The Auk 110(3):529-540, 1993

USE OF AN EXOTIC BY BORNEAN LOWLAND

SHAIBAL S. MITRA • AND FREDERICK H. SHELDON 2 1Committeeon EvolutionaryBiology, University of Chicago,Chicago, Illinois 60637, USA; and 2Departmentof Ornithology,Academy of Natural Sciences, 1900 BenjaminFranklin Parkway, Philadelphia, Pennsylvania 19103, USA

ABSTRACT.--DuringMay to July 1982,we surveyedbirds in primary forestand in different- aged grovesof the exotictree Albiziafalcataria at SabahSoftwoods, a lowland tree plantation in Sabah,East (formerly British North ).We found that the Albiziawas in general attractive to many native birds. About 60% of primary-forestspecies used the plan- tation, and the frequencyat which individuals were observedin the oldestgroves was almost twice that of nearby primary forest. The Albiziaattracted birds because,as an extremely fast- growing legume with thin leaves,it permitted the rapid development of a well-structured secondaryforest. It alsowas heavily infestedwith pestinsects, which providedan abundant food source. Despite its apparent richness,however, the Albizia lacked several important featuresof natural forest(e.g. canopyfruits and nestholes). As a result,some primary-forest groupswere poorly represented(e.g. large canopy frugivores and flycatchers)and others could make only limited use of the plantation(e.g. ).In addition, the Albiziais expectedto loseits diversityas the plantationas a whole ages.Many of the plantationbirds were transientsfrom nearbyforest that visited daily to feed, and someprobably had been displacedby intense logging. The number of daily transientsshould decreaseas primary forestrecedes due to logging and development.Refugee diversity should sufferfrom attrition as the plantation is cropped and predation and age take their toll. Received9 March 1992, accepted20 November1992.

As THERESULT of development,many tropical forestcommunities (e.g. as a sourceof food or rain foreststhroughout the world face rapid competitors)?Answers to such questionsare and generallydestructive change. One form of neededbecause they not only bear on issuesof tropical land development that has become in- tropicalcommunity ecology, but they relatedi- creasinglypopular in the last 20 years is the rectly to practicalproblems of plantation man- planting of exotic, fast-growingtrees for chip agementand the conservationof organismal and pulp production.Tracts of primary rain for- diversity.Although plantationsare inevitable, estin ,Africa, and the Neotropicsthat have they canbe developedto preserveand exploit been exploited for timber subsequentlyhave natural .For example, that been clearedand replacedwith plantationsof are intercalatedwith or surroundedby natural treessuch as Eucalyptus deglupta, Gmelina arborea, forestalmost certainly benefit from biological Albiziafalcataria, Acacia manglum, Pinus carabaea, control of infesting pests(e.g. insectsand ro- P. oocarpa,and Anthocephaluschinensis. Unlike dents)by forest-dwellingpredators. The useof agricultural crops, such plantations do not re- forestconservation in plantationmanagement quire good soilsor intensive management(e.g. and the influenceof plantationson forestcom- of understorygrowth). Thus, they offer an at- munitieswill dependupon many potentialfac- tractive alternative for sustainedyield of rev- tors, including the types,ages, and mixture of enues. that are cultivated, crop rotation, prox- Although the prospectof sterile monocul- imity of primaryforest, plantation physiogno- tures replacing diverse, rich forestshas raised my (e.g. presence of streams, cliffs, etc.), and concerns(e.g. Andrews 1973, Wells 1985, Shel- timing and complexityof pestinfestations. don 1986), the ecologicalconsequences of this To understand elements of the interaction be- type of land use have received little attention, tween plantation and forest bird communities, and many basic questions remain to be an- we censused birds at Sabah Softwoods Planta- swered. In particular, how will native tion in Sabah,East Malaysia (formerly British and respond to plantations,and how North Borneo)during May to July 1982. Ours will plantationcommunities influence adiacent is one of only a few suchstudies that have been

529 530 MITRA^NO SHELDON [Auk, Vol. 110

TABLE1. Dates and times of surveys. of our survey, about 25,000 ha of a 61,000-ha concessionhad been replanted,principally with No. the exotic treesAlbizia falcataria, Acacia mangium, Plot type Plot sur- Early/ (age) no. veys Dates late" Gmelinaarborea, and Eucalyptusdeglupta. Anoth- er 3,300ha were planted with agriculturalcrops Primaryforest 1 2 10, 15 June 2/0 such as cocoa and coffee (Duff et al. 1984). Cir- 2 ! 10 June 0/! cumscribedstands of primary forest persisted Albizia about 1 to 2 km from the plantation in 1982, !975(7years) ! 8 28 May-25 July 4/4 2 3 29 May; 8, 14 June !/2 and logging was active at distancesfrom 5 to 3 1 20 July 1/0 20 kin. !977(5years) ! 2 30 May; 4 June !/1 We were interestedin assessing(1) the extent 2 3 2, 3, 12 June 2/! to which primary-forestbirds used the planta- 3 2 22 July 1/ 1 tion, (2) how the bird community in the plan- 4 4 24, 30 July 2/2 tation developed through time, and (3) ecolog- !979(3years) ! 6 3, !3 June;17, 3/3 ical factors that played a role in this 29 July 2 4 7 June;18 July 2/2 development. To this end, we censusedbirds !981 (! year) ! 5 6, !1 June;23 July 3/2 in different age groups of one speciesof plan- 2 3 6, 9, 1! June 1/2 tation tree, Albizia falcataria.We selected this All surveys 44 28 May-30 July 23/21 speciesbecause preliminary observations(e.g. "Early (0600-0900)/late(0930-1230). A. G. Davies unpubl. manuscript) indicated that Albiziagroves were more attractiveto birds than were those of other plantation trees. conducted in Asia. Others include Davies (un- Locally called "Batai," Albiziafalcataria is na- publ.manuscript), Davies and Payne(1982), Duff tive to the Molucca Islands, New Guinea, New et al. (1984), and Stuebing and Gasis (1989), Britain, and the Solomon Islands, but has been which were surveysmainly of mammalsin Sa- introduced to most parts of Malaysia (Cockburn bah Softwoods, and Wilson and Johns (1982), 1976).Synonyms are A. falcataand A. moluccana; which examined a restricted set of mammal and sometimesthe speciesis placed in the bird speciesin a similar plantation in east Ka- Serianthes(Mabberley 1987).Albizia is a popular limantan (Indonesian Borneo). These investi- plantation tree becauseit grows fast,gaining as gations complement data collected in recent much as 6 m in height and over 12.5 cm in girth years on the effectsof logging on Malayan and per year in its first few years.It is alsoa legume Bornean bird and mammal communities (e.g. that improves soil quality (Cockburn 1976, K.- Wilson and Wilson 1975, Johns 1983,1986,1989, C. Tan pers. comm.). Wong 1985,Lambert 1992). Beehler et al. (1987) also surveyed birds in several plantation and METHODS man-disturbed forest in the Eastern Ghats of . However, the age and config- Data were collectedin each of the following five habitatsor "plot types":one-, three-, five-, and seven- uration of the forestsand plantations they stud- year-old Albizia,and primary forest (summarizedin ied (including coffeeand groves)differed Table !). To comparerichness and diversity, we com- from the newer exotic-treeschemes of Malaysia piled accuratelists of the kinds and numbersof birds and . present in each plot type and converted these into Sabah Softwoods is located in the Kalabakan frequency values. For each plot type we surveyed Forest Reserve in southeastern Sabah (4ø37'N, replicate "plots." To ensure consistencyamong plots, 117ø45'E,altitude < 300 m). The plantation was we selected areas with similar altitudes, terrains, and begun as a joint venture by the SabahFoun- distancesfrom streamsand primary and logged forest. dation and North Borneo Timber Company in In each plot, a quadrat trail of 300 m on a side was cut; one of the four sides consisted of an access road. 1972. Prior to development, most of the region To census,we walked the trails,allowing 3 h for each had been covered with primary forest, domi- quadrat.Surveys were conductedat two differenttimes nated by trees of the family of day: 0600 to 0900 and 0930 to 1230 standardtime. (Wood and Meijer 1964, Whitmore 1984). This We consideredall surveys of a given plot to be rep- forest was heavily logged in the early and mid- licates,but endeavoredto complete equal numbers of 1970s, and a large area was clear-felled and early- and late-morningsurveys for eachplot and plot burned to establish the plantation. At the time type. Every bird that was seen or heard was noted. July 1993] BorneanPlantation Birds 531

