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Herpetology Notes, volume 14: 1009-1025 (2021) (published online on 29 July 2021)

Creation of an and atlas for the Indonesian islands of and reveals gaps in sampling effort

Mirza D. Kusrini1,*, Amir Hamidy2, Lilik B. Prasetyo1, Rizky Nugraha1, Diana Andriani1, Nuzulul Fadhila1, Eko Hartanto1, and Andri Afrianto1

Abstract. Java and Bali are important as the most populated islands and as economic centres in , and Java is the seat of the national government. Although a series of reports exists to document the richness of and on those islands, there has so far been no unified compilation of their spatial distributions. The aim of this study was to quantify sampling effort and species richness for the herpetofauna of Java and Bali from specimen collections as well as published and unpublished literature sources, and to develop a map of spatial distributions as of 31 December 2017 – the first herpetofaunal atlas for any part of Indonesia. We found that the western part of Java has been better sampled than all other areas. Amphibian and reptile species richness on both islands is correlated with sampling effort, and sampling has occurred primarily in conservation areas. New species are still being described, not only from conservation areas but also from human- dominated landscapes. There is a need to increase the sampling effort on both islands, including outside of conservation areas. Development of a citizen science program focused on amphibians and reptiles would aid in increasing our understanding of species distributions in these islands and throughout Indonesia.

Keywords. Biodiversity, distribution map, herpetofauna, museum specimens, species richness, Southeast

Introduction in 1894 (Goss, 2004) to serve as temporary depository for specimens destined for Dutch museums (Iskandar It is well known that the collection of museum and Mumpuni, 2002), the Museum Zoologicum specimens is important for biodiversity assessment, Bogoriense (MZB) in Bogor, Indonesia, likely houses since the collection of a voucher specimen creates a the most complete collection of Indonesian amphibians permanent record of a location with a specific date and reptiles. Beginning in 2015, the development of attached to each specimen (Ponder et al., 2001). an Indonesian atlas of amphibians and reptiles is now Georeferencing of specimens has been a perennial underway and, so far, we have completed the work for task in many museums around the world, and these such an atlas for Java and Bali. data have been used for various purposes, including As the drivers of Indonesia’s economy, society, for analyses of species occurrence for many amphibian politics, and culture, Java (area ca. 128,300 km2) and and reptile groups (e.g., Herczeg et al., 2004; Horn et Bali (area ca. 5780 km2) have been transformed by al., 2007), predictions of their spatial distributions human occupation for hundreds of years (Whitten et al., (Ray et al., 2002; Penman et al., 2007), or to document 1996). The islands are both positioned on an extension biodiversity and hotspots in specific regions (Chen and of the continental shelf known as the Sunda Shelf and Bi, 2007; Sillero et al., 2014; Vasconcelos et al., 2014). were joined until the end of the last ice age, when they Developed as the Landbouw Zoologisch Laboratorium became separated by the Bali Strait, which is only about 2.4 km wide at its narrowest point (Clements et al., 2009). Bali represents the southeasternmost part of Sundaland (Johnson, 1964), separated from 1 Faculty of Forestry & Environment, IPB University, Darmaga, Lombok Island by the biogeographic divide known as 46, Cibinong 16911, Indonesia Cibinong 16911, Indonesia. 2 Museum Zoologicum Bogoriense, Research Center for Wallace’s Line (Hall, 2012). Although both islands are Biology, Indonesian Institute of Sciences, Widyasatwaloka already more developed than other islands of Indonesia, Building, Jalan Raya Jakarta Bogor Km. 46, Cibinong 16911, economic growth has not slowed in recent years (Hill Indonesia et al., 2008). As a result, both Java and Bali have lost * Corresponding author. E-mail: [email protected] most of their natural forest (Smiet, 1990; Whitten et al., © 2021 by Notes. Open Access by CC BY-NC-ND 4.0. 1996), yet the remaining natural areas, mostly under 1010 Mirza D. Kusrini et al. the protection of government as conservation areas or amphibians and reptiles on Java and Bali is needed as protected areas, are still important as the last strongholds reference to aid decision-making related to landscape of important endemic wildlife, including the Javan changes and species management. This knowledge Rhino (Rhinoceros sondaicus) in Ujung Kulon (Java) will support many applications in the future, including and the Balinese Starling (Leucopsar rothschildi) on spatial planning, environmental impact assessments, Bali Barat. and implementation of conservation measures. The study of amphibians and reptiles on Java and The objective of our study was to quantify and Bali dates to Dutch colonial times. Amphibians and identify patterns in species richness for amphibians reptiles of Java and Bali were listed and detailed and reptiles on Java and Bali, and to determine the in the earliest complete publications on Indonesian relationship between sampling effort and any observed herpetofauna written by Van Kampen (1923) and De patterns. To do this, we compiled information from Rooij (1915, 1917), respectively, and the work of De museum specimens, journal articles, and the grey Rooij was subsequently complemented and updated by literature to assess the distribution of amphibians and a checklist of (De Haas, 1950). After Indonesian reptiles on Java and Bali, and to estimate variation independence in 1945, reports on the herpetofauna in sampling effort. This study is part of a project of Java and Bali were scarce until the 1990s. It was to develop an atlas of amphibians and reptiles of not until 1998 that a book on amphibians of Java and Indonesia that can be used for future conservation and Bali was published (Iskandar, 1998). Subsequently, species management initiatives. several guidebooks about amphibians and reptiles were published for select locations on Java, in English as well Material and Methods as Indonesian (Kurniati, 2003; Yanuarefa et al., 2012; Kusrini, 2013; Rusli, 2016). A more comprehensive Study area. Java and Bali are two important islands in treatise on the herpetofauna of Bali and several other the western part of Indonesia and are administratively Lesser Sunda Islands was published by Mertens (1930) divided into seven provinces (Fig. 1). The current after his travels to the region, and field guides to the administrative area of Java includes six provinces herpetofauna of Bali were published by McKay (2006) (Banten, Jakarta, West Java, Central Java, Yogyakarta, and Somaweera (2017). and East Java), 86 districts and 28 municipalities, Most of the older publications on the herpetofauna whereas Bali has only one province (Bali) with eight of Java and Bali are based on specimens deposited districts and one municipality. in museum or university collections. There certainly Presence location data. Data for amphibians and is a wealth of information available from museum reptiles were compiled using data mostly from Museum specimens, in scientific journals, and in the grey Zoologicum Bogoriense (MZB) in Bogor (80.4% of literature that can be used to assess species numbers amphibian records and 85.6% of reptile records), with and distributions across the islands. Yet, no atlas of additional data from the scientific literature, amphibian the herpetofauna on these islands, nor for any island or reptile online databases, and unpublished theses, in Indonesia, has been published. The International reports and articles (e.g., the Indonesian herpetological Union for the Conservation of Nature (IUCN) has society newsletter, Warta Herpetofauna). Only 5% produced species distribution maps in their Red List of our compiled data are georeferenced (latitude and assessments, using a minimum convex polygon to longitude), whereas about 75% consist only of vague connect known localities and refining the distribution information (i.e., at the level of village or sub-district, according to specific criteria (i.e., suitable ; known as kecamatan). IUCN Standards and Petitions Subcommittee, 2017). For specimens collected in conservation areas (i.e., The resulting maps do not address if a given species national parks, nature reserves, wildlife reserves, is distributed uniformly within a polygon, occurs botanical gardens), we centred a reference point in everywhere in the polygon, or has a fragmented and the area and used it as the locality. Older museum skewed distribution, and they do not indicate areas that specimens are often labelled with generalized localities have not been surveyed. With Java and Bali currently (e.g., “Surabaya”, “Bogor”) or with only the name experiencing rapid infrastructure development that of the island (i.e., “Java” or “Bali”). We removed produces degradation of natural environments, such imprecise locality data from the analysis and detailed knowledge of the spatial distribution of distribution map. Gaps in Java and Bali Amphibian and Reptile Atlas 1011

