dan reptilia meliputi: 13 jenis katak, 29 jenis bengkarung, 29 jenis ular, 1 jenis
kura-kura air tawar dan 1 jenis buaya. Ada jenis-jenis endemik 38 percent ...
Herpetofaunal Richness and Community Structure of Offshore Islands of Sulawesi, Indonesia Author(s): Graeme Gillespie, Sam Howard, David Lockie, Michael Scroggie, Boeadi Source: Biotropica, Vol. 37, No. 2 (Jun., 2005), pp. 279-290 Published by: The Association for Tropical Biology and Conservation Stable URL: http://www.jstor.org/stable/30043182 . Accessed: 19/05/2011 08:53 Your use of the JSTOR archive indicates your acceptance of JSTOR's Terms and Conditions of Use, available at . http://www.jstor.org/page/info/about/policies/terms.jsp. JSTOR's Terms and Conditions of Use provides, in part, that unless you have obtained prior permission, you may not download an entire issue of a journal or multiple copies of articles, and you may use content in the JSTOR archive only for your personal, non-commercial use. Please contact the publisher regarding any further use of this work. Publisher contact information may be obtained at . http://www.jstor.org/action/showPublisher?publisherCode=tropbio. . Each copy of any part of a JSTOR transmission must contain the same copyright notice that appears on the screen or printed page of such transmission. JSTOR is a not-for-profit service that helps scholars, researchers, and students discover, use, and build upon a wide range of content in a trusted digital archive. We use information technology and tools to increase productivity and facilitate new forms of scholarship. For more information about JSTOR, please contact
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BIOTROPICA 37(2): 279-290 2005
10.1111/j.1744-7429.2005.00038.x
HerpetofaunalRichness and CommunityStructureof OffshoreIslands of Sulawesi, Indonesial Graeme Gillespie2,6,Sam Howard3,David Lockie4, MichaelScroggie2, and Boeadi5 2ArthurRylahInstitutefor Environmental Research, PO Box 137, Heidelberg3084, Victoria,Australia 357 WedderburnRoad, Harrogate,NorthYorkshire,HG27QQ, UnitedKingdom 4LarkRise, FranksField,Peaslake, Guildford,Surrey,GU5 9SS, UnitedKingdom 5MuzeumZoologicumBogorinse,Sibinung,West Java, Indonesia
ABSTRACT We examinedthe distributionand broadhabitatassociationsof the herpetofaunaon threeoffshoreislandsof southeastSulawesi,Indonesia.A total of 74 amphibianand reptile taxawererecorded,comprising13 frogs,29 lizards,29 snakes,1 freshwaterturtle,and 1 crocodile.Of the totaltaxa,38 percentwereendemicto Sulawesi,13 werenew undescribed taxa. Rangeextensionswere also recordedfor one taxon previouslynot known from Sulawesi.Herpetofaunaof these islandsis largelyderivedfrom that of mainlandSulawesi, and as for Sulawesigenerally,is depauperatecomparedwith herpetofaunalassemblagesin Borneo,Java,and Thailand.Taxonrichnesswas much higherin minimallydisturbed forestand foresthabitatswith only moderatedisturbancelevels than in highly modifiedor disturbedhabitats,such as secondaryforests,plantations,and villages.Disturbed by endemicand habitat-specialist taxa.Littlediscriminationof by widespread,habitatgeneralistsand humancommensals.Forestswerecharacterized habitatswerecharacterized taxon compositionor endemismwas found betweenminimallyand moderatelydisturbedforesthabitats.These resultsreaffirmthe need for moregeneralbiologicalsurveyand researchin this region.Taxamost likelyto be displacedby human impactstend to be endemictaxa,for which thereexistslittle or no ecologicalinformation.The similarityin the herpetofaunalcommunitystructurebetweenhabitatswith minimaland moderatedisturbancelevelshasimportantimplicationsfor our understandingof ecologicalresilience in tropicalherpetofaunalcommunities.
RESUMEN Herpetofaunadi kepulauanlepaspantaiSulawesiTenggara:Kekayaanjenis,asosiasinyadenganhabitatsertahubunganantarastrukturkomunitasnyadangangguanulahmanusia. Telahditelitiluassebarandan asosiasinyadenganhabitatherpetofaunadi tiga buahkepulauanlepaspantaidi SulawesiTenggara,Indonesia.Tercatatsejumlah74 taksaamfibia dan reptiliameliputi:13 jenis katak,29 jenis bengkarung,29 jenis ular,1 jenis kura-kuraair tawardan 1 jenis buaya.Adajenis-jenisendemik38 percentsedangkanl3taksadi antaranyamerupakanjenis baruyang belum dipertelakan.Ada satu jenis rekorbaruyang sebaransebelumnyatak di kenaldi Sulawesi.Herpetofaunadi kepulauanini berasal daridaratanSulawesisedangkanjenis-jenisamfibia- reptiliadi Sulawesisendirisangatsedikitjumlahnyabila dibandingkandengangabunganjumlahyang adadi Kalimantan, Jawadan Siam. Di hutan yang mengalamisedikitgangguandan di habitatdengantingkatperubahansedangterdapatlebih kayakayajenis daripada di kawasanyang sudah tergangguatauhabitatyang sudahbanyakberubahsebagaihutan sekunder,kebundan pedesaan.Di habitatyang telah terganggumakaditemukanjenis-jenisyang umum dan biasaterdapatdi hunianmanusia.Di hutanyang masihutuh ditemukanjenis-jenisendemikdan jenis yang khasmenuruthabitatnya.Di antarahabitathutan yang menglami sedikitgangguandanyangmendapatgangguansedangtidakterdapatperbedaannyatakompsisijenismaupunendemismejenis.Kesimpulanini mengukuhkanperlunyaagarlebih menaruhperhatiandilakukansurvaidan penelitianalamhayatidi kawasanini. Karenaulahmanusiacenderungberdampakpadajenis-jenisendemikdi kawasanini tersingkirkan tanpaatausedikitsekaliketeranganmengenailingkunganhidupnya.Adanyakesamaanstrukturkomumitasherpetofaunayang adadi antarahabitatyang mengalamigangguan minimaldan gangguansedangbermaknapentingsekalidalampengertianguna upayapemulihanlingkunganbagikomunitasherpetofaunadi kawasantropis. tropical Southeast Asia;Sulawesi;taxonrichness; reptiles; Keywords:amphibians;conservation; forest.