TABLE2. Feeding-guild classificationwith examples.

1 Raptors (r): Hawks and owls. 2 Terrestrial insectivores (ti): , some babblers, and thrushes. 3 Bark-gleaninginsectivores (bgi): Woodpeckers,, and somebabblers. 4 Foliage-gleaninginsectivores (fgi): Many babblersand warblers. 5 Sallying substrate-gleaninginsectivores (ssgi): , some , broadbills, and . 6 Sallying insectivores(sai): Flycatchers. 7 Nectarivores (n): , , Loriculusgalgulus, and Dicaeumtrigonostigma. 8 Arborealfrugivores (af): Most pigeons,most barbets, Calyptomena, Irena, crows, mynas, and mostbulbuls. 9 Arboreal frugivores/faunivores(aft): and Megalaimachrysopogon. 10 Terrestrial frugivores (tf): Phasianidsand . 11 Aerial insectivores(ai): Swifts, swallows, and nightjars. 12 Miscellaneous(m): Somekingfishers, a ,predatory waterbirds, and rails.

The censusingtook placefrom 28 May to 30 July 1982. make pairwise comparisonsbetween plot types on During this period, no northern migratory species the basisof their food-guild frequencies: were present in the area. Table 1 shows the datesand times of all surveys.The Appendix lists the species and numbers of individuals recorded. In addition to data recordedduring formal surveys, we compiled many distributional recordswhile col- where x, is the frequency of guild i in plot type x, y, is the frequency of guild i in plot type y, and s is the lectingbirds in primary forestand other Albiziagroves number of guilds. This index varies from 0 for cases during March and April 1977, and May to July 1982. when sets share no common members to 1 when sets These records have been separated from the survey are identical. data, but we have used them to obtain lists of the total To portray the toleranceof bird speciesto Albizia, number of speciesfor the plot types.Nonsurvey spe- cies are marked with a "P" in the Appendix. To dis- we computedT, the inverseof Simpson's(1949) mea- tinguish between survey data and the combination sure of concentration(Lovejoy 1974): of surveyand nonsurveydata, we use the terms"spe- T = 1/• P? (3) cies surveyed"and "total species." Speciesrichness in each plot type is expressedsim- where P, is the proportion of individuals of a species ply asthe number of species.Abundance or encounter found in i. P, is calculated as the encounter frequency was computedby dividing the total num- frequencyin habitat i divided by the sum of encoun- ber of individuals recorded on a survey or group of ter frequenciesin all five plot types.The T-valuesare surveysby the number of survey hours. To portray listedfor eachsurveyed species in the Appendix.They the distribution of speciesacross different environ- range upward from 1.00for the leasttolerant species. ments, we counted the number of speciesrestricted to each plot type (unique species)and the proportion of total individuals accountedfor by the five com- RESULTS toonestspecies in each plot type (dominant species). Diversity in eachplot type is expressedby the Shan- Our 44 surveys yielded 4,889 records of 140 non-Wiener index (H'): speciesand 6,741 individual birds. Most of the recordswere basedon identification by song or H'= -• p•logp,, (1) call (65%). Only 27% were basedon sightings, and 8% on both callsand sightings.Nonsurvey where p, is the proportion of individuals in speciesi recordsyielded data on an additional 60 species. and n is the number of speciessurveyed. Of the 200 total species,59 were found only in To evaluatethe more obviousecological differences primary forestand 38 only in the Albiziagroves. distinguishinghabitats, we divided bird speciesand About 50% of the 38 speciesrestricted to the individuals into categoriesbased on their feeding Albiziawere obligate secondary-forestor open- guildsand thencompared relative frequencies of these land species.Of the 162 speciesrecorded in pri- groupsin each plot type. The 12 feeding guilds were basedon their main food type (Table 2), rather than mary forest, 103 were also found in the Alb&ia a combination of foods, as is often done (Karr 1980, groves (see Appendix). Johns1986, Lambert 1992). Thus, we categorizedsun- In general, within the plantation, plot types birds as nectarivores,even though they also eat in- varied predictably in respectto speciesrichness, sects.We used Horn's (1966) index of overlap (C) to encounter frequency, uniqueness, dominance, 532 MITRA AND SHELDON [Auk, Vol. 110

TABLE3. Summary of data for Albizia groves and primary forest.

