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ZOOTAXA
ISSN 1175-5326 (print edition)
ISSN 1175-5334 (online edition)
Accepted by A. Bauer: 26 Mar. 2019; published: 17 May 2019
Licensed under a Creative Commons Attribution License http://creativecommons.org/licenses/by/3.0
155
Zootaxa 4608 (1): 155–173
https://www.mapress.com/j/zt/
Copyright © 2019 Magnolia Press Article
https://doi.org/10.11646/zootaxa.4608.1.9
http://zoobank.org/urn:lsid:zoobank.org:pub:62E65783-1E5F-434E-B6CC-CBD611EF6C74
Hiding in plain sight on Gunung Muria: A new species and first record of rock
gecko (Cnemaspis Strauch, 1887; Squamata, Gekkonidae) from Java, Indonesia
AWAL RIYANTO1,3, MISBAHUL MUNIR1 , ANDRI I. S. MARTAMENGGALA2,
YULI SULISTYA FITRIANA1 & AMIR HAMIDY1
1 Museum Zoologicum Bogoriense, Research Center for Biology – Indonesian Institute of Sciences, Widyasatwaloka Building, Jl. Raya
Jakarta Bogor, Km.46, Cibinong, West Java, Indonesia
2 Gaia Consulting, CEO building, lv 12, Jl. TB Simatupang 18C, Jakarta, Indonesia
3 Corresponding author. E-mail: awal_lizards@yahoo.com
Abstract
We describe a new species of rock gecko of the genus Cnemaspis from Java, Indonesia, representing the first record of
the genus for this Island. The new species was collected from the southern slopes of Gunung Muria, a dormant volcano
in Central Java. The new species is easily distinguished from all congeners by having a maximum SVL of 58.1 mm in
males and 56.9 mm in females; a pair of sharp conical tubercle clusters on the occiput; a warty bridge on the nuchal loop,
extending from the upper tympanum and curving to the nape; dorsal tubercles not linearly arranged; 18–20 paravertebral
tubercles; postmentals separated by one scale; gular, pectoral and abdominal scales, ventral scales of fore- and hindlimbs,
and subcaudal scales keeled; no tubercles on lower flank; precloacal and femoral pores absent; enlarged submetacarpal
scales present on the first digit of the manus; 38–40 ventral scales; 31–35 lamellae under fourth toe; two postcloacal
tubercles on each side; enlarged median subcaudal scales row present; caudal tubercles encircling tail; and a sexually
dimorphic ventral color pattern, with males having a yellow belly and females white and the ventral surface of the
tail in males yellow proximally changing to white at mid-length, whereas in females, alternating black and white rings
completely encircle the tail, which is black distally.
Key words: Central Java, Cnemaspis, first record, Mount Muria, new species
Abstrak
Kami mempertelakan spesies baru cicak batu dari genus Cnemaspis dari Jawa, Indonesia, yang juga sebagai catatan
pertama genus di pulau ini. Spesies baru dikoleksi dari lereng selatan Gunung Muria, yang merupakan gunung api dorman
yang berada di Jawa Tengah. Spesies baru ini mudah dibedakan dari semua kerabatnya dengan memiliki SVL maksimum
58,1 mm pada jantan dan 56,9 mm pada betina; sepasang struktur tuberkular seperti kerucut pada kepala bagian belakang;
alur berkutil pada nuchal loop, membentang dari atas tympanum melengkung ke tengkuk; tuberkular dorsal tidak tersusun
secara linier; 18–20 baris tuberkular sepanjang paravertebral; postmental dipisahkan oleh satu sisisk; gular, pektoral,
abdominal, sisi ventral kaki depan dan belakang, serta sisik subcaudal berlunas; tidak ada tuberkul di lipatan paha; tidak
mempunyai pori-pori prakloakal maupun femoral; pada jari pertama tungkai depan terdapat struktur submetakarpal yang
membesar; pada jari pertama tungkai belakang terdapat sisik submetatarsal yang membesar; 38–40 sisik perut; 31–35
lamella di bawah jari keempat tungkai belakang; pada kanan kiri kloaka terdapat masing dua struktur tuberkular; terdapat
satu baris sisik subcaudal yang membesar ditengah ventral ekor; tuberkular membentuk formasi cincin di sepanjang ekor;
dan terdapat dimorfisme seksual warna pada ventral tubuh yaitu pada jantan perut dan pangkal ekor berwarna kuning
dengan setengah panjang ekor hingga ujung berwarna putih, sedangkan pada betina perut berwarna putih dan ekor dengan
pola cincin hitam putih yang berselang-seling dengan ujung berwarna hitam.
Kata kunci: Jawa Tengah, Cnemaspis, catatan pertama, Gunung Muria, spesies baru
Introduction
The African-Asian rock geckos of the genus Cnemaspis Strauch, 1887 comprise about 133 species (Uetz et al. 2018).