Figure 1. Map of Java and Bali, Indonesia, showing the seven provinces in the region.

We compiled the general distribution of the species Distribution maps. Maps were developed using and the most recent using the databases of ArcGIS, version 10.5 (ESRI). We did not apply a Uetz et al. (2017), Frost (2017), and IUCN Redlist species modelling approach in the present study, (2017), with a cut-off date of 31 December 2017. instead we used 5 x 5 km grids as the smallest unit Data from museum vouchers were omitted from the for mapping the data on sampling effort and species dataset in cases of doubtful origin (e.g., when the richness onto the Indonesian Topography Map specimen would probably have come from a zoo or developed by the Geospatial Information Agency law enforcement confiscation). We omitted specimens (Badan Informasi Geospasial / BIG). Sampling effort of species not yet described, but we included the was estimated from the available data for each grid number of specimens from undescribed species for cell. Species richness was estimated as the count of mapping the sampling effort. We used the accession species that occurred in each grid cell. In total, Java date of museum specimens as a proxy for historical and Bali were divided into 7261 cells. sampling effort by grouping the accession dates into three periods: before 1945 (during the colonial period), Results 1946–80 (after independence up to start of local We collected 15,594 records, of which we omitted 2061 sampling efforts), and 1981–2017 (the modern era, amphibian and 1158 reptile records due to questionable coinciding with the establishment of Indonesian local authenticity or imprecise locality information. Of the herpetology). Using data grouped into districts and remaining 12,375 records, amphibians make up a municipalities, we compared sampling effort between greater portion (57%) than reptiles (43%). provinces using chi-square tests and examined the The percentage of grid cells on Java with the relationship between sampling effort and species occurrence of at least one amphibian species is 17.8%, richness using a regression analysis. and 53.4% of cells contain at least one reptile species. 1012 Mirza D. Kusrini et al.