Alb&ia grove' Primary 1975 1977 1979 1981 forest (7 years) (5 years) (3 years) (1 year) Species surveyed 69 98 73 56 38 Total speciesb 162 122 92 63 45 Survey records 234 1,651 1,381 1,123 5OO Individuals surveyed 338 2,200 1,792 1,591 82O Individuals/observation 1.44 1.33 1.30 1.42 1.64 Individuals/hour c 37.6 61.1 49.8 53.0 34.2 Unique speciesa 59 5 0 5 2 Dominant species* 26.6 30.4 43.7 61.0 75.0 Diversity (H') 3.823f 3.867 3.334 2.915 2.360 • Plot type (age). t'Includes species recorded outside formal surveys. ßEncounter frequency. dNumber of speciesrestricted to given plot type (38 speciesrestricted to Albiziagroves as a whole). "Proportion of total individuals accountedfor by five commonestspecies in eachplot type. • Diversityfor primary forestartifically depressed because of relativelysmall sample size. and diversity (Table 3). The number of species mary forest were artificially depressedfor two and diversity increasedwith plot age. Encoun- reasons:(1) detecting birds was more difficult ter frequency increasedbetween youngest and in primary forest than in the plantation (this oldest plots, despite similar values for three- was a consequenceof the higher canopy and and five-year-old plots. There was no obvious greater structural complexity of natural forest) pattern in individuals per observation,and no and (2) we surveyed the forest formally only reason to expectany. Similarly, the number of three times. To achieve results comparable to unique specieswas not related to plot age.With the plantation data, we needed more primary- the exceptionof a few open-country or wetland forest surveys. Nevertheless, the total number birds (e.g. Gallirallusstriatus, Rallina fasciata, To- of speciesrecorded in the primary forest (162), dirhamphuschloris, and Lonchuramalacca), the which was compiled over a much longer period speciesunique to given plantation plot types than the plantation survey, is reasonably ac- were uncommon, patchily distributed, or dif- curate. This number is only slightly less than ficult-to-identifyforest birds that probablywere that of nonmigratory, closed-canopyspecies overlooked in our primary forest work (e.g. Spi- found in the well-studied, nearby Ulu Segama zaetusalboniger, minutillus, region (e.g. Lambert 1992, A.D. Johns and F. archipelagicus, raffiesii, agile, D. H. Sheldon unpubl. manuscripts). concolor,and Arachnotherarobusta). Dominance To check for bias in our sampling methods, by individual speciesdecreased with plot age we examined variation among plots within plot as the proportion of abundant early colonists types.We found that plots varied significantly and openland speciesdecreased (e.g. Macronous in numbers of species recorded per survey gularis,Prina fiaviventris, Orthotomus sericeus, and (ANOVA, P < 0.025). Such variation may in- Pycnonotusgoiavier). This pattern prevailed de- dicate ecologicalheterogeneity among plots, but spite the increasing abundance of some domi- alsoit appearsto be influencedby survey dates. nant forest speciesin the older plots (e.g. Di- Even though we tried to sample over similar caeumtrigonostigma, simplex,and datesfor plot types,the averagedates of surveys particularly Arachnotheralongirostra). for individual plots within plot types typically Trends noted acrossAlbizia plot types seemed were different (Table 1). The differencesamong to hold for the primary forest in the casesof plots over time indicate not only an increasein species richness (total species) and relative number of species surveyed, but also an in- dominance of species.In contrast,trends in di- crease in encounter frequencies. Possible ex- versity and encounter frequency did not extend planations for these increases are growth in to the primary forest.We believe, however, that populationsfollowing nesting,or that a change the numbers of species and individuals sur- in logging intensity causedgreater dispersalof veyed and diversity of speciesfound in the pri- birds from forestedareas. It seemsmore likely, July1993] BorneanPlantation Birds 533

TAI•LE4. Feedingguild frequencies(in percent).Percent species calculated as number of speciesrepresenting eachguild divided by total species(including nonsurveyspecies) in a given plot type. Percentindividuals calculatedas number of individualsrepresenting each guild dividedby totalnumber of individualssurveyed in plot type.

Albiziagraves a Primary forest 1975 (7 years) 1977 (5 years) 1979 (3 years) 1981 (1 year) Individ- Individ- Individ- Individ- Individ- Guild uals Species uals Species uals Species uals Species uals Species lr 0.3 6.8 0.5 5.7 0.1 4.3 0.2 4.8 0.0 0.0 2ti 5.3 8.0 6.9 7.4 4.3 5.4 3.0 3.2 2.4 4.4 3bgi 7.4 9.9 1.5 9.0 0.8 5.4 0.6 3.2 0.7 4.4 4 •i 46.2 21.6 43.8 26.2 52.9 31.5 57.2 31.7 60.4 28.9 5ssgi 7.1 12.3 3.4 9.0 0.8 6.5 0.1 1.6 0.0 0.0 6sai 12.4 9.9 5.6 4.9 4.5 6.5 2.8 3.2 1.8 8.9 7 n 4.7 4.9 16.4 10.7 11.6 12.0 11.6 17.5 2.0 13.3 8af 10.7 16.7 19.6 17.2 23.2 16.3 23.6 17.5 27.7 24.4 9aft 5.3 3.1 0.6 1.6 0.2 1.1 0.0 0.0 0.0 0.0 10tf 0.6 3.1 1.1 1.6 1.7 2.2 1.6 3.2 5.0 2.2 11ai 0.0 3.7 0.7 5.7 0.0 7.6 0.1 9.5 0.0 11.1 12 m 0.0 0.0 0.1 1.6 0.0 1.1 0.2 6.3 0.0 2.2

Plot type (age). however, that our skill at detectingbirds simply cies in terms of quantity and evennessof dis- improved as we conducted more surveys. persion in the five plot types were the small Feeding-guild frequenciesare listed in Table babblerStachyris ruJ•rons (T = 4.67)and the large 4. The degree of domination by the two most Copsychusrnalabaricus (4.60). They were prominent guilds (foliage-gleaning insecti- followed by Cacornantisrnerulinus (3.97), Ortho- vores and arboreal frugivore-faunivores) tornussericeus (3.94), Chloropsissonnerati (3.85), showed an inverse relationship to plot age. To- Stachyriserythroptera (3.82), Pellorneurncapistra- gether,these guilds accountedfor the following turn (3.78), and Orthotornusruficeps (3.73). With percentagesof individuals and species:57% and the exception of the generalist feeder C. son- 38% for primary forest;63% and 43% for seven- nerati,all of thesespecies are insectivores.Most year-old plots; 76% and 48% for five-year-old of the tolerant speciesare insectivoresor gen- plots;81% and 49% for three-year-oldplots; and eralists.