In Southeast Asia, with 50 species, this genus is distributed in central and southern Vietnam, marginal areas of the
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Gulf of Thailand, through the Malay Peninsula, Seribuat Island, the Riau Archipelago (Anambas, Natuna, Tembelan
and Belitung islands), Borneo (including the Karimata Islands), and Sumatra (including the Mentawai archipelago:
Simeleu, Nias, Siberut, Pagai and Enggano islands) (Gray, 1845; Smith, 1925; Das & Bauer, 1998; Das, 2005; Gris-
mer & Chan 2009; Grismer et al. 2014; Amarasinghe et al. 2015; Iskandar et al. 2017; Riyanto et al. 2017; Kurita et
al. 2017). In the Greater Sundas, in Sumatra and the Riau and Mentawai archipelagoes at least 13 species have been
encountered: Cnemaspis aceh Iskandar, Amarasinghe & McGuire, 2016; C. andalas Iskandar, Amarasinghe & Mc-
Guire, 2016; C. dezwaani Das, 2005; C. jacobsoni Das, 2005; C. minang Iskandar, Amarasinghe & McGuire, 2016;
C. modiglianii Das, 2005; C. mumpuniae Grismer, Wood, Anuar, Riyanto, Ahmad, Muin, Sumontha, Grismer, Chan,
Quah & Pauwels, 2014; C. pagai Iskandar, Amarasinghe & McGuire, 2016; C. purnamai Riyanto, Hamidy, Sidik &
Gunalen, 2017; C. rajabasa Amarasinghe, Harvey, Riyanto & Smith, 2015; C. sundainsula Grismer, Wood, Anuar,
Riyanto, Ahmad, Muin, Sumontha, Grismer, Chan, Quah & Pauwels, 2014; C. tapanuli Iskandar, Amarasinghe &
McGuire, 2016; and C. whittenorum Das, 2005; and five species are known to occur on Borneo: C. dringi Das &
Bauer, 1998; C. kendallii (Gray, 1845); C. leucura Kurita, Nishikawa, Matsui & Hikida, 2017; C. nigridia (Smith,
1925); and C. paripari Grismer & Chan, 2009. Until now there were no records of this genus from Java. Field sur-
veys in July and August 2018 revealed nine specimens of the genus Cnemaspis from Gunung Muria, Central Java,
Indonesia. These represent a population sufficiently different from all currently recognized species of Cnemaspis,
on the basis of color pattern, size, and scalation, to warrant description as a new species.
Material and methods
Specimen collection. A herpetofaunal field survey was conducted in early July 2018 by one of the authors (AISM)
in Gunung Muria, Kudus Regency, Central Java, during which seven specimens of rock geckos, genus Cnemaspis,
were obtained. Twenty-five days later, AR and MM, collected two more specimens at the same location. The speci-
mens were captured by hand, euthanized with 20% benzocaine diluted in 70% ethanol. Liver tissue was removed
for DNA extraction and stored in 95% ethanol. Voucher specimens were fixed with 10% buffered formalin, and later
stored in 70% ethanol. Latitude, longitude, and elevation of the collecting localities were recorded on the WGS 84
map datum system using Garmin GPSmap 60CSx. All type specimens were deposited at the Museum Zoologicum
Bogoriense (MZB), Indonesian Institute of Sciences (LIPI), Cibinong, Indonesia.
Morphological analysis. Most morphological characters, external measurements and scalation counts fol-
lowed Grismer et al. (2014). Measurements were taken using dial calipers (Mitutoyo) to the nearest 0.1 mm, under
a dissecting microscope (AmScope) on the right side. Measurements: snout–vent length (SVL, from tip of snout
to anterior margin of vent); tail length (TailL, from vent to tip of tail); head length (HeadL, from posterior edge of
retroarticular process of lower jaw to tip of snout); head width (HeadW, measured in a straight line at angle of jaws);
head depth (HeadD, maximum height of head from occiput to throat); snout length (SnoutL, from anterior most edge
of orbit to tip of snout); eye to ear distance (EED, from posterior edge of orbit to anterior edge of ear opening); ear
length (EarL, maximum length of ear opening); orbital diameter (OD, horizontal diameter of eye ball); axilla–groin
length (AGL, from posterior margin of forelimb, at insertion point on body, to anterior margin of hind limb, at in-
sertion point on body). We also measured some characters on the ventral surface: antebrachium length (AntBraL,
from posterior margin of elbow, while flexed, to inflection of flexed wrist); brachium length (BracL, from axilla to
inflection of flexed elbow); thigh length (ThighL, from anterior margin of hind limb, at insertion on body, to knee,
while flexed); and tibia length (TibiaL, from posterior surface of knee, while flexed 90º, to base of heel).
Meristic data collected were: supralabial (SupL) and infralabial (InfL) scales (counted from below the middle
of the orbit to the rostral and mental scales, respectively); number of paravertebral tubercles (PVT, counted between
limb insertions, in a straight line immediately right of the vertebral column); and number of subdigital lamellae
under fourth toe (counted from base of first phalanx to distalmost lamella, excluding claw sheath). We noted the
texture of scales (smooth, unicarinate or tricarinate) on the gular, pectoral, and abdominal regions on the ventral side
of the fore- and hindlimbs and on subcaudal surfaces. The presence of tubercles along the ventral edge of the body
(flank) between limb insertions, and in whorls encircling the tail was noted as were enlarged submetacarpals and
submetatarsals on the first finger and toe, respectively. The presence of enlarged unpaired median subcaudals was
also noted. Following Harvey et al. (2015) we applied methylene blue in 70% ethanol to stain specimens and better
visualize some minute structures, such as subdigital keels and pores.