Occurrence records for Bali are slightly higher with chalconota, Huia masonii, 42.2% for amphibians and 72.4% for reptiles. For Java, kuhlii, aurifasciatus, and Rhacophorus the highest numbers exist for the western part of Java margaritifer. Most of these species are widespread on (West Java and Banten Provinces), whereas for Bali the Java. Only F. limnocharis and H. chalconota are also highest number of records is from Gianyar Municipality. found on Bali. Both H. masonii and L. kuhlii occur Amphibian and reptile species richness of Java primarily in stream habitats near forested areas where and Bali. Based on current taxonomy, the herpetofauna they are considered common. of Java and Bali comprises a total species number of Ten species of reptiles accounted for 35% of records 238 (Table 1), including 43 species of amphibians from Java and Bali. This list includes (presented from (Appendix 1) and 195 species of reptiles (Appendix highest to lowest frequency): multifasciata, 2). Almost all Bali amphibians are also found on Java, Pseudocalotes tympanistriga, Hemidactylus frenatus, with the exception of Oreophryne monticola, a regional mutilata, haematopogon, Cyrtodactylus endemic of Bali and Lombok. Similarly, almost all of marmoratus, prasina, , Bali’s reptiles are present on Java, except for four species jubata, and . All of these, of (Cryptoblepharus renschi, Dibamus taylori, except for P. tympanistriga, D. haematopogon, and T. , Sphenomorphus vanheurni), three sexlineatus, are only distributed on Java, where they are species of snakes ( rynchops, Hypsiscopus considered common, and not on Bali. plumbea, Laticauda laticaudata), and one species of sea Sampling effort. Based on specimens in the MZB, the turtle (Caretta caretta). Three reptile species occur only oldest records for reptiles are from 1901 (one specimen on two small offshore islands. fusca (MZB Lace of Dogania subplana and two specimens of Malayemys 853, 11811–22) and Sphenomorphus melanopogon subtrijuga, collected by the naturalist Pieter Ouwens in (MZB Lace 848, 850, 12701–05, 12662–80) are Jakarta) and the oldest amphibian record is from 1903 (a reported only from Tinjil Island, Banten Province (16 specimen of Occidozyga lima from Sukabumi, collected km south of Java), and caudolineatus by an unknown collector identified only by the single has been collected (MZB Lace 1069) only on Kangean name Alie; MZB 21189). Sampling effort for reptiles and Island, East Java Province (120 km north of Bali). amphibians differed among periods (c2 = 285.5, df = 2, Six species represented > 50% of records in the n = 15,439; P < 0.001). There are more specimens from amphibian database. These include (listed from the modern period (10,524) than from both the post- high to lowest frequency): Fejervarya limnocharis, independence (3806) and the colonial periods (1109). Sampling effort for amphibians differed among provinces (c2 = 25.06, df = 6, n = 203; P < 0.001). Table 1. Number of species of amphibian and reptiles recorded on Java and Bali. Species occurring only on The highest sampling effort for amphibians on Java one island are listed under that island’s name, whereas occurred primarily in the western part of the island species found on both islands are in the J+B column. (Fig. 2A), including in conservations areas such as Mount Halimun Salak National Park (2543 collected Taxon Java Bali J + B specimens) and Mount Gede Pangrango National Park (1653) in West Java Province, and in Ujung Kulon National Park (423) in Banten Province. West Java AMPHIBIANS 41 15 43 and Banten (7869) accounted for 90% of the entire 40 15 42 sampling effort for amphibians, whereas numerous Caecilians 1 0 1 gaps occurred in other provinces. Sampling of amphibians on Bali generated 297 specimens, with the REPTILES 187 72 195 highest proportions from Tabanan (42.8%), Buleleng Turtles 14 8 16 (24.6%) and Gianyar (12.8%) Municipalities. Crocodiles 3 1 3 Sampling of reptiles (6435 specimens) was slightly and 52 27 56 lower than for amphibians (9123) and more evenly Monitor lizards 3 1 3 distributed (Fig. 2B). For reptiles, we found no Snakes 115 35 117 difference in sampling effort among provinces (c2 = 10.5, df = 6, n = 332; P = 0.100), although there is a TOTAL 228 87 238 general trend indicating higher sampling effort in West

Gaps in Java and Bali Amphibian and Reptile Atlas 1013

Figure 2. Sampling effort for amphibians (A) and reptiles (B) on Java and Bali, inferred from the number of collected specimens in the MZB collection augmented by selected data from unpublished and published reports. Each pixel represents the number of specimens collected within a 5 x 5 km grid cell. The time span for data collection is 113 years for amphibians and 115 years for reptiles.

Java (3476). Similarly, the highest sampling effort for Relationship between species richness and reptiles was in the western part of Java, specifically in the sampling effort. We found a moderate but significant national parks of Gede Pangrango (638), Halimun Salak positive relationship (r2 = 0.263, P < 0.001) between (415), and Ujung Kulon (522). West Java and Banten sampling effort and amphibian species richness on Java Provinces (4189) accounted for about 66% of the entire and Bali. Species richness increased with sampling sampling effort for reptiles. Sampling effort for reptiles effort. West Java and Banten Provinces exhibited in the eastern part of Java was higher (13.5%) than that the highest amphibian species richness on Java. The for amphibians (3.5%). Sampling effort in East Java was hotspots of amphibian species richness on the island also concentrated in conservation areas, such as Alas were the national parks of Halimun Salak (29 species) Purwo National Park (86). Sampling effort for reptiles and Gede Pangrango (27) in West Java, followed by on Bali was highest in three municipalities: Klungkung Ujung Kulon in Banten (19) and Alas Purwo in East (19.4%), Gianyar (17.2%), and Tabanan (16.7%). Java (14). Bali generally has a lower amphibian species 1014 Mirza D. Kusrini et al.