88% and 53% for one-year-old plots. The next- DISCUSSION most-prominent plantation guild was nectari- vores, which when added to the previous two The impressive diversity of birds found in guilds yielded: 62% and 43% for primary forest, Sabah Softwoods'Albizia, especially the older 79% and 54% for seven-year-old plots, 88% and 60% for five-year-old plots, 93% and 67% for TABLE5. Degree of feeding-guild overlap between three-year-old plots, and 90% and 67% for one- bird communitiesof five plot typesas indicated by year-old plots. Harn's (1966) index. Cells in upper-right basedon Horn's indexes of ecologicaloverlap are list- individual counts, and those in lower-left based on ed in Table 5. This index provides a measureof speciescounts. the ecologicalsimilarity of plot typesunder the Albizia gravea assumption that the degree of similarity be- tween different bird communities(compared Pri- 1975 1977 1979 1981 pairwise in terms of their feeding guild fre- mary (7 (5 (3 (1 forest years) years) years) year) quencies)reflects ecological similarity between the habitats in which the communities are found. Primary forest -- 0.935 0.931 0.915 0.892 In general, the index indicated that plot types 1975 (7) 0.961 -- 0.979 0.962 0.914 of similar age are most similar in bird com- 1977 (5) 0.914 0.979 -- 0.997 0.976 munity structure. 1979 (3) 0.813 0.929 0.968 -- 0.984 Habitat tolerancesare listed for all surveyed 1981 (1) 0.841 0.921 0.955 0.958 -- speciesin the Appendix.The mosttolerant spe- Plot type (age). 534 MITRA AND SHELDON [Auk, Vol. 110

TABLE6. Averagecanopy heights and basalareas of understory, which became more herbaceousin Albizia(K.-C. Tan pers. comm.). three to five years.After five to sevenyears, the Albizia acquired an impressive undergrowth, Canopy height Basalarea Plot type (age) (m) (m2/ha) which was dominatedby woody plants,had a canopy of its own at about 5 m, and featured 1981 (1 year) 5.5 4 many plants in flower and fruit (see Tables 6 1979 (3 years) 17 18 1977 (5 years) 24 28 and 7). 1975 (7 years) 28 32 Changesin the bird communityfrom youn- ger to older grovesof Albiziaappeared to mirror, in part, the reassemblageof the arthropodcom- groves,indicates that many native forest birds munity. Arthropods would be expectedto re- made at least some use of this artificial habitat. turn as the temperatureand humidity in the Four attributesappear to be responsiblefor the forestbecame buffered by canopydevelopment attractivenessof Albizia:(1) By virtue of its rapid (Wong 1985, D. R. Wells pers.comm.). Foliage growth and thin canopy, Albizia provided the gleaners in one- and three-year-old plots (e.g. spaceand light for the development of a sub- Cacomantismerulinus, Macronous gularis, stantial secondary forest replete with erythroptera,S. rufifrons,Prinia fiaviventris, tailor- and food. (2) The Albiziathemselves were birds, and some sunbirds and spiderhunters) infestedwith caterpillars,which attractedbirds. probablysubsisted largely on the lepidopteran (3) The plantation was adjacentto primary for- larvae that form the first wave of invad- estand near to areasof active logging and, thus, ers. In the later stagesof Albiziagrowth, there had a ready source of avian forest species.(4) was an influx of larger faunivores,such as mal- The plantation as a whole (including non-A/- kohas, trogons, Ceyx erithacus,broadbills, and bizia groves) was young; thus, there may not drongos. Other kinds of insectivores also in- have been enough time for birds displacedby creasedin older growth, including bark-glean- logging to be depleted, and there was still mi- ing woodpeckers,terrestrial-feeding pittas, and crohabitat structure in the plantation (e.g. foliage-gleaningcampephagids (e.g. Coracina) stumpsand logs)left from clearingprimary for- and babblers. est. Raptors (including owls) were relatively Secondaryforest and prey communitydevelop- common in Albizia, probably in response to ment.--The rapid successionaldevelopment of abundantprey. In addition to birdsand , Albizia groves accounted in large measure for their potential prey included large numbersof the diversity of the plantation'sbird commu- mammals. Stuebing and Gasis (1989), for ex- nity. In general, the youngestplots had a grassy ample, found that six- and seven-year-oldAl-

TAI•LE7. Descriptionof changesin understorycomposition in Albiziagroves (K.-C. Tan pets. comm.).

One-year-old plots Grasses:Paspalum conjugatum, Imperata cylindrica, Eleusine indica, Ottochloa nodosa, etc. Herbaceous:Mostly Eupatoriumodoratum. Ferns: Nephrolepsisspp. Climbers:Mainly Mikania spp. Three-year-old plots Grasses and ferns: Less common than above. Climbers: As above. Herbaceous:Same as above, except Eupatoriummore common. Woody plants: Solanumwrightii, Melastoma malabathrium. Five-year-old plots Grasses: Less common than above. Woody plants: More common than in three-year-oldplots, including, Credrelaglaziorii, Trema tomentosa, Leea indica,Dillenia suffruticosa. Seven-year-oldplots Woody plants:Even more prominent, including Macarangagigantea, Macaranga hypoleuca, Mallotus panicula- tus, Anthocephaluschinensis. July1993] BorneanPlantation Birds 535 bizia (the only ages they surveyed) contained from surrounding areas. Vagile and nomadic unusually large numbers of the nocturnal rat birds, suchas hornbills, Loriculusgalgulus, bul- Maxomyswhiteheadi, as well as the treeshrews buls, Zosteropseveretti, , sunbirds, Tupaiaglis and T. tana. Graculareligiosa, and Corvusenca would be ex- The successionalchanges described above pectedto migrate daily from forestedareas. Fur- were accompaniedby a decreasein someearly thermore, a proportion of the plantation species colonists.The grasslandspecies Centropus ben- probablyhad been displacedby logging of their galensis,for example, though common in the home forest. one- and three-year-oldplots, declined and dis- The age of the plantation as a whole also appeared thereafter. Pycnonotusgoiavier, Prinia should influence the richnessand diversity of fiaviventris,and Lonchurafuscans were common its bird community. In 1982, when the survey throughout the Albizia plots, but decreasedin was conducted,the plantation was young. As frequency in the older plots. Somespecies that the surrounding primary forest recedesdue to were abundantin young plot types(e.g. Macro- logging and development,visits by daily mi- nousgularis and Orthotomussericeus) were less grants and infusions of refugee speciesshould numerousin older plots,apparently having been decline. Moreover, at the time of the survey, replacedby their congeners(e.g.M. ptilosusand the plantation groves were still littered with O. ruficeps).It is possible that this pattern of stumps and logs left from clearing of primary congenericreplacement was in some casesan forest. These stumps could be used as nesting artifact of plantation development. It has been sites by such taxa as woodpeckersand Gracula suggested,for example,that M. gularismay have religiosa,and feeding sitesby bark gleaners,in- retreated to the canopy in older groves where cluding woodpeckers,some babblers (e.g. Stach- it would have been more difficult to observe. yris maculataand Pomatorhinusmontanus), and While this retreat may have occurred,we prob- Sittafrontalis. Through time, however,such lin- ably did not miss this speciesin older groves gering microhabitatsand their dependentbird given that most of our recordsof it were based specieslikely will disappear.Finally, at the time on calls, not sightings. of our study, the Albizia had not yet been Pest insects.--Important attractants for many cropped.Thus, the effectsof harvestingon the birds were the lepidopteran larvae that infested bird communitydid not have time to take effect. the Albizia leaves (e.g. the pierid butterfly Eu- Birdgroups not found in Albizia groves.--In the rema blanda).The existence of this superabun- plantation, we failed to record speciesrepre- dant food supply requires that our food-guild senting the bulk of several families, guilds, or data be interpreted with caution. Many birds, subguilds.Some of thesesimply may have been including flowerpeckersand sunbirds,that are overlooked. ,for example, were diffi- normally characterized as frugivores and nec- cult to record unlesscalling. Members of other tarivores,were feeding on caterpillars.We ob- groupswere rare even in the surrounding forest served,for example,during nonsurveyperiods (e.g. the trogons Harpactesorrhophaeus and H. flocks of 10 to 20 Dicaeumagile gleaning cater- oreskios).Other groups may have required spe- pillars in the seven-year-oldAlbizia canopy. Be- cial subhabitats(e.g. large rivers and cliffs) that causebirds assignedto different guilds actually were not found on the plantation. An obvious may have had similar diets, trophic diversity in example would be the large piscivorous king- the plantation may not be as great as indicated fishers,which were notably lacking. Regardless in Tables 4 and 5. of such extenuating circumstances,several for- Plantationlocation and age.--Even though the est-dwelling groups were underrepresentedin Albizia groves at Sabah Softwoods in 1982 the plantation, apparently becauseof funda- showeda remarkablediversity of birds, sucha mental properties of the Albizia groves them- finding should not be expectednecessarily of selves. Albiziaplantations in other areasor even of Sa- In the absenceof large canopy fruits (e.g. bah Softwoods in 1993. The proximity of the strangler figs), there was a dearth of large can- plantation to primary forest and active logging opy frugivores, including hornbills, pigeons, of that forest certainly influenced the size and and barbers.The only exceptionswere smaller constitution of the plantation's bird commu- species (e.g. Megalaima australis) or species nity. In morning surveys, we regularly ob- thought to consumea relatively large propor- servedflocks of birds flying into the plantation tion of animal prey (e.g.Anorrhinus galeritus and 536 MrrRAAND SHELDON [Auk,Vol. 110

