Creation of an amphibian and reptile atlas for the Indonesian islands of Java and Bali reveals gaps in sampling effort
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Herpetology Notes, volume 14: 1009-1025 (2021) (published online on 29 July 2021)
Creation of an amphibian and reptile atlas for the Indonesian
islands of Java and Bali 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 Indonesia, and Java is the
seat of the national government. Although a series of reports exists to document the species richness of amphibians and
reptiles 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published and unpublished literature sources, and to develop a map of spatial distributions as of 31 December 2017 – the first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 Asia
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,
46, Cibinong 16911, Indonesia Cibinong 16911, Indonesia.
Lombok Island by the biogeographic divide known as
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: mirza_kusrini@yahoo.com most of their natural forest (Smiet, 1990; Whitten et al.,
© 2021 by Herpetology 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 snakes (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 habitats; 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 taxonomy 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 Hylarana chalconota, Huia masonii, Limnonectes
42.2% for amphibians and 72.4% for reptiles. For Java, kuhlii, Philautus 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): Eutropis multifasciata,
2). Almost all Bali amphibians are also found on Java, Pseudocalotes tympanistriga, Hemidactylus frenatus,
with the exception of Oreophryne monticola, a regional Gehyra mutilata, Draco haematopogon, Cyrtodactylus
endemic of Bali and Lombok. Similarly, almost all of marmoratus, Ahaetulla prasina, Takydromus sexlineatus,
Bali’s reptiles are present on Java, except for four species Bronchocela jubata, and Gekko gecko. All of these,
of lizards (Cryptoblepharus renschi, Dibamus taylori, except for P. tympanistriga, D. haematopogon, and T.
Draco lineatus, Sphenomorphus vanheurni), three sexlineatus, are only distributed on Java, where they are
species of snakes (Cerberus 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. Carlia 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 Dendrelaphis 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
Frogs 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
Skinks and Geckos 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 WestGaps 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. ThisGaps 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 Borneo (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 Sumatra 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 synonym 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 Microhyla 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 habitat for many species Indonesian). Data from the grey literature must be used (big cats in India – Athreya et al., 2013; birds – 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 Calamaria albiventer, Oligodon signatus, and (Lithobates catesbeianus). Introduced to Indonesia Sibynophis collaris, which were reported by student as part of the effort to increase the production of frog
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 Chinese Softshell Turtle (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 finding Duttaphrynus 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
Exploration – Asia to the Java-Bali Herp CARE (Conservation,
Java and predicted that the species would spread more
Awareness and Research) Initiatives program (grant number
widely. The result of our study show that the toad
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). Karraker for constructive discussions and an anonymous reviewer
and H. Kaiser during the development of this manuscript.
The database developed as a part of our research
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Gaps in Java and Bali Amphibian and Reptile Atlas 1019
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
Ingerophrynus 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
Phrynoidis 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
Megophrys montana Kuhl & Van Hasselt, 1822 + 1, 2
Microhylidae
Kaloula 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
Chalcorana 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, 21020 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
Agamidae
Bronchocela cristatella (Kuhl, 1820) + + 1, 2
Bronchocela jubata Duméril & Bibron, 1837 + + 1, 2
Calotes versicolor (Daudin, 1802) + 1, 2
Draco cornutus Günther, 1864 + 3
Draco fimbriatus 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
Gonocephalus 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 + 2Gaps 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
Gehyra mutilata (Wiegmann, 1834) + + 1, 2
Gekko gecko (Linnaeus, 1758) + + 1, 2
Gekko monarchus (Schlegel, 1836) + 1, 2
Gekko smithii 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
Hemiphyllodactylus typus Bleeker, 1860 + + 1, 2
Lepidodactylus lugubris (Duméril & Bibron, 1836) + 1
Lacertidae
Takydromus sexlineatus Daudin, 1802 + 1, 2
Scincidae
Carlia fusca (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
Dasia olivacea Gray, 1839 + + 1
Emoia atrocostata (Lesson, 1830) + 1
Emoia caeruleocauda (De Vis, 1892 + 1
Eutropis macrophthalma (Mausfeld & Böhme, 2002) + 3
Eutropis multifasciata (Kuhl, 1820) + + 1, 2
Eutropis rudis (Boulenger, 1887) + + 1, 2
Eutropis rugifera (Stoliczka, 1870) + + 1, 2
Lamprolepis leucosticta (Müller, 1923) + 3
Lamprolepis smaragdina (Lesson, 1826) + 1, 2
Lipinia relicta (Vinciguerra, 1892) + 1
Lygosoma 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) + + 31022 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
Acrochordus granulatus (Schneider, 1799) + + 1, 2
Acrochordus javanicus Hornstedt, 1787 + 1
Colubridae
Ahaetulla mycterizans (Linnaeus, 1758) + 1
Ahaetulla prasina (Boie, 1827) + + 1, 2
Boiga cynodon (Boie, 1827) + + 1, 2
Boiga dendrophila (Boie, 1827) + + 1, 2
Boiga drapiezii (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
Calamaria lumbricoidea 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
Chrysopelea 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
Dendrelaphis formosus (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
Gonyosoma oxycephalum (Boie, 1827) + + 1, 2Gaps in Java and Bali Amphibian and Reptile Atlas 1023
Appendix 2. cont.