SULAWESIIS A LARGE EQUATORIALISLAND IN THE INDONESIAN
ARCHIPELAGO lying between Borneo and the Maluku (Mollucas)islands. geological complex origins and geographicalposition have resultedin Its a distinctivemixtureof Australasianand Asian floraand faunaelements, with high levels of endemism acrossall vertebrategroups (Whitten et al. 2002, Evans et al. 2003). However, despite the biogeographicalsignificance of Sulawesi,little is known about the biology of most organisms occurring in the region. More than 115 reptile and 40 frog taxa have been recordedfrom the island to date, many of which are endemic (Iskandar& Tjan 1996). However, relativelyfew surveys have been undertakenof the herpetofauna of Sulawesi and associated smaller islands, and these figures are 1Received26 August2003; revisionaccepted3 September2004. 6 Correspondingauthor. Currentaddress:Zoos Victoria, P.O. Box 74, Parkville, Victoria3052, Australia;e-mail:
[email protected] 279
likely to be conservativeestimatesof the taxon richness.Approximately half of the frog taxa known from Sulawesihave been discoveredin the past 15 yr, and many are yet to be formallydescribed.New reptile taxa have also been discoveredin the recent years (Iskandar& Tjan 1996). Systematicsof several other reptile groups requiresfurther resolution, and the status of a number of taxa in the region requiresclarification (Iskandar& Tjan 1996, Evanset al. 2003, Gillespie et al. 2004). Conservationstatus of many poorly known taxa is also unclear. Knowledge of biodiversityand organizationof communities is essential to develop conservation policy and sustainable environmental management systems. Such knowledge provides the basis for identifying important areasto focus limited conservationresources,and identifying threats that require mitigation. This may only be achieved if a sound knowledge exists of systematics,taxon distributions,and habitat associations. This information is lacking for herpetofaunathroughout much of Southeast Asia. As for other areas in Southeast Asia, habitat
280 Gillespie,Howard,Lockie,Scroggie, and Boeadi
destruction threatens the biodiversity of Sulawesi (Bawa et al. 1990). Habitat destruction and fragmentation are the leading causes of amphibian and reptile decline around the world (Blaustein et al. 1994, Sarre et al. 1995). It is therefore imperative to adequately document the biodiversity of this region and to gain a better understandingof community organizationand the impact of disturbanceprocesses. Extensiveareasof forestpersiston some of the largeroffshoreislands in southeast Sulawesi.These forestedareashave the potential to provide a basis for a robust conservation reserve system in the region. However, knowledge of the flora and fauna of these islands is poor. Whereas Iskandarand Tjan (1996) presented a limited broad-scaleherpetofaunal inventory for Sulawesi, specific information on the major islands of Sulawesi Tenggarais limited. Biodiversity surveys of the islands of SulawesiTenggaracommenced in 2000 and this paper presents the results of the herpetofaunalcomponent of this program. Our objectives were to (1) document taxaoccurringin the region, (2) assessthe relative effectivenessof various survey techniques, and (3) examine patterns of community composition, habitatassociations,and the impact of human disturbanceon community composition.
METHODS STUDY AREA.-SulawesiTenggaracomprisesthe southeasternarm of Sulawesi, along with a cluster of smaller islands, including the Tukang Besi/Wakatobi Archipelago (Fig. 1). The main islands are Kabaena (approx.1000 km2), Muna (2900 km2), and Buton (approx.4400 km2). All these islands are separatedfrom the mainland by shallow seas and
would have been connected to Sulawesi during the Pleistocene (Voris 2000). Buton is the largestoffshore Sulawesiisland and the closest to the mainland. It has a hilly terrain,rising to 900 m in elevation, dominated by intensively karstified Cenozoic coral limestones (J. Milsom, pers. comm.). Buton has so far retained a large proportion of forest cover, some of which is contained within severalforest reserves.Vegetationon Buton is tropical monsoonal rainforest.Limited rainfall data available for Buton indicate annual precipitationat sea level of 1500-3000 mm (Departmentof Agriculture,Bau Bau, SulawesiTenggara,pers. comm.). Monthly rainfallvaries considerablyannually but there is a monsoon season from November to January and a pronounced dry season in August to October. Kabaenahas a mountainous terrain,rising to 1650 m in elevation, dominated by a major oceanic overthrust (ophiolite, J. Milsom, pers. comm.). Much of the lowlands of Kabaenahave been deforested, but significant areas of forest persist in the mountainous center and lessaccessiblevalleys.Vegetationon Kabaenais characterizedby more open forests and savanawoodlands with grassy understories,and grasslands occur on exposed ridges. Closed canopy rainforestis more restrictedto montane areasand lower valleys. Muna Island has been largely deforested, as a result of its flatter terrainof less than 100 m elevation, and is now a mosiac of plantations, crops, and secondaryscrub.
SURVEYSITES.-The
survey focused on six areas, five of which were
established on Buton and Kabaena (Fig. 1). The sixth sampling area
Southeast Sulawesi
0°
Buton 4
5 6 (Hoga)
Kabaena Muna 50 S
2
100 km 1200 E
1250 E
FIGURE 1. Mapsof Sulawesiand SulawesiTenggarashowingthe studyareasamplingsites.
Herpetofaunaof OffshoreIslandsof Sulawesi 281
was on HogaIsland,a small,relativelyremote,coralkarsteatollin the Archipelago.Collectively,these areasprovided TukangBesi/Wakatobi somegeographic coverageandenabledsamplingof bothlargeandsmall islandsas well as coastal,lowland,and mountainousareas,anda range of modifiedandintacthabitats.
Timed diurnal searches of 1-h duration were also undertaken at point localities in all habitat classes, including each pitfall trap site. These searcheswere undertaken by two people slowly walking across an areaof broadlyconsistent habitat type, recordingactive and basking taxa, and searchingunder rocks, logs, and within litter and debris for shelteringor cryptic taxa. Limited road networksexist on Buton and KabaenaIslands.Roads were traversedat night in a slow-moving vehicle to detect any taxa on or crossingroads.Incidentalobservationsof herpetofaunawere collected over a wider areaof the region. These included field observationsmade while accessingsampling sites, reconnaissancewalks, road-killedspecimens and, although not encouraged,specimens presentedto us by local people. It was not possible to standardizesampling effort among sampling techniques,but an effortwas made to ensurethat a reasonablelevel of effort was invested in each sampling approachthroughout the survey in each study area,and within each broadhabitat type. Voucher specimens of most taxa were collected to aid the identification of unknown taxa or suspected new taxa, and to collect material for taxonomic groups requiringfurthersystematicresolution. All specimens collectedhave been lodged at the Muzeum Zoologicum Bogorinse, Indonesia (Appendix).