M. chrysopogon;Lambert 1992).Even thesewere Minister's, Sabah Chief Minister's, and Sabah Forest not common and were recorded only in the Departments and Patrick Andau for permission to oldest groves. Duff et al. (1984) and Stuebing undertake research in Sabah. Financial support was and Gasis (1989) discovered a similar dearth of provided by the Western Foundation of Zoology (WFVZ). For logistical help at BrumasCamp canopy mammals in the plantation. and Sabah Softwoods, we thank the North Borneo Large terrestrial frugivores also were rare in Timber Company, the SabahFoundation, and partic- the plantation, with the exception of the ubiq- ularly: P. Cassels,R. Ibbotson,M. McMyn, K.-C. Tan, uitousforest pigeon Chalcophapsindica. The only and L.-K. Wong. J. Ewel, G. Davies, F. Lambert, C. phasianidswe recordedwere Arborophilacharl- Marsh, R. Stuebing,and D. Wells advised or helped tonii and the grasslandspecialist Coturnix chi- with variousaspects of the project.M. Bull, G. Falxa, nensis.Phasianids, which are generalist feeders, A. Mack, Taising bin Mattanggal, J. Schmitt, and R. appear to be fairly common in recently logged Sembahelped collect many of the nonsurveyspecies forest (F. Lambert unpubl. manuscript),and we records. A. Johns, F. Lambert, C. Marsh, A. Mostrom, may have missed them in the Albizia because, G. Schnell, D. Wells, and an anonymous reviewer like cuckoos,they are difficult to find unless commentedon the manuscript.This is paper no. 15 of the WFVZ's Sabahbird project. calling. Large-sized and many medium-sized wood- peckers were missing from the plantation, as LrrEP.ATtm• CrrED expectedin a forest composedof thin, young trees. Seven smaller specieswere ANDREWS,N, 1973. Tropical forestry:The timber in- recorded in the older Albizia stands, but few dustry finds a new last stand. Sierra Club Bull. 58:4-9. woodpeckers would be likely to survive exclu- BEEHLER,B. M., K. S. R. KRISHNA RAJU, AND S. ALl. sively in the plantation becausethe trees are 1987. Avian use of man-disturbed forest habitats cropped at about 10 years (i.e. before they de- in the eastern Ghats, India. Ibis 129:197-211. velop nest holes).Albizia trees 20 or more years COCKBURN,P.F. 1976. Trees of Sabah, vol. 1. Sabah in age, however, are attractive to nesting wood- Forest Record No. 10, Borneo Literature Bureau, peckers(D. Wells pers.comm.), a fact that should Kuching, Sarawak. be considered in managing plantation wildlife. DAWES,A. G., AND J. PAYNE. 1982. A faunal survey Flycatchers,with the exception of most mon- of Sabah. IUCN/WWF Project No. 1692, World archs (Rhipidura,Philentoma, Hypothymis, and Wildlife Fund (Malaysia), Kuala Lumpur. Terpsiphone),were notably fewer in the Albizia DUFF, A. B., R. A. HALL, AND C. W. MARSH. 1984. A than in the primary forest, despite the large survey of wildlife in and around a commercial tree plantation in Sabah.Malays. For. 47:197-213. number of mosquitoesand other flying insects. HORN, H. S. 1966. Measurement of "overlap" in Studiesof the effect of logging on primary for- comparativeecological studies. Am. Nat. 100:419- est bird communities in Malaysia and Borneo 494. also have shown that flycatcher diversity is re- JOHNS,A.D. 1983. Ecologicaleffects of selectivelog- duced (Johns 1986, 1989, Wong 1986, Lambert ging in a west Malaysian . Ph.D. thesis, 1992). Why this occurs is not clear, but specu- Univ. Cambridge, Cambridge. lations include a reduction in foraging area for JOHNS,A.D. 1986. Effectsof selectivelogging on the birds (e.g. causedby overcrowding of scrub- the ecological organization of a peninsular Ma- by vegetation), marked change in insect-com- layman rainforest avifauna. Forktail 1:65-79. munity composition, subtle microhabitat dif- JOHNS,A.D. 1989. Recoveryof a peninsular Malay- sian rainforest avifauna following selective tim- ferences affecting both birds and insects (e.g. ber logging: The first twelve years.Forktail 4:89- warmer forest-floortemperatures), and extreme 105. specialization on the part of the birds (Wong KARR,J.R. 1980. Geographicalvariation in the avi- 1986,Lambert 1992).These factors also may have faunas of tropical forest undergrowth. Auk 97: contributed to the absence of other insectivo- 283-298. rous birds in the Albiziagroves, notably the ter- LAMBERT,F.R. 1992. The consequencesof selective restrial speciesPitta baudii,P. guajana,Kenopia logging for Borneanlowland forest birds. Philos. striata,and Napotheraatrigularis. Trans. R. Soc. Lond. B Biol. Sci. 335:443-457. LOVEJOY,T. E. 1974. Bird diversity and abundance ACKNOWLEDGMENTS in Amazon forest communities. Living Bird 13: 127-191. We are particularly indebted to Jody Kennard for MABBERLEY,D.J. 1987. Plant-book. Cambridge Univ. help with the survey.We thank the MalayManPrime Press, New York. July 1993] BorneanPlantation Birds 537