Figure 3. Percentage of amphibian (A) and reptile (B) species richness on Java and Bali inferred from the number of collected specimens in the MZB augmented by data from unpublished and published reports. Each pixel represents the number of species recorded within a 5 x 5 km grid from the earliest georeferenced specimens until 2016. richness, and the highest species richness was found in Bali, on Nusa Penida Island (22), and in the western Gianyar Municipality (11 species; Fig. 3A). part of the Gianyar Municipality (15; Fig. 3B). There was also a strong relationship between sampling effort and species richness for reptiles (r2 = 0.674, P < Discussion 0.001). Species richness again increased with sampling Our results show a relationship between sampling effort. The highest reptile species richness on Java effort and species richness and, on the other hand, was found in West Java and Banten Provinces. The reveal sampling gaps, particularly in the eastern part of hotspots of reptile species richness on Java were the Java. Sampling gaps and biased sampling effort have national parks of Ujung Kulon in Banten (57 species), been demonstrated for other groups in other parts of the followed by Gede Pangrango (56) and Halimun Salak world. For instance, Oliveira et al. (2017) studied the (56) in West Java, and Alas Purwo in East Java (34). spatial distribution of spider species in Brazil and found Bali has an overall lower reptile species richness, and a high concentration of records in only a few locations, the highest species richness was found in Bali Barat with higher sampling effort evident near cities where National Park (25 species) at the northwestern tip of museums and major universities were located. This Gaps in Java and Bali Amphibian and Reptile Atlas 1015