ROCK GECKO FROM JAVA, INDONESIA Zootaxa 4608 (1) © 2019 Magnolia Press · 157
Color notes were taken based on digital images obtained from living specimens prior to preservation.
We also obtained character and distribution information for other Southeast Asian Cnemaspis species from Gris-
mer et al. (2014), Amarasinghe et al. (2015), Iskandar et al. (2017), Kurita et al. (2017) and Riyanto et al. (2017).
FIGURE 1. Bayesian Inference tree based upon ~954 bp NADH dehydrogenase 2 gene sequences for representative Cne-
maspis species. Values above or below branches indicate Bayesian Posterior Probability (BPP) and Maximum Likelihood Boot-
srap Proportion (MLBP), asteric indicates strongly supported clade (BPP=100 and MLBP=100).
Phylogenetic analysis. We extracted genomic DNA from liver stored in 95% using standard phenol-chlo-
roform method (Sambrook et al. 1989). DNA amplification of the NADH dehydrogenase 2 (ND2) and partial
flaking tRNAs were carried out in Polymerase Chain Reactions (PCR) using the primers M112F (5-AAGCTTTC-
GGGGCCCATACC-3) and M1123R (5-GCTTAATTAAAGTGTYTGAGTTGC-3) designed in the flanking me-
thionine and alanine tRNAs (Oliver et al. 20016). The PCRs were performed in 25 µl total volumes using Top TaqTM
by Qiagen comprising 1.0 µl DNA template, 2.5 µl 10x Top Taq PCR bufferTM, 0.5 µl 10 mM dNTP mix, 2.5 µl 10x
CoralLoad, 5 µl 5x Q solution, 1.0 µl light strand primer, 1.0 µl heavy strand primer, 0.125 µl Top Taq DNA poly-
merase with appropriate buffer and ddH2O to volume. PCR reactions were executed on an Eppendorf Mastercycler
under following conditions: initial denaturation at 94°C for 9 min, followed by a second denaturation at 94°C for 45
s, anneling at 60°C for 45 s, followed by a cycle extension at 72°C for 1 min, for 35 cycles. The purified PCR prod-
ucts were sequenced directly by 1st Base Asia facility (Singapore). Sequence data were validated using Chromas pro
software (Technelysium Pty Ltd, Australia) and aligned with published data from GenBank (Table 1) using Clustal
W in MEGA 6.06 (Tamura et al. 2013). Phylogenetic analyses of 954 base pairs (bp) of ND2 and its flankig tRNAs
were performed using Maximum Likelihood (ML) and Bayesian Inference (BI). We used the Akaike criterion as
implemented in Kakusan 3 (Tanabe 2007) to select the best model of evolution for both BI and ML analyses. We
conducted our BI analysis using MrBayess 3.2.6 (Ronquist & Huelsenbeck 2003) under GTR+Gamma as the best-
fitting model, for 10 million generations with parameter and topology sampling every 1000 generations and Markov
Chain Monte Carlo (MCMC) diagnosis frequency of 100,000 and later discarded 25% of the first analysis as burn-
in. For the likelihood analysis, we used RAxML version 8.2.10 (Stamatakis 2014) implementing the GTR+Gamma
model with 1000 bootstrap replicates on the CIPRES Science Gateway platform (Miller et al. 2010). Genetic dis-
tances (uncorected p-distance) were estimated using MEGA 6.06 (Tamura et al. 2013). The two new sequences of
Cnemaspis from Java were submitted to GenBank under the accession numbers MK161507 and MK161508.
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158 · Zootaxa 4608 (1) © 2019 Magnolia Press
TABLE 1. A list of the samples used in the molecular analyses with GenBank accession numbers. Abbreviated for voucher
numbers are as follows: LSUHC, La Sierra University Herpetological Collection; AA, Rohan Pethiyagoda field series;
AMB, Aaron M. Bauer field series; KUHE, Graduate School of Human and Environmental Studies, Kyoto University;
KUZ, Zoological collection of Kyoto; MVZ, Museum of Vertebrate Zoology University of California, Berkeley; MZB,
Museum Zoologicum Bogoriense, Indonesia; and WHT, Wildlife Heritage Trust, Sri Lanka.