M^CAR?I-IL•R,R. H., AND J. W. M^CAR?I-IL•R. 1961. On WILSON, C. C., AND W. L. WILSON. 1975. The influ- bird speciesdiversity. Ecology42:594-598. ence of selectivelogging on primates and some SHELDON,F. H. 1986. Habitat changespotentially other animals in east Kalimantan. Folia Primatol. affectingbirdlife in SabahEast Malaysia. Ibis 128: 23:245-274. 174-175. WILSON,W. L., ^ND A.D. JOHNS.1982. Diversity and SIBLEY,C. G., AND B. L. MONROE, JR. 1990. Distri- abundance of selected animal speciesin undis- bution and taxonomyof birds of the world. Yale turbedforest, selectively logged forest and plan- Univ. Press, New Haven, Connecticut. tations in east Kalimantan, Indonesia. Biol. Con- SIMPSON,E. H. 1949. Measurementof diversity. Na- serv. 24:205-218. ture 163:688. WoNt, M. 1985. Understory birds as indicatorsof SMYrI-IIES, B. E. 1981. The birds of Borneo, 3rd ed. regenerationin a patchof selectivelylogged west SabahSociety and Malayan Nature Society,Kuala MalaysJanrainforest. International Council for Lumpur. Bird Preservation Tech. Publ. 4:249-263. STUEBINC,R. B.,•NO J.G^sls. 1989. A surveyof small WONG,M. 1986. Trophic organizationof understory mammals within a Sabah tree plantation in Ma- birds in a MalaysJandipterocarp forest. Auk 103: laysia. J. Trop. Ecol. 5:203-214. 100-116. WELLS, D. R. 1985. The forest avifauna of western WOOD,G. H. S., m's•DW. MFaJER.1964. Dipterocarps Malesia and its conservation. International of Sabah (North Borneo). Sabah Forest Record Council for Bird Preservation Tech. Publ. 4:213- No. 5, Sabah Forest Dept., Sandakan,Sabah. 232. WHrrMoRE,T. C. 1984. Tropical rain forests of the Far East. Clarendon, Oxford.

APPENDIX.Species recorded in SabahSoftwoods Albizia groves and adjacentprimary forest. List does not includemigrants or speciesrecorded in otherhabitats. Numbers indicate individuals surveyed in eachplot type; "P" indicatesnonsurvey records. List sequencedaccording to Smythies(1981), and namesfollow Sibley and Monroe (1990).

Hab- Pri- Albiziagroves a itat mary 1975 1977 1979 1981 toler- Name forest (7) (5) (3) (1) ance 1 Bat Kite (Macheirhamphusalcinus) 2 Jerdon'sBaza (Avicedajerdoni) -- 4 -- 1 -- 1.55 3 Brahminy Kite (Haliasturindus) P -- -- 1 -- 1.00 4 CrestedGoshawk (Accipiter trivirgatus) P 4 ------1.00 5 Blyth's Hawk- (Spizaetusalboniger) -- 2 ------1.00 6 Wallace'sHawk-Eagle (S. nanus) P P 1 -- -- 1.00 7 Rufous-belliedEagle (Hieraaetuskienerii) 8 LesserFish-Eagle (Ichthyophagahumilis) 9 CrestedSerpent-Eagle (Spilornis cheela) 1 P -- 1 -- 1.55 10 Blue-breasted Quail (Coturnixchinensis) -- -- 4 1 17 1.41 11 Chestnut-necklacedPartridge (Arborophila charltonii) P 4 ------1.00 12 CrestedPartridge (Rollulusrouloul) 13 CrestedFireback (Lophura ignita) 14 Great Argus (Argusianusargus) 2 .... 1.00 15 Slaty-breastedRail (Gallirallusstriatus) ------1 -- 1.00 16 Red~leggedCrake (Rallinafasciata) -- 1 ------1.00 17 White-breastedWaterhen (Amaurornisphoenicurus) -- p P P P -- 18 Large Green-Pigeon(Treron capellei) 19 Little Green-Pigeon (T. olax) 20 Green Imperial-Pigeon (Duculaaenea) P -- 2 -- -- 1.00 21 Common Emerald-Dove(Chalcophaps indica) P 13 7 1 -- 2.05 22 Blue-rumpedParrot (Psittinuscyanurus) 23 Blue-crownedHanging-Parrot (Loriculus galgulus) 3 12 23 39 -- 2.93 24 MoustachedHawk- (Cuculusvagans) 1 .... 1.00 25 Indian Cuckoo(C. micropterus) 3 .... 1.00 26 BandedBay Cuckoo (Cacomantissonneratii) 1 -- 1 -- 2 2.46 27 (C. merulinus) P 42 35 37 27 3.97 28 Rusty-breastedCuckoo (C. sepulcralis) 1 .... 1.00 29 Little Bronze-Cuckoo (C. minutillus) 30 ( xanthorhynchus) -- 1 5 -- -- 1.38 538 MITRAAND SHELDON [Auk,Vol. 110

APPENDIX. Continued.

Hab- Pri- Albiziagroves a itat mary 1975 1977 1979 1981 toler- Name forest (7) (5) (3) (1) ance