“museum effect” of increased sampling effort in close expeditions (HIMAKOVA, 2011, 2014) from our list vicinity to major museums has been described in detail of species from Java and Bali. (Ponder et al., 2001; Werneck et al., 2011; Oliveira et Increasing sampling effort has led to new distribution al., 2017). A similar pattern is obvious in West Java, records and new species of amphibian and reptiles on where the MZB, the leading institution in Indonesia for Java and Bali, including new records of Polypedates natural history since the late 19th Century, is located, otilophus, a species previously thought to be an endemic and where the highest number of amphibian and reptile of (Riyanto et al., 2009). Increased sampling species for Java have been recorded. effort has also confirmed the occurrence of several Another possible explanation for sampling bias is the amphibian species thought to have vanished, including tendency to survey conservation areas and areas with Ichthyophis hypocyaneus (Kusrini et al., 2017), easy access. On both Java and Bali, the areas of highest Philautus pallidipes (Mumpuni, 2002), and Chiromantis sampling effort are conservations areas. In addition, vittiger (Wostl et al., 2017). After the completion of the higher sampling effort on Bali is also located in our analysis, a new species, Leptophryne javanica, was areas considered scenic or attractive to tourists, such as described from Mount Ciremai National Park (Hamidy Tabanan Municipality, where the Bali Botanic Garden et al., 2018) and a new species of rock gecko, Cnemapis is located, and Gianyar Municipality, where the famed muria, was described from Mount Muria in Central Java Ubud Village is located. (Riyanto et al., 2019). Other studies have also improved Higher sampling effort in the modern era might also taxonomic knowledge of other amphibian and reptiles in be partly caused by the availability of identification Java. For instance, P. otilophus is now considered only books, especially in Indonesian. However, there are distributed in Borneo, and species in Java and more identification books on the amphibian fauna formerly known as P. otilophus have been assigned to (e.g., Liem, 1971; Iskandar, 1998; Kurniati, 2003; P. pseudotilophus (Matsui et al., 2014). Cyrtodactylus Kusrini, 2013) than the reptile fauna. The increase in klakahensis is now considered a junior of C. the number of studies on amphibians and reptiles by petani (Riyanto et al., 2020). local university students since the 2000s correlates with Although our results highlight the importance of the foundation of the Indonesian Herpetological Society national parks for the maintenance of herpetofaunal (Eprilurahman, 2008). Most of the research is carried out diversity, new species, such as Chiromantis trilaksonoi by universities on Java, including extensive exploration (Riyanto and Kurniati, 2014), Fejervarya iskandari and documentation of herpetofaunal diversity in various (Veith et al., 2001), and orientalis locations. Only a small part of this research is actually (Matsui et al., 2013) were all found in rice paddies. published in journals, with most of the findings stored Anthropogenically altered landscapes also need to be as grey literature (i.e., students’ reports written in sampled, as they can serve as for many species Indonesian). Data from the grey literature must be used (big cats in – Athreya et al., 2013; – Daily with care as these often do not give enough evidence for et al., 2001), including amphibians (e.g., Hodgkison et the occurrence of certain species. al., 2007) and snakes (Shine and Koenig, 2001). Even in the published literature there are some There is a possibility that some species have been uncertainties regarding the occurrence of certain extirpated from areas or are extinct. Several amphibian species on Java and Bali. For instance, McKay (2006) species have low recorded numbers, such as Hylarana reported the occurrence of Psammophis indochinensis baramica (one specimen at MZB from Bogor, West Java) in Negara (Bali), but Hartmann et al. (2011) questioned or Philautus jacobsoni (no specimens at MZB). About this record and stated that molecular studies are 20% of the reptiles on Java and Bali are represented needed to confirm the occurrence of the species in by only a single record, and in some cases the voucher Indonesia. Therefore, and following Uetz et al. (2017), specimens are held in museums outside of Indonesia, we omitted P. indochinensis from our list. The absence such as Sphenomorphus necopinatus (specimens in of voucher specimens may also cause difficulties with Leiden, The Netherlands) and Harpesaurus tricinctus species confirmation, especially when photographs (specimens in London, United Kingdom). of species do not show diagnostic features for There is also evidence of an increase in the distribution identification to species level. For this reason, we also of some invasive species, such as the American Bullfrog omitted albiventer, signatus, and (Lithobates catesbeianus). Introduced to Indonesia , which were reported by student as part of the effort to increase the production of 1016 Mirza D. Kusrini et al. meat for human consumption, L. catesbeianus were incorporate strategies to increase the accuracy of the data formerly raised in closed farms (Kusrini and Alford, (Freitag et al., 2016). Web-based programs that also act 2006). Iskandar (1998) reported that the species could as data platforms, such as iNaturalist, allow verification be found in many cities but did not indicate whether by other users, especially for identification of species. it was found in the wild or on farms. Our data show However, both Freitag et al. (2016) and Aceves-Buenos that this frog can nowadays be found in the wild, both et al. (2017) highlighted the need for training sessions on Java and Bali. Other prospective invasive species when employing citizen scientist to enhance accuracy include the (Trionyx chinensis) and credibility. We recommend that the herpetological and the Red-eared Slider (Trachemys scripta elegans). community in Indonesia, which primarily consists of Both species are now captively bred in Indonesia, as young university students, adopt this approach when food and pets (Kusrini, 2011). There is a report of developing the amphibian and reptile citizen science T. chinensis (Sunyoto, 2012) and there are informal program. This proposed program for Indonesia will reports of T. scripta elegans occurring in the wild. increase the capacity of young herpetologists and The availability of historical data is invaluable, as it also build networks between professional and non- can indicate the presence or absence of a species in a professional herpetologists in accordance with the certain location at a particular time and can give clues to recommendation of Theobald et al. (2015) regarding changes in the range of a species in the recent past. For the need for collaboration between scientists and non- instance, Church (1960) reported findingDuttaphrynus scientists to combat large-scale environmental issues. melanostictus on the western tip of Bali but not in other parts of the island. He concluded that this observation Acknowledgements. This work was made possible by funding in 1958 was evidence of a new dispersal event from from the National Geographic Foundation for Science and Java and predicted that the species would spread more Exploration – Asia to the Java-Bali Herp CARE (Conservation, Awareness and Research) Initiatives program (grant number widely. The result of our study show that the Asia-16-15 to M.D. Kusrini). We are grateful for the assistance of has spread widely on Bali, even to the island of Nusa staff at the Museum Zoologicum Bogoriense for access given to Penida. More recently, D. melanostictus has expanded the team, the IPB 2017 mobility program (BPPTN BH 2017), and its range to other parts of Wallacea, raising concerns Jodi Rowley who enabled M.D. Kusrini to write the early draft of that the presence of this toad may have an impact on this manuscript at the Australian Museum, Sydney. We thank N. local endemic species (Reilly et al., 2017). 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Appendix 1. List of amphibian species found on Java and Bali, based on specimens deposited in the MZB collection (1) or literature reports (2). We use the plus sign (+) to denote the presence of a species. The total number of amphibian species on these islands is 43, with several species occurring on both islands.

Taxon Java Bali Source

FROGS Bufonidae Duttaphrynus melanostictus (Schneider, 1799) + + 1, 2 biporcatus (Gravenhorst, 1829) + + 1, 2 Ingerophrynus parvus (Boulenger, 1887) + 1, 2 Leptophryne borbonica (Tschudi, 1838) + 1, 2 Leptophryne cruentata (Tschudi, 1838) + 1, 2 Leptophryne javanica Hamidy et al., 2018 + 1, 2 asper (Gravenhorst, 1829) + 1, 2

Dicroglossidae Fejervarya cancrivora (Gravenhorst, 1829) + + 1, 2 Fejervarya iskandari Veith et al., 2001 + 2 Fejervarya limnocharis (Gravenhorst, 1829) + + 1, 2 Limnonectes kuhlii (Tschudi, 1838) + 1, 2 Limnonectes macrodon (Duméril & Bibron, 1841) + 1, 2 Limnonectes microdiscus (Boettger, 1892) + 1, 2 Occidozyga lima (Gravenhorst, 1829) + + 1, 2 Occidozyga sumatrana (Peters, 1877) + + 1, 2

Megophryidae Leptobrachium hasseltii Tschudi, 1838 + + 1, 2 montana Kuhl & Van Hasselt, 1822 + 1, 2