Species Vouchers Locality ND2 GenBank
Accession
C. baueri LSUHC 7274 Malaysia: Johor, Pulau Aur KM024699
C. baueri LSUHC 7303 Malaysia: Johor, Pulau Aur KM024697
C. bidongensis LSUHC 11444 Malaysia: Trenggani, Pulau Bidong KM024703
C. bidongensis LSUHC 11446 Malaysia: Trengganu, Pulau Bidong KM024705
C. boulengeri LSUHC 9278 Vietnam: Ca Mau Province, Con Dao Archipelago KM024710
C. dringi KUHE 56015 Malaysia: Serawak, Gunung Mulu LC158337
C. grismeri LSUHC 9970 Malaysia: Perak, Lenggong KM024723
C. kallima AA 82 Sri Lanka: Matale District, Rattota, Gammaduwa KY037970
C. kandiana AMB 7487 Sri Lanka, Kandy District, Gampola KY037972
C. kendallii LSUHC 5317 Indonesia: Riau Province, Pulau Serasan KM024738
C. kendallii LSUHC 9174 Malaysia: Sarawak, Gunung Gading KM024742
C. kendallii LSUHC 9179 Malaysia: Sarawak, Santubong KM024744
C. kendallii KUHE 57123 Malaysia: Serawak, Kuching LC205718
C. kendallii KUHE 57350 Malaysia: Serawak, Kuching LC205720
C. kendallii KUHE 57343 Malaysia: Serawak, Kuching LC205719
C. kendallii KUZ(JPN): R27060 Malaysia: Serawak, Kuching LC158342
C. leucura KUHE 57422 Malaysia: Serawak, Kuching LC205724
C. leucura KUHE 57423 Malaysia: Serawak, Kuching LC205723
C. limi LSUHC 3774 Malaysia: Pahang, Pulau Tioman KM024749
C. limi LSUHC 3888 Malaysia: Pahang, Pulau Tioman KM024750
C. hangus HC 0225 Malaysia: Pahang, Bukit Hangus KM024729
C. mcguierei LSUHC 8855 Malaysia: Perak, Bukit Larut KM024753
C. mumpuniae MZB Lace 10167 Indonesia: Pulau Natuna KM024761
C. modiglianii MVZ 239314 Indonesia: Pulau Enggano KY037977
C. modiglianii MVZ 239315 Indonesia: Pulau Enggano KY037978
C. muria sp.nov. MZB Lace 14571 Indonesia: Java, Gunung Muria MK161507
C. muria sp.nov. MZB Lace 14572 Indonesia: Java, Gunung Muria MK161508
C. nigridia KUZ(JPN): R27145 Malaysia: Serawak, Kuching, Lundu, Gunung
Gading
LC158342
C. nigridia LSUHC 9170 Malaysia: Serawak, Gunung Gading KM024772
C. paripari KUHE 57285 Malaysia: Serawak, Kuching LC205725
C. paripari LSUHC 9184 Malaysia: Serawak, Gua Paripari KM024781
C. pava AMB7494 Sri Lanka, Nuwara Eliya District, Labookellie KY037980
C. peninsularis LSUHC 5184 Malaysia: Johor, Pulau Seribuat KM024796
C. peninsularis LSUHC 5185 Malaysia: Johor, Pulau Seribuat LSUHC 5185
C. peninsularis LSUHC 53017 Malaysia: Johor, Pulau Aceh
C. peninsularis LSUHC 3773 Malaysia: Pahang, Pulau Tioman, Malaysia KM024789
C. peninsularis LSUHC 6381 Malaysia: Johor, Pulau Ibol KM024801
C. pemanggilensis LSUHC 80312 Malaysia: Johor, Pulau Pemanggil KM024786
C. pemanggilensis LSUHC 80313 Malaysia: Johor, Pulau Pemanggil KM024787
C. pseudomcguirei LSUHC 9146 Malaysia: Perak, Bukit Larut KM024825
C. samanalensis AMB7505 Sri Lanka: Nuwara Eliya District KY037983
C. sundainsula MZB Lace 10156 Indonesia: Riau Province, Pulau Natuna Besar,
Ceruk Forest
KM024845
C. sundainsula MZB Lace 10157 Indonesia: Riau Province, Pulau Natuna Besar,
Ceruk Forest
KM024846
C. temiah LSUHC 9160 Malaysia: Pahang, Cameron Highlands, Tanah
Rata
KM024849
C. upendrai WHT 7258 Sri Lanka, Nuwara Eliya District, Punduloya KY037985
ROCK GECKO FROM JAVA, INDONESIA Zootaxa 4608 (1) © 2019 Magnolia Press · 159
Results
Both ML and BI analysis yielded nearly the same phylogenetic tree topology, with only minor differences in the
placement of species with low bootstrap values (Fig. 1). Peninsular Malaya, Bornean and Javan Cnemaspis formed
a monophyletic group with strong support only from Bayesian posterior probabilities (BPP=100) but was not sup-
ported under Maximum Likelihood bootstraps. The new species from Java was placed with low support in the C.
kendalii group but was strongly supported (1.00/100) as the sister taxon of C. bidongensis Grismer, Wood, Ahmad,
Sumarli, Vazquez, Ismail, Nance, Mohd-amin, Othman, Rizaijessika, Kuss, Murdoch & Cobos, 2014 from Pulau
Bidong in Peninsular Malaysia. The uncorrected p-distance between the new species and their congeners from Pen-
insular Malaysia, Borneo and their surrounding islands is large (range = 4.4–34.2 and while interspecies distances
among C. kendalii group are quite high (range = 4.4–18.2%) (Table 2). The diagnostic characters revealed in the
morphological analysis (Tables 4, 5) supported by the molecular results, support the distinctiveness of the Javan
Cnemaspis, which is described below as a new species.