31 DrongoCuckoo (Surniculus lugubris) 7 16 1 -- -- 1.95 32 Raffies'sMalkoha (Phaenicophaeus chlorophaeus) P 11 5 -- -- 1.75 33 Black-belliedMalkoha (P. diardi) 34 Red-billedMalkoha (P.javanicus) I I P -- -- 1.47 35 Chestnut-breastedMalkoha (P. curvirostris) 2 8 5 3 -- 3.65 36 GreaterCoucal (Centropus sinensis) -- 19 22 28 1 2.93 37 LesserCoucal (C. bengalensis) -- -- 3 23 18 2.21 38 CollaredScops-Owl (Otus lempiji) 39 BuffyFish-Owl (Ketupa ketupu) P P .... 40 Brown Hawk-Owl (Ninoxscutulata) 41 Brown Wood-Owl (Strix leptogrammica) P P P -- _ _ 42 MalaysianEared-Nightjar (Eurostopodus temminckii) P P P -- _ _ 43 Large-tailedNightjar (Caprimulgusmacrufus) -- P P 1 -- 1.00 44 Black-nestSwiftlet (Collocaliamaximus) P P P P P -- 45 White-bellied Swiftlet (C. esculenta) P P P P P -- 46 Brown-backedNeedletail (Hirundapusgiganteus) P P P P P -- 47 Silver-rumpedSwift (Rhaphiduraleucopygialis) P P P P P -- 48 WhiskeredTreeswift (Hemiprocnecomata) 11 .... 1.00 49 Grey-rumpedTreeswift (H. longipennis) 50 Diard's (Harpactesdiardii) P 7 2 -- -- 1.53 51 Red-napedTrogon (H. kasumba) I 2 ------1.80 52 Scarlet-rumpedTrogon (H. duvaucelii) 5 4 ------1.38 53 Cinnamon-rumpedTrogon (H. orrhophaeus) 54 Orange-breastedTrogon (H. oreskios) 55 BandedKingfisher (Lacedo pulchella) 56 Rufous-collaredKingfisher ( concretus) 57 Collared (Todirhamphus chloris) ------I -- 1.00 58 Blue-earedKingfisher (Alcedo meninting) 59 Blue-bandedKingfisher (A. euryzona) 60 Oriental Kingfisher (Ceyxerithacus) P 4 I -- -- 1.47 61 Red-beardedBee-eater (Nyctyornis amictus) 62 Bushy-crestedHornbill (Anorrhinusgaleritus) 2 10 4 -- -- 2.69 63 Wreathed (Acerosundulatus) 64 RhinocerosHornbill (Bucerosrhinoceros) 5 .... 1.00 65 Helmeted Hornbill (B. vigil) 10 .... 1.00 66 MalaysianHoneyguide (Indicator archipelagicus) ------I -- 1.00 67 Brown Barbet(Calorhamphus fuliginosus) 8 .... 1.00 68 Gold-whiskeredBarbet (Megalaima chrysopogon) I 2 ------1.80 69 Red-throatedBarbet (M. mystacophanos) I .... 1.00 70 Yellow-crowned Barbet (M. henricii) 71 Blue-eared Barbet (M. australis) 3 2 ------1.32 72 Rufous (Sasiaabnormis) P 8 8 8 I 3.30 73 Crimson-wingedWoodpecker ( puniceus) 4 .... 1.00 74 Checker-throatedWoodpecker (P. mentalis) 75 Banded Woodpecker (P. miniaceus) 76 RufousWoodpecker ( brachyurus) 3 2 ------1.32 77 Grey-cappedWoodpecker (Dendrocopos canicapillus) P 5 ------1.00 78 Buff-rumpedWoodpecker ( tristis) I P I -- -- 1.47 79 Buff-neckedWoodpecker (M. tukki) P I 2 -- -- 1.80 80 Grey-and-buffWoodpecker (Hemicircus concretus) I I ------1.47 81 Olive-backedWoodpecker (Dinopium raffiesii) 82 White-belliedWoodpecker ( javensis) 83 ( pulverulentus) 84 ( rubiginosus) 6 .... 1.00 85 Orange-backedWoodpecker (Reinwardtipicus validus) 4 .... 1.00 86 Green Broadbill(Calyptomena viridis) P 7 ------1.00 87 Black-and-yellowBroadbill ( ochromalus) 9 19 -- I -- 1.90 88 BandedBroadbill (E. javanicus) 2 6 2 -- -- 2.46 July 1993] Bornean Plantation Birds 539

APPENDIX. Continued.

Hab- Pri- Albiziagroves • itat mary 1975 1977 1979 1981 toler- Name forest (7) (5) (3) (1) ance

89 Dusky Broadbill (Corydonsumatranus) 90 Garnet (Pitta g,ranatina) P 11 ------1.00 91 Blue-headed Pitta (P. baudii) 92 BandedPitta (P. guajana) 2 .... 1.00 93 Hooded Pitta (P. sordida) -- 17 7 -- -- 1.70 94 Pacific Swallow (Hirundo tahitica) -- p P P P -- 95 Large Wood-shrike (Tephrodornisvirgatus) P P 1 -- -- 1.00 96 LesserCuckoo-shrike (Coracina fimbriata) 5 11 6 -- -- 2.46 97 Black-winged Flycatcher-shrike(Hemipus hirundinaceus) P 40 21 27 6 3.31 98 Bar-winged Flycatcher-shrike(H. picatus) 1 4 6 -- -- 2.88 99 Fiery (Pericrocotusigneus) 7 -- 4 -- -- 1.28 100 ScarletMinivet (P. fiammeus) 8 3 P 1 -- 1.27 101 Green Iora (Aegithinaviridissima) 4 27 32 9 -- 3.46 102 (A. tiphia) 103 LesserGreen (Chloropsiscyanopogon) 8 3 P -- -- 1.19 104 (C. sonnerati) 5 26 11 3 21 3.85 105 Asian Fairy-Bluebird (Irena puella) 15 21 5 -- -- 1.82 106 Puff-backed (Pycnonotuseutilotus) P P P -- _ __ 107 Black-and-white Bulbul (P. melanoleucos) P -- 2 -- -- 1.00 108 Black-headedBulbul (P. atriceps) P 127 95 30 8 2.74 109 Scaly-breastedBulbul (P. squamatus) 110 Grey-bellied Bulbul (P. cyaniventris) 111 Straw-headed Bulbul (P. zeylanicus) -- 20 ------1.00 112 Yellow-vented Bulbul (P. goiavier) -- 38 155 269 207 3.02 113 Red-eyedBulbul (P. brunneus) -- 48 54 1 -- 2.04 114 Cream-ventedBulbul (P. simplex) -- 5 3 1 1 3.04 115 SpectacledBulbul (P. erythropthalmos) 6 84 86 18 7 3.28 116 Grey-cheekedBulbul (Alophoixusbres) 5 2 ------1.20 117 Yellow-bellied Bulbul (A. phaeocephalus) P .... 1.00 118 Finsch'sBulbul (A. finschii) P 1 .... 119 Hairy-backed Bulbul (Tricholastescriniger) 1 13 2 -- -- 1.91 120 Buff-vented Bulbul ( olivacea) P P .... 121 Rufous-tailedShama (Trichixos pyrropygus) P 1 ------1.00 122 White-rumpedShama (Copsychus malabaricus) 9 43 39 27 10 4.60 123 White-crowned Forktail (Enicurusleschenaulti) P 2 1 -- -- 1.80 124 Chestnut-napedForktail (E. ruficapillus) 125 Chestnut-cappedThrush (Zootherainterpres) P P .... 126 Black-cappedBabbler (Pellorneum capistratum) 5 48 24 8 10 3.78 127 Short-tailed Babbler (Malacocincla malaccense) 2 24 7 4 1 2.87 128 White-chested Babbler (Trichastomarostratum) P 19 3 -- -- 1.31 129 Ferruginous Babbler (T. bicolor) P 29 6 -- -- 1.40 130 Horsfield'sBabbler (Malacocincla sepiarium) P 15 4 2 -- 1.86 131 Rufous-crownedBabbler (Malacopteron magnum) 3 27 7 -- -- 2.30 132 Scaly-crownedBabbler (M. cinereum) 6 5 ------1.40 133 MoustachedBabbler (M. magnirostre) 7 2 ------1.14 134 Sooty-cappedBabbler (M. affine) -- 30 12 11 -- 2.50 135 Chestnut-backed Scimitar-Babbler (Pomatorhinusmontanus) P 1 2 -- -- 1.80 136 Bornean Ground-Babbler (Ptilocichlaleucogrammica) 2 1 ------1.25 137 Striped Wren-Babbler (Kenopiastriata) 138 Black-throatedWren-Babbler (Napothera atrigularis) 139 Striped Tit-Babbler (Macronousgularis) -- 149 168 224 59 3.48 140 Fluffy-backedTit-Babbler (M. ptilosus) 4 33 28 6 3 3.57 141 Grey-headed Babbler (Stachyrispoliocephala) P 3 P -- -- 1.00 142 Black-throatedBabbler (S. nigricollis) -- 2 P 1 -- 1.88 143 White-necked Babbler (S. leucotis) 144 Chestnut-rumpedBabbler (S. maculata) 3 6 ------1.80 145 Chestnut-wingedBabbler (S. erythroptera) 40 114 99 52 8 3.82 146 Rufous-frontedBabbler (S. rufifrons) 9 21 39 27 13 4.67 540 MITRA AND SHELDON [Auk, Vol. 110