FIELD SAMPLING.-Field surveyswereconductedfromJuneto September in eachyearfrom2000 to 2003. Thissamplingperiodroughlycoincided with the transitionfromthe end of the wet seasonto the dry season. Samplingeffortvariedbetweenareasbecauseof access,seasonalfactors, andtimeconstraints. Priorto thisstudy,the effortrequiredto complete comprehensive inventoriesof herpetofaunal communitiesin the region was unknown.To assessthis, and as partof moredetailedstudies,we samplingone locality(Area1) in assignedpriorityto comprehensively a muchhighersamplingeffortwasundertaken at all4 yr.Consequently, Area1 thanthe otherareas. possible,samplesites To samplethe widestrangeof environments werechosenin a varietyof differenthabitats,suchas forestwithvarying and at sitesin highlydisturbedhabitats, levelsof humandisturbance, clearedforestandplantations. Otherhabitats,such suchas regenerating as streams,swamps,coastaland estuarineareas,and caves,were also specifically investigated. Standardmethodswere employedto samplethe herpetofauna ANALYSES.-Eightbroad habitat types were identified: minimally diswithineacharea,includingpitfalltrapping,diurnalandnocturnalcenturbed forest, moderatelydisturbedforest, secondaryforest, plantations suses,andopportunistic searches.Initially,intensivesearches,employing and farmland, villages and towns, coastal and estuarine habitats, and litter removaland destructivesearchesof logs and crevices,were atcaves. Minimally disturbed forest was defined as forest areas remote temptedin 5 x 5 m quadrats at severalsites(afterLloydetal. 1968),but (>4 km) from human settlement or roads, and although some disturtheseprovedunproductive and labor-intensive, andwerediscontinued bance may have occurred historically,little or no human disturbance in favorof othertechniques. was evident during the study. Moderately disturbed forest comprised Pitfalltrapscomprisedfiveplasticbuckets,embeddedin theground forestedareaswith evidence of disturbancefrom logging of selected trees 40 cmhighand25 m long, 4-5 m apart,witha driftfence,approximately and rattan (climbing palm) harvesting.This habitat was distinguished passingacrossthe middleof eachbucket.Driftfenceswereeitherplastic from minimallydisturbedforest by the presenceof scatteredtreestumps sheetingor 2-mm-gaugewiremesh,with the bottomedge embedded and rattan cutting trails or foot tracks. Secondaryforest was defined as in the ground.Bucketswereeither40 or 60 literin volumeand 57 or regeneratingforest historicallyclear-felledeither for timber or farming. 60-cmdeep,respectively; dueto limitedlocalsuppliesnot enoughof one This habitat was distinguished from the former two by the presence sizewasavailable. in trapsuccess,eachpit site To controlfordifferences of young trees with uniform height, absence of old trees, and simple hadthesameratioof bucketsizes,andthepositionof bucketsizeswithin structure.Plantationsand farmlandincluded all areasunder cultivation the traparraywas randomlyassignedto eachsite. Trapswerechecked outside of villages, including coconut, cocoa and cashew nut plantadailyin the morning. tions, rice paddies, and various other food crops. Coastal and estuarine Nocturnalcensusesof 20-min durationwereundertakenat point habitatscomprisedall coastalforeshorehabitats, mangroves,and bracklocalitiesin all habitatclassesexceptstreams,includingeachpitfalltrap ish stream estuaries,with both relativelyhigh and low levels of visible human disturbance. site. This censusdurationwas adequateto searchhabitatswithin apwas proximately a 20 m radiusof pitfalltrapsites,andthismethodology The presence/absenceof each taxon was recorded in each of the eight (G. habitat types, and a matrix of dissimilaritiesin habitat occupancy as a part of other Gillespie, concurrent studies required sampling searches, among taxa was computed using the Bray-Curtisindex. Agglomerative pers.comm.).Timed searches,ratherthan area-constrained hierarchichalclustering(average-linkagemethod) was used to group taxa wereundertakenbecauseof the difficultyin achievingconsistentsamon the basisof similarityin habitatutilization,with the resultsdisplayed pling effortby areabetweenhabitatswith largedifferencesin strucas a clusterdendrogram. turalcomplexity,suchas forestsandplantations.Censuseswereundertakenby two peoplewearingheadlamps,slowlywalkingacrossan area following literaturewas consulted for taxonomy with broadlysimilarhabitattype,searchingthe ground,undergrowth, NOMENCLATURE.-The and overstoryfor animals.Nocturnalcensusesof streamswereunderand nomenclature:Boulenger (1897), Rooij (1915), Kampen (1923), sampling,as Greer (1977), Bosch (1985), Iskandarand Tjan (1996), Manthey and takenalong200 m transectsratherthantime-constrained Grdssmann (1997), Iskandarand Colijn (2000), Honda et al. (2001), a greaterlevelof consistencywasachievableand this methodologywas asa partof otherconcurrent required samplingstudies(seeGillespieetal. Iskandarand Colijn (2001), Das and Greer (2002), and Mausfeld and 2004). Bohme (2002). Because of the poor resolution and confusion of the
282 Gillespie,Howard,Lockie,Scroggie, and Boeadi
systematics of several groups in Sulawesi, we also consulted relevant expertson appropriatenomenclature (R. Brown, I. Das, D. Iskandar,J. Mc Guire, & G. Shea, pers. comm.).