Systematics
Cnemaspis muria sp.nov.
Muria Rock Gecko
Cicak Batu Gunung Muria
Figures 2–5
urn:lsid:zoobank.org:act:066FE9E7-7916-4E8E-AE81-CEFCCB8901C8
Holotype. MZB. Lace. 14571 (Fig. 2A), an adult male from the river bank at Gunung Muria, Kajar (village), Dawe
(District), Kudus (Regency), Jawa Tengah (Province), Indonesia (06o39’47.4” S; 110o53’22.9” E; elevation 599 m
asl), collected on 11 August 2018 by Awal Riyanto, Misbahul Munir, Rubby Alfian, Lianita Rarasandy and Rega
D. Ganiarta.
Paratypes. MZB. Lace. 14564 (Fig. 2B), an adult female; MZB. Lace. 14565–70, six adult males from the
same locality as holotype (06˚39’33.5” S; 110˚53’20.6” E; elevation 650 m asl), collected on 17 July 2018 by Andri
I.S. Martamenggala, and MZB. Lace.14572 (Fig. 2C), an adult male, with the same data as holotype (6°39’36.9” S;
110o53’20.0” E; elevation 646 m asl).
Diagnosis. Cnemaspis muria sp. nov. differs from its congeners in Southeast Asian by the following combina-
tion characters: (1) maximum SVL of at least 58.1 mm in males and 56.9 mm in females, (2) a pair of sharpe conical
tubercle clusters on occiput, (3) nuchal loop bearing a bridge of warts from the upper tympanum to the nape and
made in a curved line, (4) dorsal tubercles not linearly arranged, (5) 18–20 paravertebral tubercles, (6) postmental
separated by a single scale, (7) gular, pectoral, abdominal, subantebrachial, subbrachial, subfemoral, subtibial and
subcaudal scales keeled, (8) no tubercles on lower flank, (9) both precloacal and femoral pores absent, (10) enlarged
submetacarpal scales present at the base of first finger, (11) enlarged submetatarsal present at the base of first toe ,
(12) 38–40 ventral scales, (13) 31–35 lamellae under fourth toe, (14) two postcloacal tubercles on each side, (15)
enlarged median subcaudal scale row present, (16) caudal tubercles encircling tail, (17) sexually dimorphic in color
pattern: males with a yellow belly and the proximal subcaudal surfaces yellow becoming white distally, female with
a white belly with proximal subcaudal surface of alternating white and black rings and black coloration distally.
Description of holotype. An adult male, 56.8 mm SVL; head oblong in dorsal profile, moderately large in size
(HeadL/SVL 0.28), elongate (HeadW/HeadL 0.65), narrow and flattened (HeadD/HeadL 0.36), distinct from neck;
snout short (SnoutL/HeadL 0.43), slightly concave in lateral profile; snout longer than the distance between eye and
ear (SnouL/EarEye 1.67); canthus rostralis smoothly rounded; eye large (ED/HeadL 0.21; ED/SnoutL 0.50); pupil
round; ear opening oval, taller than wide; rostral concave, dorsal 80% divided by longitudinal groove; rostral width 1.66
times of it’s length; rostral bordered posteriorly by two small supranasals and two large scales between the supranasals
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TABLE 2. Uncorected p-distance (percent difference) based on 954 bp NADH dehydrogenase 2 gene sequences for representative Cnemaspis kendalii group. Ranges distance in parentheses.
No
Species
1
2
3
4
5
6
7
8
1
Cnemaspis muria sp. nov.
0.0
2
Cnemaspis bidongensis
4.4–4.5
0.1
3
Cnemaspis mumpuniae
6.7
6.1–6.2
–
4
Cnemaspis peninsularis
6.4–8.3
6.4–7.9
6.7–8.6
0.0–7.1
5
Cnemaspis baueri
14.8–14.9
15.0–15.2
14.5–14.6
14.3–16.0
0.2
6
Cnemaspis pemanggilensis
13.5–13.7
13.5–13.7
14.7–14.9
14.9–15.8
16.1–16.5
0.6
7
Cnemaspis kendalii
17.4–17.7
17.3–17.6
18.0–18.4
17.5–18.8
18.2–19.0
18.7–19.6
0.2–1.8
8
Cnemaspis sundainsula
18.1–18.2
16.7–16.9
17.4–17.5
17.5–18.7
19.5–19.7
18.3–18.6
17.3–18.0
0.1
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FIGURE 2. Types specimens of Cnemaspis muria sp. nov. A. Holotype (MZB.Lace.14571), male. B. Paratype (MZB.
Lace.14564), female. C. Paratype (MZB.Lace.14572), male. Photos A and C by A. Riyanto, B by A. I. S. Martamenggala.