APPENDIX. Continued.

Hab- Pri- Albiziagroves a itat mary 1975 1977 1979 1981 toler- Name forest (7) (5) (3) (1) ance

147 Brown Fulvetta (Alcippebrunneicauda) 11 30 3 -- -- 2.08 148 White-bellied Yuhina (Yuhina zantholeuca) 149 Yellow-bellied Prinia (Priniafiaviventris) -- 11 87 139 184 2.62 150 Dark-necked (Orthotomusatrogularis) P 3 ------1.00 151 Rufous-tailed Tailorbird (O. sericeus) 3 157 214 235 138 3.94 152 Ashy Tailorbird (O. ruficeps) 6 107 147 107 20 3.73 153 Spotted Fantail (Rhipiduraperlata) 4 .... 1.00 154 Pied Fantail (R. javanica) -- -- P -- 5 1.00 155 Grey-headedCanary-flycatcher (Culicicapa ceylonensis) 13 .... 1.00 156 Verditer Flycatcher(Eumyias thalassina) P ------2 1.00 157 White-tailed Flycatcher (Cyornisconcretus) 158 Pale-blue Flycatcher (C. unicolor) 159 Sunda Blue-Flycatcher(C. caerulatus) 1 .... 1.00 160 Bornean Blue-Flycatcher(C. superbus) 161 Rufous-chestedFlycatcher (Ficedula dumetoria) 162 Grey-chestedJungle-Flycatcher (Rhinomyias umbratills) 4 .... 1.00 163 Rufous-winged Philentoma (Philentomapyrhopterum) P P 4 -- -- 1.00 164 Maroon-breasted Philentoma (P. velatum) P 2 ------1.00 165 Black-napedMonarch (Hypothymisazurea) 1 51 45 17 2 3.01 166 Asian Paradise-Flycatcher(Terpsiphone paradisi) 7 27 4 -- -- 2.28 167 Velvet-fronted (Sittafrontalis) 3 8 2 1 5 3.49 168 Scarlet-breastedFlowerpecker (Prionochilusthoracicus) 169 Yellow-rumped (P. xanthopygius) P 23 3 1 1 1.52 170 Yellow-breasted Flowerpecker (P. maculatus) P 3 ------1.00 171 Yellow-vented Flowerpecker (Dicaeumchrysorrheum) -- p -- _ p -- 172 Thick-billed Flowerpecker (D. agile) -- 41 ------1.00 173 (D. concolor) -- I ------1.00 174 Orange-bellied Flowerpecker (D. trigonostigma) 1 49 27 16 1 2.88 175 Plain (Anthreptessimplex) 1 93 12 8 6 1.81 176 Brown-throated Sunbird (A. malacensis) -- 14 6 5 -- 2.52 177 Red-throated Sunbird (A. rhodolaema) -- 4 -- -- 4 1.92 178 Ruby-cheekedSunbird (A. singalensis) P 25 32 5 -- 2.36 179 Purple-napedSunbird (Hypogrammahypogrammicum) P 16 8 -- I 2.02 180 Purple-throatedSunbird (Nectariniasperata) -- 5 2 2 -- 2.54 181 Crimson Sunbird (Aethopygasiparaja) -- -- 3 3 2 2.98 182 Scarlet Sunbird (A. mystacalis) I P I -- -- 1.47 183 Little (Arachnotheralongirostra) 9 122 89 70 1 3.55 184 Thick-billed Spiderhunter (A. crassirostris) -- 8 3 5 -- 2.65 185 SpectacledSpiderhunter (A. fiavigaster) P P -- 1 -- 1.00 186 Long-billed Spiderhunter (A. robusta) -- P -- -- 1 1.00 187 Yellow-eared Spiderhunter (A. chrysogenys) P -- 1 1 2 2.36 188 Grey-breastedSpiderhunter (A. affinis) 1 7 ------1.86 189 Everett's White-eye (Zosteropseveretti) -- 21 -- 30 -- 1.87 190 Hill Myna (Graculareligiosa) 2 21 8 45 -- 2.38 191 BorneanBristlehead (Pityriasis gymnocephala) 192 Dusky Munia (Lonchurafuscans) -- 8 19 23 22 3.37 193 Chestnut Munia (L. malacca) .... 2 1.00 194 Bronzed (Dicrurusaeneus) P P .... 195 Greater Racket-tailedDrongo (D. paradiseus) 4 12 ------1.96 196 Black-hooded Oriole (Oriolus xanthornus) ------1 -- 1.00 197 Dark-throated Oriole (O. xanthonotus) 10 20 9 -- -- 2.29 198 Crested Jay (Platylophusgalericulatus) 199 Black Jay (Platysmurusleucopterus) 200 Slender-billed Crow (Corvusenca) P 17 P 7 I 2.00 Total species 162 122 92 63 45 -- Total individuals 338 2,200 1,792 1,591 820 --

Plot types (age in years).