RESULTS A total of 74 amphibianand reptiletaxawere recordedduring the study (Table 1) comprising 13 anurans, 29 lizards, 30 snakes, 1 freshwater turtle, and 1 crocodile. Twenty-eight (38%) of these taxa are considered endemic to Sulawesi. Severalnew and undescribedtaxa were also discovered, including at least two frogs, eight lizards,and three snakes. typus, Range extensionswere recordedfor one taxon, Hemiphyllodactylus previouslyknown from the SundaShelf west of Sulawesi,but not known from Sulawesi. Range extensions were also recorded for a further two frog, six lizard,and six snake taxa previouslyknown from Sulawesi,but not documentedfrom the SulawesiTenggara. Markeddifferenceswere found in taxon richnessamongst the three islandssampled(Fig. 2); however,samplingeffortalso variedbetween the islands (Table2). The most taxa (67) were found on Buton, followed by Kabaenawith 36, then Hoga with 12. The number of endemic taxawas proportionallyhigher on Buton (43%) comparedwith Kabaena(29%) and Hoga (17%) (Fig. 2). Two taxa, one undescribed,were found only on Kabaena,similarlywith Hoga. Taxaaccumulationcurvesfor Area 1 show that after2 yr most taxa likely to occur in the areawere detected (Fig. 3). However,a significant number of additional taxa (6.9 %) were also added to the inventory for Area 1 in the third year of sampling. No additional taxa were detected in the fourth year. The most productive sampling techniques for inventory were pit trapping,nocturnalcensuses,and incidental observations(Fig. 4). Diurnal censuses and road searcheswere comparativelyunproductive.They yielded fewer taxa than other approaches across all major taxonomic groups, and only one lizard taxa, Emoia atrocostata,was detected more frequentlyby diurnalcensus than other approaches(Fig. 4b). Nocturnal censuses and pit traps were the most productive approachesfor frogs (Fig. 4a). All taxa were detected with these techniques combined, and only one taxon, Fejervaryacancrivora,was detected more frequentlyby incidental observation.The patternsfor lizardsand snakeswere similar, with most taxa detected by incidental observationfollowed by pit traps (Figs. 4a and 4b). Pit traps, nocturnal censuses, and incidental observations all yielded similar number of taxa not detected by any other approaches. The greatesttaxon richnesswas found in minimallyand moderately disturbed forests (48 and 52 taxa, respectively;Fig. 5). These habitats also contained relativelyhigh proportionsof endemic taxa (56 and 54%, respectively).Fourteen endemic taxa were restrictedto these habitats, three of which were only found in minimally disturbed forest. Several nonendemic arborealtaxawere also restrictedto these habitats(Table 1). Although lower in taxon richness,riparianhabitatsalso had a high proportion of endemic taxa, two of which were restrictedto this habitat type. Highly modified and disturbed habitats (i.e., secondary forest, farmland/plantation,and village/towns) had much lower taxon richness with low levels of endemicity. The endemic taxa in these habitatswere all found in a wide range of alternativehabitat types. These modified
habitats,along with estuarineand coastalhabitats,were characterizedby widespreadSoutheastAsian and human commensal taxa, such as Fejervaryacancrivora,Hemidactylusfrenatus, and Eutropismultifasciata.Caves had the lowest taxon richness with only three taxa detected, although two were endemics, one of which (Cyrtodactylusnew sp. 1) was only found in caves. The resultsof the hierarchicalclusteringprocedurearepresentedas a cluster dendrogramwith the taxa names superimposed(Fig. 6). Taxa exhibited a strong clusteringbased upon broad habitatassociations.The largestdissimilarityoccurredbetween the one cave-restrictedtaxon and the rest, followed by taxa restricted to coastal and estuarine habitats, porosusand the taxa restrictedto Hoga island; Cyrsuch as Crocodylus cf monticolasp. 2 todactylusnew sp. 2. Enhydrisspp. and Rhacophorous were the third group separated,which were the only taxa restrictedto streams.Other stream-dwellingtaxa,such as the freshwaterturtle Curora amboinensisand stream-breedingfrogsLimnonectes cf grunniensand Rhacophoruscf. monticolasp. 1, were also observedaway from streams. The fourth major groupings distinguished taxa restrictedto minimally or moderatelydisturbed forest habitats from those occurring in the more disturbedhabitats.Apart from the few taxa restrictedto either minimallydisturbedforest(Calamarianuchalis,Calamarianew sp. 2 and new sp. 2) or moderatelydisturbed forest (Bronchocoela Sphenomorphus cristellataand Ptychozoonkuhli), thereis no discriminationbetween these two habitats.The remainderof this group is furtherdivided on the basis of occurrence in riparianhabitat within forest. The disturbed habitat grouping comprises a large group of habitat generaliststhat occurred across a range of disturbed and forest habitats, and a smaller group of human commensal taxa restrictedmostly to villages. With few exceptions, endemic taxa were predominantlyassociated with minimally or moderately disturbed forest habitats. The one cave specialistwas also found in a minimallydisturbedhabitat.Most endemic taxa occurring in riparian habitats, including those restrictedto this habitat, were only found in streams within minimally or moderately disturbedforest.
DISCUSSION ISLAND TAXONCOMPOSITION.-The herpetofaunaof the study islands is largely derived from that of mainland Sulawesi (see Iskandar & Tjan 1996). Excluding new, undescribed taxa and others previously not recorded from Sulawesi, the taxa recorded in this study represent 42 percent of the taxa previouslyknown from Sulawesi (Iskandar & Tjan 1996). Seventy-five percent of taxa previously known from Sulawesi Tenggarawere also recorded (Iskandar& Tjan 1996). In addition, 26 taxa, including 13 undescribedtaxa that have not previously been recordedin SulawesiTenggara,were found on these islands.In view of the limited intensive sampling in the region, it is likely that many of these taxa are more widespreadthan currentlydocumented. The findings of this limited geographicstudy suggest that the herpetofaunaof Sulawesiand associatedislandsis considerablyundersampled.Terrestrial mammal surveys conducted concurrentlywith this study also detected severalundescribedtaxa (N. Grimwood & G. Gillespie, pers. comm.). Greatersurvey effort is thereforenecessaryto document more fully the terrestrialvertebratebiodiversityof this region.
Herpetofaunaof OffshoreIslandsof Sulawesi 283
TABLE1.
Taxalist showingendemicity,habits,and occurrences acrossthe islandssurveyedand the habitatssampled.Habitsindicatethosetypicalof eachtaxon: T-Terrestrial; AQ-aquatic; AR-arboreal;F-fossorial. Habitat Minimally Moderately Estuarine/ Secondary Plantation/ Village/ disturbed disturbed forest Habit Endemic Kabaena Buton Hoga forest town coastal-foreshoreCave Riparian forest farm Island
Amphibia Microhylidae Kaloulabaleata Oreophryne new sp. Ranidae Fejevarya cancrivora cf. grunniens Limnonectes Limnonectes cf. modestus Ranacelebensis Ranachalconota Ranaeurythraea Rhacophoridae leucomystax Polypedates Rhacophorus georgii Rhacophorus cf. monticolasp. 1 cf. monticolasp. 2 Rhacophorus
T AQ/T T AR/T AR/T T
+ + +
+ + +
AR
+ + +
+ +
+
+
+ +
AR
+
T
+
Dibamidae Dibamusnovaeguineae
F
+
Agamidae cristellata Bronchocoela Draconew sp.