Microhylidae baleata (Müller, 1836) + + 1, 2 Kalophrynus minusculus Iskandar, 1998 + 1, 2 Kalophrynus pleurostigma Tschudi, 1838 + 1, 2 Microhyla achatina Tschudi, 1838 + 1, 2 Microhyla orientalis Matsui et al., 2013 + 1, 2 Microhyla palmipes Boulenger, 1897 + + 1, 2 Oreophryne monticola (Boulenger, 1897) + 1, 2

Ranidae Amnirana nicobariensis (Stoliczka, 1870) + + 1, 2 chalconota (Schlegel, 1837) + + 1, 2 Chalcorana rufipes (Inger et al., 2009) + 1, 2 Pulchrana baramica (Boettger, 1900) + 1 Hylarana erythraea (Schlegel, 1837) + 1, 2 Huia masonii (Boulenger, 1884) + 1, 2 Lithobates catesbeianus (Shaw, 1802) + + 2 Odorrana hosii (Boulenger, 1891) + 1, 2

Rhacophoridae Chiromantis trilaksonoi Riyanto & Kurniati, 2014 + 1, 2 Chiromantis vittiger (Boulenger, 1897) + 1, 2 1020 Mirza D. Kusrini et al.

Appendix 1. cont.

Taxon Java Bali Source

Rhacophoridae (cont.) Nyctixalus margaritifer Boulenger, 1882 + 1, 2 Philautus aurifasciatus (Schlegel, 1837) + 1, 2 Philautus jacobsoni (Van Kampen, 1912) + 1, 2 Philautus pallidipes (Barbour, 1908) + 1, 2 Polypedates leucomystax (Gravenhorst, 1829) + + 1, 2 Polypedates pseudotilophus Matsui et al., 2014 + 1, 2 Rhacophorus margaritifer (Schlegel, 1837) + 1, 2 Rhacophorus reinwardtii (Schlegel, 1840) + 1, 2

CAECILIANS Ichthyophis hypocyaneus (Van Hasselt in Boie, 1827) + 1, 2

TOTALS 41 15 43

Appendix 2. List of the reptile species found on Java and Bali, based on specimens deposited in the MZB collection (1), on literature reports (2), or on the information in Uetz et al. (2017) (3). We use the plus sign (+) to indicate the presence of a species. The total number of reptile species on these islands is 195, with several species occurring on both islands.

Taxon Java Bali Source

CROCODILES Crocodylidae Crocodylus porosus Schneider, 1801 + + 2 Crocodylus siamensis Schneider, 1801 + 3 Tomistoma schlegelii (Müller, 1838) + 3

LIZARDS (Kuhl, 1820) + + 1, 2 Duméril & Bibron, 1837 + + 1, 2 Calotes versicolor (Daudin, 1802) + 1, 2 Draco cornutus Günther, 1864 + 3 Kuhl, 1820 + + 1, 2 Draco haematopogon Gray, 1831 + 1, 2 Draco lineatus Daudin, 1802 + 2 Draco maximus Boulenger, 1893 + 3 Draco volans Linnaeus, 1758 + + 1, 2 chamaeleontinus (Laurenti, 1768) + 1, 2 Gonocephalus kuhlii (Schlegel, 1848) + 1, 2 Harpesaurus tricinctus (Duméril, 1851) + 3 Pseudocalotes tympanistriga (Gray, 1831) + 1, 2 Pseudocophotis sumatrana (Hubrecht, 1879) + 3

Dibamidae Dibamus taylori Greer, 1985 + 2 Gaps in Java and Bali Amphibian and Reptile Atlas 1021

Appendix 2. cont.

Taxon Java Bali Source

Gekkonidae Cnemaspis muria Riyanto et al., 2019 + 1, 2 Cyrtodactylus fumosus (Müller, 1895) + + 1, 2 Cyrtodactylus marmoratus Gray, 1831 + + 1, 2 Cyrtodactylus petani Riyanto et al., 2015 + 1, 2 Cyrtodactylus semiadii Riyanto et al., 2014 + 1, 2 Gekko kuhli (Stejneger, 1902) + 1, 2 (Wiegmann, 1834) + + 1, 2 Gekko gecko (Linnaeus, 1758) + + 1, 2 (Schlegel, 1836) + 1, 2 Gray, 1842 + 1, 2 Gekko vittatus Houttuyn, 1782 + 1 Hemidactylus frenatus Duméril & Bibron, 1836 + + 1, 2 Hemidactylus garnotii Duméril & Bibron, 1836 + + 1, 2 Hemidactylus platyurus (Schneider, 1797) + + 1, 2 typus Bleeker, 1860 + + 1, 2 (Duméril & Bibron, 1836) + 1