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162 · Zootaxa 4608 (1) © 2019 Magnolia Press
FIGURE 3. Close up of the head of Cnemaspis muria sp. nov. showing a pair of sharpe conical tubercles clustered on the
occiput (yellow arrows) and nuchal loop bearing a bridge of warts in a curved line from the upper tympanum to the nape (red
arrows). Photo by A. Riyanto.
and laterally by first supralabials; 10 supralabials on each side; 9 infralabials on each side, decreasing in size slightly
posteriorly; nostrils elliptical, oriented posterodorsally, bordered posteriorly by small, granular, postnasal scales; a
pair of bridges of warts present on occiput; nuchal loop present, bearing a bridge of warts extending from upper
tympanum to the nape in a curved formation (Fig. 3).
Mental large, sub triangular, elongate (MentL/MentW 1.11), and extending posteriorly to a point equal to the
anterior part of third infralabial, laterally in contact with first infralabial, posterolaterally bordered by three post-
mental scales; asymmetric arrangement of the postmentals, slightly damaged on the right side of the postmental,
postmentals bordered posteriorly by 11 weakly keeled scales; scales on throat raised and weakly keeled.
Body slender, elongate (AGL/SVL 0.41); dorsal body covered by pointed-weakly unicarinate scales which
homogeneous in size; dorsal tubercles moderately prominent and randomly distributed; absent tubercles on flank;
dorsal scales at mid-body smaller than ventral at same level; 20 PVT, flat, sub-pyramidal and weakly tricarinate,
each about two or three times as large as granules separating them; abdominal and ventral scales sub-equal in size,
ovoid, tricarinate, juxtaposed; pre-cloacal and femoral pores absent; enlarged femoral scales absent.
Forelimbs short (AntBraL/SVL 0.18; BracL/SVL 0.17); dorsal scales on both upper and lower arm, same size as
abdominal scales; ventral scales on lower arm slightly larger than ventral scales on upper arm, unicarinate; ventral
scales on lower arm weakly tricarinate; digits well developed, elongate, slender, all bearing slightly recurved claws;
enlarged sub-metacarpal present at base of first finger (Fig. 4A); relative length of fingers 4˃3˃5˃2˃1.
Hindlimbs relatively long (TibiaL/SVL 0.26; ThighL/SVL 0.23); dorsal scales on both tibias and thighs weakly
tricarinate, size relatively homogeneous; subtibial sligthly smaller than subfemoral scales, enlarged scales absent,
both tricarinate; digits well developed, elongate, slender, all bearing slightly recurved claws; enlarged submetatarsal
present at the base of first toe (Fig. 4B); relative length of toes 4˃3˃5˃2˃1; 31 lamellae under fourth toe.
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FIGURE 4. Palmar and tarsal views of the holotype Cnemaspis muria sp. nov. (MZB.Lace.14571), showing the enlarged meta-
carpal scale on the first finger (A) and the enlarged metatarsal scale on the first toe (B). Bar = 10 mm. Photos by A. Riyanto.
FIGURE 5. Ventral tail view of Cnemaspis muria sp. nov., holotype (MZB.Lace.14571) showing the unicarinate subcaudal
scales, and the interrupted median row of enlarged unicarinate subcaudals. Bar = 10 mm. Photos by A. Riyanto.
Tail original (TailL 73.5 mm), long (TailL/SVL 1.29), base swollen; two conical postcloacal tubercles present
on each side; a distinct furrow on the lateral surface of the tail present; caudal tubercles encircling tail in whorls; tail
segmented with 21 whorls of tubercles, each whorl consisting of ten enlarged keeled tubercles separated from one
another by 1 to 8 small scales; each whorl separated from adjacent whorls by 5 to 7 small scales; subcaudal scales
unicarinate; interrupted median row of enlarged unicarinate subcaudals (Fig. 5).
Color pattern in life. Dorsal ground color brick-red; rostrum mixed yellowish; supralabials, infralabials and
superciliaries yellow; a yellowish white continuous nuchal loop present, extending from middle posterior margin of
one orbit to the other; a yellow line present, extending from lower posterior margin of orbits to the lower anterior
margin of ears; white ocelli in shoulder region; three black weakly longitudinal spots arranged in a transverse row
on the nape and shoulder region; seven blackish blotches intermixed with six white blotches along spine extending
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164 · Zootaxa 4608 (1) © 2019 Magnolia Press
from shoulder to hindlimb insertion; six white dorsolateral blotches between limb insertions; flanks bearing small,
round, yellow spots that extend onto along lateral margins of abdomen; dorsal ground color of tail dorsum is brick-
red, interrupted by blackish and white transverse bands and the apex of the tail white.
In mental region ventral part of infralabials yellow; gular region yellowish white; pectoral and abdominal re-
gions yellow; subantebrachials, subbrachials, precloacal region, subtibials and subfemorals yellowish; palms and
tarsals yellow; proximal subcaudals yellow until about the ninth whorl, then white distally.
Variation. Meristic data for the single female are similar to those of males and measural data are included within the
range of males (see Table 3), which led us to consider them as conspecific, despite the differences in coloration. The new
species exhibits sexual dimorphism in ventral color pattern. Males have a yellow belly whereas it is white in the female and
the ventral surface of tail in males is proximally yellow and distally white (changing at approximately mid-length) but in
the female alternating black and white rings completely encircle the tail, which is black distally (Fig. 6).