AR AR
+
+
+ +
AR AR/T
+
+
+ +
+
+
+
+
+ +
+ +
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+ +
+
+
+
+
+
+
+
+
+ + + +
+ +
+ + +
+
AR AR
Bufonidae Bufocelebensis
+ +
+ +
T T
+
+ +
Reptilia
+
Geckonidae platyurus Cosymbotus Cyrtodactylusjellesmae Cyrtodactylus new sp. 1 new sp. 2 Cyrtodactylus Gekkogecko Gehyramutilata Gehyranew sp. Hemidactylusfrenatus Hemiphylodactylus typus Ptychozoon kuhlii
Cave AR/T AR AR AR AR AR AR
Varanidae Varanus salvator Scincidae Emoiaatrocostata Eutropismultifasciata Eutropisrudis Eutropisnew sp.
AR/T T T
AR/T AR Lamprolepis smaragdinum cf. smaragdinum AR Lamprolepis Lipinacf. quadrivittata AR/T T bowringii Lygosoma
+ + +
+ + + + + +
+
+
+ + +
+ + + + +
+
+ +
+ + +
+ +
+
+ + +
+
+ +
+
+
+ +
+ +
+ +
+
+
+
+
+
+ +
+ + +
+ + + +
+ + + +
+
+ +
+
+
284 Gillespie,Howard,Lockie,Scroggie,and Boeadi
TABLE1. Continued. Habitat Minimally Moderately
Island
disturbed
disturbed
forest
forest
+ + + + +
+ + +
+
+
+ +
+ +
+
+ + + + +
+ +
+
+ +
+ +
+ + +
+ +
+ + + + +
+
+
+
+
+
Habit Endemic Kabaena Buton Hoga quadrupes Lygosoma cf. temminki Sphenomorphus tropidonotus Sphenomorphus variagatum Sphenomorphus new sp. 1 Sphenomorphus new sp. 2 Sphenomorphus Sphenomorphus new sp. 3 Acrochordidae granulatus Acrochordus
Village/
farm
town
+
+
+
+ +
+ +
forest
Estuarine/
coastal-foreshoreCave
T/F T/F AR/T T T T
+ + +
+
+
+
+
T/F AQ
AR Amphiesmacelebica T Boigadendrophila AR AR/T Boigairregularis Calamariabrongersmai T Calamarianuchalis T T Calamarianew sp. 1 Calamarianew sp 2 T/F rynchops Cerberus AQ Chrysopelea paradisicelebensis AR T/F melanotus Cylindrophis Dendrelaphis AR pictuspictus Elapheeuruthreacelebensis T T Elaphejansenii Enhydrismatannensis AQ Enhydris plumbea AQ T aulicus Lycodon T Oligodonwaandersi pulverulentusAR/T Psammodynastes chrysargos Rhabdophis A/T Rhabdophis chrysargoides A/T Xenochrophis trianguligerus A/T
+
+
+
+ +
+
Colubridae Ahaetullaprasina
Xenopeltidae Xenopeltisunicolor
Secondary Plantation/
Riparian
+ + + + + +
+ + + + + +
+
+
+
+ +
+
+
+
+
+
+
+
+
+ +
+
+
+
+
+ + +
+ + +
+
+
+
+
+ +
+
T/F
+ +
+ + +
+
Crotalidae wagleri Tropidolaemus Elapidae Ophiophagus hannah
AR
+
+
+
T
+
+
+
+
+
+
+
+
+
+
Pythonidae Pythonreticulatus Typhlopidae Cyclotyphlops deharvengi braminus Ramphotyphlops new sp. Typhlops Bataguridae Curoraamboinensis
AR/T
+
+
F AR/F
+
AR/F
AQ
+
+
+
+
+ + +
+
+
+
+
+
+
+
+
+
+
Crocodylidae Crocodylus porosus
AQ
+
+
+
Herpetofaunaof OffshoreIslandsof Sulawesi 285
70
a. Frogs
12
60
10
50 40 30 20 10
Pit trap
Hoga
Kabaena
Buton
FIGURE2. Numberoftaxafoundon eachisland.Solidportionsof the barsrepresent taxaendemicto Sulawesi;open portionsof barsarenon-endemictaxa.
Diurnal Nocturnal search search
Road search
Incidental
b. Lizards
25
20 TABLE2. Samplingeffortin eachsurveyarea.
Area
15
No. of
No. of
No. of
pitfalltrap-nights
day/nightcensuses
persondaysin area
4592
370
336
0
15 58
222 126
10 37 22
25 78
176 0
10
50 20
25
Pit trap
Nocturnal search
Road search
Incidental
c. Snakes
25
70
Diurnal search
Frogs 60
Lizards 50 40
Snakes
15
Total herpetofauna
10
30 20 10
Pittrap 2000
2001
2002
Diurnal Nocturnal Road search search search
Incidental
2003
FIGURE3. Taxonaccumulationcurvesfor variousherpetofaunalgroupsdetected at Site 1, CentralButon Island.The total also includesthe freshwaterturtle Curora
FIGURE4. Numbersof taxarecordedby varioussamplingmethods.Solidportions of barsaretaxadetectedexclusivelyby thespecifiedsamplingmethod;hatchedportions aretaxadetectedmost frequentlyby the method;open barsareall othertaxadetected
whichwas firstdetectedin 2000. amboinensis,
by the method.
286
Gillespie, Howard, Lockie, Scroggie, and Boeadi
60
50
40
30
20
10
FIGURE5. Numberof taxafoundin eachhabitat.Open portionsarenon-endemic taxa;solid portionsof barsareendemictaxa;stippledportionsareendemictaxaonly found in one habitat;hatched portions are non-endemictaxa only found in one habitat.