Lacertidae Takydromus sexlineatus Daudin, 1802 + 1, 2

Scincidae (Duméril & Bibron, 1839) + 1 Carlia nigrauris Zug, 2010 + 3 Cryptoblepharus balinensis Barbour, 1911 + + 1, 2 Cryptoblepharus cursor Barbour, 1911 + + 1, 2 Cryptoblepharus renschi Mertens, 1928 + 1,2 olivacea Gray, 1839 + + 1 atrocostata (Lesson, 1830) + 1 (De Vis, 1892 + 1 Eutropis macrophthalma (Mausfeld & Böhme, 2002) + 3 (Kuhl, 1820) + + 1, 2 Eutropis rudis (Boulenger, 1887) + + 1, 2 (Stoliczka, 1870) + + 1, 2 Lamprolepis leucosticta (Müller, 1923) + 3 (Lesson, 1826) + 1, 2 Lipinia relicta (Vinciguerra, 1892) + 1 bowringii (Günther, 1864) + + 1, 2 Lygosoma quadrupes (Linnaeus, 1766) + + 1, 2 Sphenomorphus melanopogon (Duméril & Bibron, 1839) + 1 Sphenomorphus necopinatus (Brongersma, 1942) + 3 Sphenomorphus puncticentralis Iskandar, 1994 + 1, 2 Sphenomorphus sanctus (Duméril & Bibron, 1839) + + 1, 2 Sphenomorphus vanheurni (Brogersma,1942) + 2 Tytthoscincus temmincki (Duméril & Bibron, 1839) + 1, 2 Tytthoscincus temmincki Dumeril & Bibron, 1839) + + 3 1022 Mirza D. Kusrini et al.

Appendix 2. cont.

Taxon Java Bali Source

Varanidae Varanus bengalensis (Daudin, 1802) + 2 Varanus nebulosus (Gray, 1831) + 1, 2 Varanus salvator (Laurenti, 1768) + + 1, 2

SNAKES Acrochordidae granulatus (Schneider, 1799) + + 1, 2 Acrochordus javanicus Hornstedt, 1787 + 1

Colubridae Ahaetulla mycterizans (Linnaeus, 1758) + 1 Ahaetulla prasina (Boie, 1827) + + 1, 2 cynodon (Boie, 1827) + + 1, 2 (Boie, 1827) + + 1, 2 (Boie, 1827) + 1, 2 Boiga jaspidea (Duméril et al., 1854) + 1 Boiga multomaculata (Boie, 1827) + + 1, 2 Boiga nigriceps (Günther, 1863) + 1, 2 Calamaria bicolor Duméril et al., 1854 + 1 Calamaria javanica Boulenger, 1891 + 3 Calamaria lateralis Mocquard, 1890 + 3 Calamaria leucogaster Bleeker, 1860 + 3 Calamaria linnaei Boie, 1827 + + 1, 2 Calamaria lovii Boulenger, 1887 + 3 Boie, 1827 + 1, 2 Calamaria modesta Duméril et al., 1854 + 1 Calamaria pavimentata Duméril et al., 1854 + 3 Calamaria schlegeli Duméril et al., 1854 + + 1, 2 Calamaria virgulata Boie, 1827 + 1 paradisi Boie, 1827 + + 1, 2 Chrysopelea pelias (Linnaeus, 1758) + 3 Coelognathus erythrurus (Duméril et al., 1854) + 1 Coelognathus flavolineatus (Schlegel, 1837) + + 1, 2 Coelognathus radiatus (Boie, 1827) + + 1, 2 Dendrelaphis caudolineatus (Gray, 1834) + 1 (Boie, 1827) + 1, 2 Dendrelaphis haasi Van Rooijen & Vogel, 2008 + 3 Dendrelaphis pictus (Gmelin, 1789) + + 1, 2 Dendrelaphis subocularis (Boulenger, 1888) + 3 Dendrelaphis underwoodi Van Rooijen & Vogel, 2008 + 1 Dryophiops rubescens (Gray, 1834) + 1, 2 Elapoidis fusca Boie, 1827 + 1, 2 Gongylosoma baliodeirus (Boie, 1827) + + 1, 2 Gongylosoma longicaudum (Peters, 1871) + 1 oxycephalum (Boie, 1827) + + 1, 2 Gaps in Java and Bali Amphibian and Reptile Atlas 1023

Appendix 2. cont.

Taxon Java Bali Source

Colubridae (cont.) (Boie, 1827) + + 1, 2 Liopeltis tricolor (Schlegel, 1837) + 1, 2 capucinus Boie, 1827 + + 1, 2 Lycodon subcinctus Boie, 1827 + + 1, 2 Oligodon bitorquatus Boie, 1827 + 1, 2 Oligodon octolineatus (Schneider, 1801) + + 1, 2 Oligodon purpurascens (Schlegel, 1837) + 1, 2 carinata (Günther, 1858) + 1 (Schlegel, 1837) + + 1, 2 (Linnaeus, 1758) + + 1, 2 Sibynophis geminatus (Boie, 1826) + + 1, 2 Sibynophis melanocephalus (Gray, 1835) + 1, 2 Tetralepis fruhstorferi Boettger, 1892 + 1 javanicus Reinhardt, 1836 + 2

Cylindropiidae ruffus (Laurenti, 1768) + 1, 2 Cylindrophis subocularis Kieckbusch et al., 2016 + 3