Taxon Java Bali Source
Colubridae (cont.)
Gonyosoma oxycephalum (Boie, 1827) + + 1, 2
Liopeltis tricolor (Schlegel, 1837) + 1, 2
Lycodon 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
Ptyas carinata (Günther, 1858) + 1
Ptyas korros (Schlegel, 1837) + + 1, 2
Ptyas mucosa (Linnaeus, 1758) + + 1, 2
Sibynophis geminatus (Boie, 1826) + + 1, 2
Sibynophis melanocephalus (Gray, 1835) + 1, 2
Tetralepis fruhstorferi Boettger, 1892 + 1
Xenodermus javanicus Reinhardt, 1836 + 2
Cylindropiidae
Cylindrophis ruffus (Laurenti, 1768) + 1, 2
Cylindrophis subocularis Kieckbusch et al., 2016 + 3
Elapidae
Aipysurus eydouxii (Gray, 1849) + 1
Bungarus candidus (Linnaeus, 1758) + + 1, 2
Bungarus fasciatus (Schneider, 1801) + 1, 2
Bungarus flaviceps Reinhardt, 1843 + 3
Calliophis bivirgatus (Boie, 1827) + 1
Calliophis intestinalis (Laurenti, 1768) + 1, 2
Hydrophis atriceps Günther, 1864 + 1
Hydrophis brookii Günther, 1872 + 3
Hydrophis caerulescens (Shaw, 1802) + 1
Hydrophis curtus (Shaw, 1802) + 1
Hydrophis cyanocinctus (Daudin, 1803) + 1
Hydrophis elegans (Gray, 1842) + 1
Hydrophis fasciatus (Schneider, 1799) + 1
Hydrophis gracilis (Shaw, 1802) + 1
Hydrophis hardwickii (Gray, 1834) + 1
Hydrophis inornatus (Gray, 1849) + 3
Hydrophis melanocephalus Gray, 1849 + 1
Hydrophis ornatus (Gray, 1842) + 1
Hydrophis platurus (Linnaeus, 1766) + 1, 2
Hydrophis schistosus Daudin, 1803 + 1
Hydrophis spiralis (Shaw, 1802) + 1
Hydrophis stokesii (Gray, 1846) + 1
Hydrophis viperinus (Schmidt, 1852) + 3
Kolpophis annandalei (Laidlaw, 1901) + 3
Laticauda colubrina (Schneider, 1799) + + 1, 21024 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
Gerrhopilus ater (Schlegel, 1839) + + 1
Gerrhopilus bisubocularis (Boettger, 1893) + 3
Homalopsidae
Cerberus schneiderii (Schlegel, 1837) + + 1, 2
Enhydris enhydris (Schneider, 1799) + 1, 2
Fordonia leucobalia (Schlegel, 1837) + 1, 2
Homalopsis 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
Rhabdophis chrysargos (Schlegel, 1837) + + 1,2
Rhabdophis rhodomelas (Boie, 1827) + 3
Rhabdophis subminiatus (Schlegel, 1837) + 1,2
Xenochrophis trianguligerus (Boie, 1827) + 1,2
Xenochrophis vittatus (Linnaeus, 1758) + 1,2
Pareidae
Aplopeltura 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) + 3Gaps 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
Indotyphlops braminus (Daudin, 1803) + + 1, 2
Ramphotyphlops lineatus (Schlegel, 1839) + 1, 2
Viperidae
Calloselasma rhodostoma (Kuhl, 1824) + 1,2
Daboia siamensis (Smith, 1917) + 1
Trimeresurus albolabris Gray, 1842 + 1,2
Trimeresurus insularis Kramer, 1977 + 1,2
Trimeresurus puniceus (Boie, 1827) + 1,2
Xenopeltidae
Xenelaphis hexagonotus (Cantor, 1847) + 3
Xenopeltis 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
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