Variationin taxon richnessamong the three islands examined conforms to island biogeographictheory (MacArthur& Wilson 1967) in that taxon richnesswas highest on the largestisland closest to the mainland (Buton), and lowest on the smallestand most remoteisland (Hoga). However,the size of islandsis also important in this theory,and as Hoga is the most distant but also the smallest island examined, the effects of distanceand island size cannot be distinguishedon the basis of available data.The taxon assemblagesof Kabaenaand Hoga areessentiallysubsets of that of Buton. They also comprise mostly "generalist"taxa that occur in a wide range of modified habitats, with wide distributions throughout Indonesia (Iskandar& Tjan 1996; Manthey & Gr6ssmann 1998; Iskandar& Colijn 2000, 2001; Thornton et al. 2002). With the exception of one undescribedtaxon (Cyrtodactylus new sp. 2), all taxa found on Hoga arehabitatgeneralistsand/or human commensals (Lim & Lim 1992, Manthey & Gr6ssmann1997, Stuebing & Inger 1999), and many readily disperse in association with humans and colonize small islands within the Indonesian archipelago(e.g., Grismeret al. 2002, Thornton et al. 2002). Although not previously documented, the occurrence of Cyrtodactylus sp. 2 on a small island such as Hoga may indicate that it is also a highly dispersivetaxon. Alternatively,this may be a new species endemic to the Tukang Besi Archipelago.This genus may be prone to rapid speciation in isolation, such as on small islands, as demonstrated on Pulau Tioman (Das & Lim 2000, Grismeret al. 2002). Unequal sampling effort undoubtedly contributedto differencesin taxon richnessobservedamong samplingsites;Buton had by farthe highest sampling effort, while Hoga had the lowest. The taxon accumulation curveforArea 1 suggeststhat furthersurveyswould uncoverother taxain
the region, particularlyon Kabaenaand other partsof Buton Island.The resultsfrom Area 1 indicate that relativelyintensive sampling for several months over at least two seasons is requiredto have some confidence of documenting most of the herpetofaunalassemblagein this region. Publishedaccounts of herpetofaunastudies that have attempted to comprehensivelysample the herpetofaunaof an areaor specificregion in SoutheastAsia are few: lowland rainforestin Borneo (Lloyd et al. 1968; Das 1995, 1996) and evergreenand deciduous forests in northeastern Thailand (Inger & Colwell 1977). Our comprehensivedata for Site 1 reveal the taxon richness here to be much lower than areaswith similar elevation in Borneo and Thailand. In Sarawak,Lloyd et al. (1968) recorded48 frog, 40 lizard, and 47 snake taxa. In Brunei Daraussalam, Das (1996) recorded 48 frog, 1 caecilian, 20 lizard, 23 snake, and 3 turtle taxa. In Thailand, Inger and Colwell (1977) recorded 24 frog, 31 lizard, 47 snake, 2 turtle, and 1 ceacelian taxa. Caecilians have not been recordedfrom Sulawesi(Iskandar& Tjan 1996). Other taxonomic groupsnotably depauperateon Buton comparedwith the sites in Borneo and Thailand include all anuranfamilies,agamids,gekkonids,colubrids, viperids,and elapids.With the exception of elapids, the most depauperate groups within these families tend to be characterizedby taxa with arborealhabits. Brown and Alcala (1961) surveyedthe herpetofaunaof submontane and montane rainforestsin the Philippines and found 12 frog, 20 lizard, and 19 snake taxa. However, their sampling effort was substantiallyless than the other studies, and taxon diversityis reduced at higher elevations (Scott 1976), so no comparison is made with our study. Although local environmentalfactorsmay contribute to constraintaxon richness and diversity,these patterns are conherpetofaunal ing sistent with the overall differencesbetween Sulawesi (Iskandar& Tjan 1996) and Borneo, Java,and Thailand (Manthey & Gr6ssmann 1997), furtherreinforcingthe notion that the herpetofaunaof Sulawesiis generally depauperatecomparedwith that of the main SundaShelf islandsand mainland SoutheastAsia. Sulawesiis geologicallyyounger than Borneo and other islands of the Sunda Shelf, and has been isolated from continentalAsia for at least 25 million yr (Audley-Charles1981, Voris2000). In contrast,Borneo was connected to continentalAsia during successive Pleistoceneglacialperiods (Morley& Fenley 1987, Inger& Voris2001). The isolation of Sulawesi from the mainland has provided the opportunity for evolution of endemic forms, and the dissected topography combined with possible Pleistocene and Holocene oceanic isolation of majorpeninsulas(Whitten et al. 2002), may have also promoted intraisland regionsof endemism within Sulawesi(Evanset al. 2003). However, there may have been less time for the evolution of more taxon-rich herpetofaunalassemblagesand less opportunityfor colonization by continental taxa, resultingin lower species richnessof communities within Sulawesi. Herpetofaunal checklists have also been reported for some other small islands in Southeast Asia, including Komodo, Indonesia (Auffenberg 1980), Singapore (Lim & Lim 1992), Tioman, Malaysia (Grismer et al. 2002), and Krakatauand Sebesi, Indonesia (Thornton et al. 2002). Taxonrichnessamongstthese islandsis highly variable,from 12 taxa on Krakatau(Thornton et al. 2002) to 163 taxa on Singapore (Lim & Lim 1992). This variabilityis influencedby biogeographicfactors (MacArthur& Wilson 1967, Inger& Voris2001), such as geologicalhistory, size, position within the archipelago,and proximity to other larger
Herpetofaunaof OffshoreIslandsof Sulawesi 287
Distance Bray-Curtis 0.0 CaveSpecialist
I
Estuary-Coastal Specialists
Riparian Specialists Moderately disturbed forest Minimally disturbed Forest
Minimally/Moderately disturbed Forest
ForestandRiparian
Generalists
HumanCommensals
0.2
0.4
0.6
0.8
1.0
newsp. 1 Cyrtodactylus Crocodylus porosus Cerberus rynchops Acrochordusgranulatus Lamprolepiscf. smaragdinum Cyrtodactylus new sp. 2 Emoia atrocostata Enyhdrisplumbea Rhacophorus cf. monticola sp. 2 Enhydrismatannensis Lygosoma quadrupes Bronchocoela cristellata Ptychozoan kuhlii Calamarianew sp 2 Sphenomorphus new sp. 2 Calamarianuchalis Xenopeltis unicolor Typhlops new sp. Cyclotyphlops deharvengi Psammodynastes pulverulentus Cylindrophismelanotus Calamariabrongersmai Amphiesma celebica Sphenomorphus tropidonotus Eutropisnew sp. Sphenomorphus new sp. 3 Lipinaquadrivittata Hemiphylodactylustypus Dibamus novaeguineae Rhacophorus georgii Kaloulabaleata Oreophrynenew sp. Tropodolaemuswagleri Calamarianew sp. 1 Oligodonwaandersi Cyrtodactylusjellesmae Boiga irregularis Rhabdophis chrysargoides Rhabdophis chrysargos Sphenomorphusnew sp. 1 Sphenomorphus variagatum Limnonectes cf. grunniens Gehyra new sp. Curoraamboinensis Xenochrophis trianguligerum Sphenomorphus cf. temmincki Rana chalconota Rhacophorus cf. monticola sp. 1 Lamprolepissmaragdinum Boiga dendrophila Fejervaryacancrivora Python reticulatus Dendrelaphispictus pictus Ahaetullaprasina Eutropisrudis Draco new sp. Varanussalvator Limnonectes cf. modestus Bufo celebensis Rana celebensis Polypedates leucomystax Hemidactylus frenatus Eutropismultifasciolata Ophiophagushannah Chrysopelea paradisi celebensis Ramphotyphlopsbraminus Gekko gecko Gehyra mutilata Cosymbotus platyurus Lygosoma bowringii Rana eurythraea Lycodon aulicus Elaphe euruthreacelebensis Elaphe jansenii
FIGURE6. Clusterdendrogramof dissimilaritiesin habitatuse amongtaxa.Bracketsand captionsat left explainthe ecologicalassociationsof majorgroupingsapparentwithin the dendrogram.