Elapidae (Gray, 1849) + 1 candidus (Linnaeus, 1758) + + 1, 2 Bungarus fasciatus (Schneider, 1801) + 1, 2 Bungarus flaviceps Reinhardt, 1843 + 3 bivirgatus (Boie, 1827) + 1 (Laurenti, 1768) + 1, 2 atriceps Günther, 1864 + 1 Hydrophis brookii Günther, 1872 + 3 (Shaw, 1802) + 1 (Shaw, 1802) + 1 Hydrophis cyanocinctus (Daudin, 1803) + 1 (Gray, 1842) + 1 (Schneider, 1799) + 1 (Shaw, 1802) + 1 Hydrophis hardwickii (Gray, 1834) + 1 Hydrophis inornatus (Gray, 1849) + 3 Gray, 1849 + 1 (Gray, 1842) + 1 Hydrophis platurus (Linnaeus, 1766) + 1, 2 Hydrophis schistosus Daudin, 1803 + 1 (Shaw, 1802) + 1 Hydrophis stokesii (Gray, 1846) + 1 Hydrophis viperinus (Schmidt, 1852) + 3 Kolpophis annandalei (Laidlaw, 1901) + 3 Laticauda colubrina (Schneider, 1799) + + 1, 2 1024 Mirza D. Kusrini et al.

Appendix 2. cont.

Taxon Java Bali Source

Elapidae (cont.) Laticauda colubrina (Schneider, 1799) + + 1, 2 Laticauda laticaudata (Linnaeus, 1758) + 1 Naja sputatrix Boie, 1827 + + 1, 2 Ophiophagus hannah (Cantor, 1836) + + 1, 2 Thalassophis anomalus (Schmidt, 1852) + 1

Gerrhopilidae ater (Schlegel, 1839) + + 1 Gerrhopilus bisubocularis (Boettger, 1893) + 3

Homalopsidae Cerberus schneiderii (Schlegel, 1837) + + 1, 2 Enhydris enhydris (Schneider, 1799) + 1, 2 leucobalia (Schlegel, 1837) + 1, 2 buccata (Linnaeus, 1758) + 1, 2 Hypsiscopus plumbea (Boie, 1827) + + 1, 2 Miralia alternans (Reuss, 1834) + 1 Myrrophis bennettii (Gray, 1842) + 3 Phytolopsis punctata Gray, 1849 + 3 Sumatranus albomaculata (Duméril et al., 1854) + 3

Natricidae Fowlea melanzostus (Gravenhorst, 1807) + + 1,2 Rhabdophis chrysargoides (Günther, 1858) + 1 (Schlegel, 1837) + + 1,2 Rhabdophis rhodomelas (Boie, 1827) + 3 (Schlegel, 1837) + 1,2 Xenochrophis trianguligerus (Boie, 1827) + 1,2 Xenochrophis vittatus (Linnaeus, 1758) + 1,2

Pareidae boa (Boie, 1828) + 1,2 Asthenodipsas laevis (Boie, 1827) + 1, 2 Pareas carinatus (Boie, 1828) + + 1,2

Pseudaspididae Psammodynastes pulverulentus (Boie, 1827) + + 1,2

Pseudoxenodontidae Pseudoxenodon inornatus (Boie, 1827) + 1, 2

Pythonidae Malayopython reticulatus (Schneider, 1801) + + 1, 2 Python bivittatus Kuhl, 1820 + + 1, 2

Typhlopidae Argyrophis fuscus (Duméril, 1851) + 3 Argyrophis muelleri (Schlegel, 1839) + 3 Gaps in Java and Bali Amphibian and Reptile Atlas 1025

Appendix 2. cont.

Taxon Java Bali Source

Typhlopidae (cont.) Argyrophis fuscus (Duméril, 1851) + 3 Argyrophis muelleri (Schlegel, 1839) + 3 braminus (Daudin, 1803) + + 1, 2 Ramphotyphlops lineatus (Schlegel, 1839) + 1, 2

Viperidae rhodostoma (Kuhl, 1824) + 1,2 (Smith, 1917) + 1 Gray, 1842 + 1,2 Kramer, 1977 + 1,2 (Boie, 1827) + 1,2

Xenopeltidae Xenelaphis hexagonotus (Cantor, 1847) + 3 unicolor Reinwardt, 1827 + 1,2

TURTLES Cheloniidae Caretta caretta (Linnaeus, 1758) + 2 Chelonia mydas (Linnaeus, 1758) + + 2 Eretmochelys imbricata (Linnaeus, 1766) + + 2 Lepidochelys olivacea (Eschscholtz,1829) + + 1, 2

Dermochelidae Dermochelys coriacea (Vandelli, 1761) + + 2

Emydidae Trachemys scripta (Thunberg in Schoepff, 1792) + 2

Geoemydidae Cuora amboinensis (Daudin, 1802) + + 2 Cyclemys dentata (Gray, 1831) + 2 Malayemys subtrijuga (Schlegel & Müller, 1845) + 2 Notochelys platynota (Gray, 1834) + 2

Testudinidae Siebenrockiella crassicollis (Gray, 1831) + 2

Trionychidae Amyda cartilaginea (Boddaert, 1770) + + 1, 2 Dogania subplana (Geoffroy Saint-Hilaire, 1809) + 2 Chitra chitra Nutaphand, 1986 + 1 Pelochelys cantorii Gray, 1864 + 2 Pelodiscus sinensis (Wiegmann, 1835) + 2

TOTALS 186 71 195

Accepted by Hinrich Kaiser