olive-green Heavy streaks Faint spots 62-68 (8) Thick Cebu, Mindanao striatissimum Luzon, Sibuyan Brownisholive Heavy streaks Faint spots 64-65 (2) Thick aColor terminologyfollows Smithe (1975). bSample sizes are given in parentheses,except in instanceswhere measurementsare taken from indepen- dent sources. ½Measurements from Ali and Ripley 1974. a Measurements from Chasen 1939b. e Measurements from Chasen 1934. complex known to Salomonsenwere D. a. re- cal to the Borneanforms of the speciesD. ev- moturnand D. e. sordidurnof Malaya. The distin- erettiand D. agile,respectively. guishing features of the three speciesare as Chasen(1935, 1939a)grouped the Sundaland follows:D. agilehas a comparativelypale olive forms, sordidurn,everetti, and bungurense,with or gray dotsurn,streaked underparts, and (ex- the Indian race, agile,in the species"Piprisoma" ceptfor finschi)white spottingat the tail tip; D. agile(Piprisorna being a subjectivesynonym of everettihas a browner dorsum, no underpart Dicaeurn,with type speciesagile), and he united streaking,and no tail spotting;and D. aerugi- remotum and modesturn in P. modesturn. Al- nosurnhas the same characteristicsas D. agile though it attempted to recognize the distinc- but generally is larger and darker (Salomonsen riveness of the sordidum-everetti-bungurense 1960). The ranges and characteristicsof these group, this arrangement was counterintuitive. races are summarized in Table 1. It ignored the dissimilaritybetween the Indian Until Salomonsen's review of Dicaeum, the and Sundalandbirds, and it createda disjunct member racesof the thick-billed flowerpecker distribution in which the continental South- complexhad beenclassified in a varietyof ways. east Asian P. modestumseparated the Indian For example,Sharpe (1885) included the 5 then- from the Sunda forms.Had Chasenunited ag- known formsas full speciesof the genusPrio- ile, remotum,and modesturnin one speciesand nochilus.Oberholser (1923) split the group into sordidurn,bungurense, and everettiin another, his 3 genera and 5 species.Mayr and Areadon classification of these taxa would have made (1947) transferredthe group from Prionochilus more morphologicaland geographicalsense. As to Dicaeumand lumped all the racesinto a sin- they were, Chasen'sgroupings did more harm gle species,D. agile.To understand the rela- than good becausethey were believed to be the tionships of the Borneanmembers of the com- product of over-splitting and, consequently, plex, the treatmentsgiven the Malayan forms, were later ignored or attacked in reviews. As sordidurnand rernoturn,must be emphasizedbe- further complications,the few existing speci- causethese Malayan birds are virtually identi- mens of sordidumand bungurense,deposited in 608 FREDERICKH. SHELDON [Auk, Vol. 102 TABLE2. Specimenrecords of Dicaeumagile collected in Borneo.a Num- Altitude bet Date Locality Habitat (m) Museum b 3 Dec 1898 Upper Mahakam R. (Kaliman- -- -- RHNL, USNM tan) 4 Dec 1947- Kelabit Plateau, Sarawak 2ø forest, fruit 1,000 AMNH, SM Jan 1948 trees 1 28 Sept 1952 Long Tinkalat, Tinjar R., Sara- 2ø forest 300 SM wak 7 Sept 1956 Long Peso,Kajan R., eastern 2ø forest, village 150-200 MHNP Borneo fruit trees 4 July 1982 SabahSoftwoods, Brumas, Sa- Albiziagroves 250 WFVZ bah l Three poorly prepared specimens(2 in the PMNH and 1 in the SM) collected 9 October 1952 on Mt. Matang, Sarawakhave been omitted from this list becausetheir identities are uncertain.They could be either D. agileor D. everetti. bRHNL = Rijksmuseumvan Natuurlikijke Historie, Leiden; USNM = U.S. National Museum;AMNH = American Museum of Natural History; SM = Sarawak Museum; MHNP = Museum d'Histoire Naturelie de Paris;WFVZ = WesternFoundation of VertebrateZoology; PMNH = PeabodyMuseum of Natural History, Yale University. the Raffles Museum in Singapore, were inac- morphological counterparts in the Indonesian cessibleduring World War II when the major archipelagoand on the Asian mainland. No review work was undertaken, and Chasen, the simple pattern of D. agile radiation could ac- only activeornithologist familiar with the Ma- count for this anomaly. Salomonsendecided layan birds, was killed. that the Philippine forms were derived from Deignan (1945), apparently in responseto D. everettion Borneo,but he placed them in a Chasen'staxonomic arrangement, condemned separatespecies (D. aeruginosum)because of their the splitting of "Piprisoma"on the Asiatic main- distinctivehess from D. everetti. To reflect the land and lumped all mainland forms in P. agile logic of his radiation scheme,however, Salo- modesturn,including by implication sordidurn. monsen chose to unite D. everetti and D. aeru- Then followed a series of important works cit- ginosumas a superspecies. ing
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