288 Gillespie,Howard,Lockie,Scroggie, and Boeadi
landmasses.Furthermore,our surveysof Buton, Kabaena,and Hoga are not consideredcomprehensive,and the comprehensivenessof other small island checklists is also likely to be highly variable.Therefore, detailed comparisonsof taxon richnessand assemblagecomposition among these small islandsacrossSoutheastAsia are not consideredmeaningful. ASSOCIATIONS.-Taxon richnesswas much higherin minimally HABITAT disturbedforestand foresthabitatswith only moderatedisturbancelevels than in highly modified or disturbedhabitats,such as secondaryforests, plantations, and villages. Coastal habitats also had low taxon richness, but these habitats were typically close to human settlement and often also highly disturbed. Pronounced shifts in herpetofaunalcommunity structurealso occurred from less disturbed forest habitats to more disturbed habitats. Most endemic taxawere restrictedto minimally or moderatelydisturbed forest. Many taxa found in these less disturbed habitatswere restricted to forest or riparianhabitatswithin forest. These include taxa that may be consideredto have relatively"specialized"habitat requirements,such as arborealtaxa, Rhacophorusspp., Tropidolaemus wagleri, Sphenomorphus tropidonotus,and Gehyranew sp., and fossorial/semifossorialtaxa, Calamaria spp., Cylindrophismelanotus,Dibamus novaeguineae,Sphenomorphusspp., and typhlopids. Most taxa found in more disturbed habitats occurred across a broader range of habitat types, with the exception of coastal habitats. Coastal habitats were dominated by taxa typicallyassociatedwith estuarineor foreshorehabitatsin SoutheastAsia (Manthey & Gr6ssmann 1997, Stuebing & Inger 1999, Thornton et al. 2002). A largegroup of habitat generalists(20 taxa), comprising mostly nonendemic taxa with wide distributions in Southeast Asia, occurred throughout forests and disturbed habitats alike. The few endemic taxa in this group, such as Bufo celebensis,appearto be ecological analogues of taxathat occupy these habitatselsewherein SoutheastAsia. Generally, disturbed habitats were characterizedby widespread habitat-generalist taxa, and forests were characterizedby endemic and habitat-specialist taxa. Most of the differencesobservedbetweenminimallyand moderately disturbed forest habitats were due to the occurrenceof rare or cryptic taxa that may be more widespread than indicated by our results. Two of the nonendemic taxa restrictedto moderatelydisturbedforest, Bronchocoelacritellataand Ptychozoonkuhlii, were found on a single occasion. Two of the three endemic taxa restrictedto minimally disturbed forest, new sp. 2 and Calamarianew sp. 2, were found on only Sphenomorphus one and four occasions, respectively.There is no clear discrimination of taxon composition or endemicity between minimally and moderately disturbed forest habitats.The forest herpetofaunacommunity may possess some resilienceto the variousdisturbanceprocessesoperatingin this region, suggesting that many forest-dwelling taxa may persist in spite of selective logging and other activities that do not greatly modify the overallstructureand microclimateof the forest. Selective timber harvestingis widespreadin this region and, apart from clearing for agriculture,is the most obvious anthropogenic disturbance to forest habitats. Clear-fellingis occurring in parts of Buton Island, but much of the timber harvestingoccurs at low intensity and involves the cutting of individualtreesof select taxaand size. Evidenceof this is most apparentin forestedareasnear human settlements. Further investigationis necessaryto determinethe impactsof variousdisturbance
processes,such as selectiveharvesting,on this herpetofaunacommunity before conclusions may be drawn about longer-term impacts on community structure. The discoveryof a significantnumber of new taxa and largerange extensions of some formerlydescribedtaxa undoubtedly reflectsundersampling of Sulawesiin the past. The resultsof this survey reaffirmthe need for more attention on this region for generalbiological surveyand research.Taxamost likely to be displacedby human impactstend to be endemic taxa, for which there exists little or no ecological information. However,seeminglyhigh levelsof persistenceof forestherpetofaunaltaxa in habitatswith moderatedisturbancelevelshave importantimplications for our understandingof ecological resiliencein tropical herpetofaunal communities. Becauseof the intractablepressuresof large human populations with economic dependencies on subsistencefarming and local forestresources,it is unrealisticto expectbiodiversityconservationobjectives to be achievedby setting aside protectedareasthat exclude harmful anthropogenicimpacts. The reality for future biodiversityconservation in SoutheastAsia is an increasingdependenceon managementof habitats with varyinglevelsof anthropogenicdisturbance.Consequently,a greater understandingof the relationshipsbetween forest structure,disturbance processes,and the biota is needed.
ACKNOWLEDGMENTS Funding and support for this work were provided by Operation Wallacea, Lincolnshire, United Kingdom and the Arthur Rylah Institute for Environmental Research, Victoria, Australia. Permission for this study was provided by Lembaga Ilmu Pengetahuan Indonesia (Indonesian Institute of Sciences) under permit nos. 3989/II/KS/2001 and 3224/SU/KS/2002, and Departemen Kehutanan DirektoratJenderal Perlindungandan KonservasiAlam (Department of ForestryDirectorateGeneralof The NaturalProtectionand Conservation),and the WallaceaDevelopment Institute,Jakarta.Additional assistanceand support were providedby PusatPenelitianPengembanganBiologi (Research and Development Centre for Biology); ForestryDepartment Southeast Sulawesi, Natural ResourcesConservation Unit of Southeast Sulawesi, Muna and Buton Section; Government of Buton Regency Culture and Tourism Department; Bidang Zoologi, Puslit Biologi-LIPI (Museum). We thankRafeBrownandJim McGuire,Universityof California,Berkeley; IndraneilDas, Universiti MalaysiaSarawak;Djoko Iskandar,Institute of Technology Bandung; Glen Shea, University of Sydney; and Allen Greer,AustralianMuseum for invaluablediscussions on the herpetofauna of Sulawesi and nomenclatorialadvice. We also thank Tim Coles, Steve Olliver,Mr. Ben, Bonny Mochamad Stafei, and Dedy Mudiana for logistical support throughout the project, and the community of Labundobundo, especially Usman and Alamin, for their invaluable field assistanceand hospitality.This manuscriptwas greatlyimprovedby comments from G. Brown, Lee Grismer,and one anonymous referee.