FIGURE 6. Sexual dimorphism exhibited in ventral views of Cnemaspis muria sp. nov. A. Male, holotype (MZB.Lace.14571),
showing the subcaudal coloration that changes from yellow to white at about the midpoint of the tail. B. Female, paratype (MZB.
Lace.14564), showing the subcaudal surface with alternating white and black bands and a black apex of the tail. Bar = 10 mm.
Photo A by A. Riyanto, B by A. I. S. Martamenggala.
Etymology. The specific epithet muria is a noun in apposition and refers to Gunung Muria, the type locality of
this species, and so far, the only known locality for the genus Cnemaspis in Java.
Distribution. Cnemaspis muria sp. nov. is only known from its type locality, in the southern foothills of Gu-
nung Muria, a dormant volcano 1602 m in maximum elevation, located in the center of the Muria Peninsula of
northern Central Java (Jawa Tengah). Gunung Muria represents the first, and so far, the only known locality for the
genus Cnemaspis in Java (Fig. 7).
Natural history. Cnemaspis muria sp. nov. is a scansorial species known only from large granite rock micro-
habitats along rivers and coffee plantations (Fig. 8) on the southern slope of Gunung Muria at middle elevations,
between 560 and 599 m. The holotype was caught at night, hanging on a tree root, 2 m above a dry river bank. The
paratypes MZB. Lace. 14564 and MZB. Lace. 14572 were also caught at night, whereas the other paratypes were
caught during the day hanging on shaded crevices of rocks. MZB. Lace. 14572 and four another specimens not col-
lected were found foraging on rock walls, and sympatric with Cyrtodactylus sp.
ROCK GECKO FROM JAVA, INDONESIA Zootaxa 4608 (1) © 2019 Magnolia Press · 165
TABLE 3. Meristic and mensural character states of the type series of Cnemaspis muria sp. nov. Meristic abbreviations are listed in the Materials and Methods. For SupL and InfL, the number
in the parentheses is left side. TailL, the measurement for the original portion of the tail is in the parenthesis and the measurement for the regenerated portion is on the right. All measurements
are in mm.
MZB.Lace,
14571
holotype
14564
14565
14566
14567
14568
14569
14570
14572
Field number
MUN263
MVR006
MVR007
MVR008
MVR009
MVR010
MVR011
MVR012
MUN364
Date collection
08/11/2018
07/17/2018
07/17/2018
07/17/2018
07/17/2018
07/17/2018
07/17/2018
07/17/2018
08/11/2018
Sex
Male
Female
Male
Male
Male
Male
Male
Male
Male
SupL
10 (10)
10 (11)
9 (10)
12 (13)
11 (12)
11 (11)
11 (12)
9 (10)
9 (9)
InfL
9 (9)
8 (9)
9 (8)
10 (10)
10 (10)
8 (8)
10 (8)
8 (10)
7 (7)
PVT
20
20
20
18
20
20
20
20
20
LamT4
31
32
33
32
30
31
30
32
31
No. of postcloacal
tubercles
2
2
2
2
2
2
2
2
2
Enlarged submetacarpal
scale on 1st finger absent
(0) or present (1)
1
1
1
1
1
1
1
1
1
Enlarged submetatarsal
scale on 1st toe absent (0)
or present (1)
1
1
1
1
1
1
1
1
1
Tubercles absent (0) or
present (1) on lower flanks
0
0
0
0
0
0
0
0
0
Gular scales smooth (0) or
keeled (1)
1
1
1
1
1
1
1
1
1
Pectoral scales smooth (0)
or keeled (1)
1
1
1
1
1
1
1
1
1
Abdominal scales smooth
(0) or keeled (1)
1
1
1
1
1
1
1
1
1
Subcaudal scales smooth
(0) or keeled (1)
1
1
1
1
1
1
1
1
1
Enlarged median
subcaudals absent (0) or
present (1)
1
1
1
1
1
1
1
1
1
Single median row of
enlarged subcaudals
smooth (0) or keeled (1)
1
1
1
1
1
1
1
1
1
Tubercles encircling tail
(1) or not (0)
1
1
1
1
1
1
1
1
1
SVL
56.8
56.9
57.3
57.4
58.0
58.6
58.5
58.1
56.4
TailL
73.5
54.1
69.1
76.2
66.9 (40.8)
72.8
68.3
59.9 (13.5)
73.0
AGL/SVL
0.41
0.43
0.41
0.44
0.45
0.43
0.43
0.43
0.43
HeadL/SVL
28.09
28.55
25.92
27.09
26.32
27.58
27.55
27.13
27.18
HeadW/HeadL
0.36
0.40
0.49
0.41
0.44
0.41
0.41
0.42
0.39
SnoutL/HeadL
0.43
0.41
0.45
0.41
0.43
0.44
0.41
0.44
0.42
SnoutL/EyeEar
1.67
1.48
1.52
1.49
1.29
1.67
1.54
1.42
1.52
SnoutL/OrbD
2.01
1.88
1.67
1.68
1.79
2.05
1.96
1.74
2.14
MentL/MentW
1.11
1.11
1.19
1.22
0.82
1.18
1.09
1.03
1.31
BracL/SVL
0.17
0.13
0.13
0.15
0.15
0.12
0.10
0.14
0.18
AntBraL/SVL
0.18
0.20
0.20
0.20
0.19
0.19
0.19
0.19
0.21
TibiaL/SVL
0.24
0.25
0.25
0.25
0.26
0.24
0.26
0.25
0.26
ThighL/SVL
0.23
0.22
0.22
0.19
0.21
0.20
0.20
0.28
0.23
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166 · Zootaxa 4608 (1) © 2019 Magnolia Press
FIGURE 7. Map of Java, showing the position of Gunung Muria as the type locality of Cnemaspis muria sp. nov. (red circle).