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APPENDIX NUMBERSOF SPECIMENS CATALOGUE LODGEDAT THE MUZEUM ZOO-
INDONESIA.-Bufocelebensis(Guenther 1859) BOGORINSE, LOGICUM MZB7034, 7035, 7686; Fejervaryacancrivora (Gravenhorst 1829) MZB7037, 7696-7699; Kaloula baleata (Mtiller 1836) MZB 7030, 7031, 7718; Limnonectes cf. grunniens (Daudin 1801) MZB7012-7018, 7028, 7029, 7709, 7710, 7724, 7726, 7732; Limnonectes cf. modestus (Boulenger 1982) MZB6995-7001, 7700-7702, 7722; Oreophryne
(Boettger 1895) new sp. MZB 7022-7027, 7711-7717; Polypedates leucomystax(Gravenhorst1829) MZB7091-7021, 7694, 7695, 8382, 8383; Rana celebensis (Peters 1872) MZB7007-7009,
7708, 8377; Rana
chalconota (Schlegel 1837) MZB7032, 7033, 7703-7707, 7721, 7723, 7728; Rana eurythraea (Schlegel 1837) MZB7690; Rhacophorus georgii (Roux 1904) MZB7689, 8329, 8381, 8378-8390,
Rhacophorus cf. mon-
ticola sp. 1 (Boulenger 1896) MZB7002-7006, 7672-7685, 76877689, 7719, 7720, 7725; Rhacophoruscf. monticolasp. 2 MZB7727; Dibamus novaeguineae(Dumiril & Bibron 1839) MZB3384, 42724279; Bronchocoelacristellata(Kuhl 1820) MZB4265; Draco (Linnaeus 3846,3847, 4266-4271; Cyrtodacty1758) new sp. MZB3704-3706, lus jellesmae (Boulenger 1897) MZB3713, 3714, 3844; Cyrtodactylus (Gray, 1827) new sp. 1 MZB3845, 4264; Cyrtodactylus new sp. 2 MZB3715, 3716; Gehyra mutilata (Wiegmann 1834) MZB3850; Gehyra (Gray 1834) new sp. MZB3696,
3848, 4263; Hemidactylus
290 Gillespie,Howard,Lockie,Scroggie, and Boeadi
frenatus (Dumrril & Bibron 1836) MZB3717, 4262; Varanussalvator ((Laurenti 1768) MZB3683, 3851; Emoia atrocostata(Lesson 1830) MZB3707, 4305; Eutropismultifasciata(Kuhl 1820) MZB3703, 3865-3868; Eutropisrudis (Boulenger 1887) MZB3697-3702, 38583864, 4306; Eutropis(Fitzinger1843) new sp. MZB3869-3874, 43074322; Lamprolepissmaragdinum(Lesson 1830) MZB3709-3711, 3854, 3855; Lamprolepiscf. smaragdinumMZB3712; Lipina quadrivittata (Peters 1867) MZB3708, 4280; Lygosomabowringii (Giinther 1864) MZB4283; Lygosomaquadrupes(Linnaeus 1766) MZB3843; Sphenomorphuscf. temminki(Duimeril& Bibron, 1839) MZB3684, 3695, tropidonotus(Boulenger 1897) MZB36923879-3882; Sphenomorphus 3694, 3853, 3856, 3857, 4290-4304; Sphenomorphusvariagatum (Peters 1872) MZB3685-3691, 3875-3878; Sphenomorphus (Fitzinger 1843) new sp. 1 MZB3883, 3852, 4281, 4282; Sphenomorphus new sp. 2 MZB4289; Sphenomorphusnew sp. 3 MZB4288; Acrochordusgranulatus (Schneider 1799) MZB2992; Amphiesmacelebica (Peters & Doria 1878) MZB2830, 2989; Boiga dendrophilaDumrril, Bibron & Dumbril 1854) DMZB2814, 2990; Boiga irregularis(Bechstein 1802) MZB2760, 2815-2817; Calamaria brongersmai(Inger & Marx 1965) MZB2758, 2759, 2843-2847, 2964-2966; Calamaria
nuchalis (Boulenger 1896) MZB2842, 28482962, 2963; Calamaria (Boie 1827) new sp. 1 MZB2753-2757, 2850-2857, 2967-2986, 2987; Calamaria (Boie 1827) new sp 2 MZB2849, 2987; Cerberus rynchops(Schneider 1799) MZB2820; Chrysopeleaparadisi celebensis (Mertens 1968) MZB2827, 2828; Cylindrophismelanotus(Wagler 1828) MZB2762, 2834, 2999; Dendrelaphispictuspictus (Gmelin 1789) MZB2811; Elaphe euruthreacelebensis(Jan 1863) MZB2761, 28242826; Elaphejansenii (Bleeker 1858)MZB2818, 2819, 2858; Enhydris matannensis(Boulenger 1897) MZB3000; Enhydrisplumbea (Boie 1827) MZB2823; Lycodonaulicus(Linnaeus1758) MZB2810; Oligodon waandersi(Bleeker 1860) MZB2812, 2831, 2832, 2991; Psammody(Boie 1827) MZB2833, 2763; Rhabdophischrysargos nastespulverulentus (GUnther 1858) (Schlegel 1837) MZB2835; Rhabdophischrysargoides MZB2809; Xenochrophistrianguligerus(Boie 1827) MZB2821-2822; waXenopeltisunicolor (Boie 1827) MZB2836, 2888; Tropidolaemus gleri (Wagler 1830) MZB2813, 2998, 3001; Ophiophagushannah (Cantor 1836) MZB2997; Cyclotyphlops deharvengi(Bosch & Ineich 1994) MZB2764-2765, 2837-2839, 3002-3004; Ramphotyphlops braminus(Daudin 1803) MZB2751, 2752, 2841, 2996, 3005; Typhlops (Oppel 1811) new sp. MZB2840, 3006