Base map modified from Wikipedia.
Comparison. Cnemaspis muria sp. nov. has numerous diagnostic characters states that separate it from con-
geners from the Malay Peninsula, adjacent small islands, Borneo, and Sumatra. Summary comparisons of the new
species with other species of the Southern Sunda clade of the C. kendalli group are presented in Tables 4 and Table
5. The new species is easily distinguished from its sister taxon, C. bidongensis, by the presence of single median row
of keeled subcaudals (versus smooth), an enlarged submetacarpal scales at the base of first finger (versus absence)
and an enlarged submetatarsal scale at the base of first toe (versus absence); from C. baueri it is distinguished by
the presence of an enlarged submetatarsal on the first toe (versus absence), absence of precloacal pores (versus pres-
ence), and presence of enlarged median row of subcaudals (versus absent); from C. kendallii it may be differentiated
by the absence of tubercles on lower flank (versus presence), presence of enlarged submetatarsal scales on the first
toe (versus absence), dorsal tubercles not linearly arranged (versus linearly arranged), ventral scales tricarinate (ver-
sus unicarinate) and single median row of keeled subcaudals (versus subcaudals not keeled); from C. mumpuniae by
the absence (versus presence) of tubercles on the lower flank and presence (versus absence) of enlarged submetatar-
sal scales on the first toe; from C. pemanggilensis Grismer & Das, 2005, by a lower number of PVT (18–20 versus
30–37), presence of enlarged submetatarsal on the first toe (versus absence); from C. peninsularis Grismer, Wood,
Anuar, Riyanto, Ahmad, Muin, Sumontha, Grismer, Chan, Quah & Pauwels, 2014 by the absence of tubercles on
lower flank (versus presence), presence of single median row of keeled subcaudals (versus absence) and enlarged
submetatarsal on the first toe (versus absence); and from C. sundainsula by absence of tubercles on lower flank
(versus presence), tubercles arranged not linearly (versus linearly arranged), lower number of PVT (18–20 versus
26–37), presence of keeled subcaudal scales (versus smooth subcaudal scales), keeled enlarged median subcaudal
row (versus smooth), and caudal tubercles encircling tail (versus not encircling).
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TABLE 4. Diagnostic characters differentiating Cnemaspis muria sp.nov. from congeners of the C. kendallii complex (sensu Grismer et al. 2014) and C. purnamai. Abbreviations: w = weak;
post = posterior.1 = maximum known SVL in mm, 2 = suplalabials, 3 = infralabials, 4 = number of paravertebral tubercles, 5 = dorsal tubercles linearly arranged (l) or random ( 0 ), 6 =
tubercles on lower flanks absent (0) or present (1), 7 = subcaudal scales smooth (0) or keeled (1), 8 = single median row of subcaudals smooth (0) or keeled (1), 9 = caudal tubercles encircling
tail (1) or not (0), 10 = enlarged median subcaudal row absent (0) or present (1), 11 = number of postcloacal tubercles (spur) in males, 12 = enlarged submetatarsal scale on 1st toe (1) or not, 13
= number lamella under 4th toes, References: A-this study, B–Amarasinghe et al. (2015), C–Grismer et al. (2014), and D–Riyanto et al. (2017).
Species
Characters
1
2
3
4
5
6
7
8
9
10
11
12
13
References
C. muria sp.nov.
58.6
9–13
7–10
18–20
0
0
1
1
1
1
2
1
31–35
A
C. baueri
67.4
11–13
8–12
18–27
0
w, 0
1
1
1
1
2
0
26–32
C
C. bidongensis
58.1
9, 10
7–9
21–26
0
0
1
0
1
1
1, 2
0
26–30
C
C. kendalli
58.4
10, 11
8, 9
18–26
w
w, 1
1
0
1
0
2
0
25–33
C
C. mumpuniae
60.9
9–12
8–11
18–25
w, 0
w, 1
1
0
1
1
1, 2
0
29–35
A, C
C. pemanggilensis
76.0
10–13
8–10
30–37
0
0, w
1
1
1
1
1, 2
0
27–34
C
C. peninsularis
60
10, 11
7–11
17–25
0
w, 1
1
0
1
0
1, 2
0
27–33
C
C. purnamai
54.1
9
8
14–15
0
0
1
1
1
1
2
1
22–24
A, D
C. rajabasa
58.7
13, 14
11, 12
20–21
0
0
1
1
1
0
2
0
28–34
B
C. sundagekko
68
11–13
8–11
20–25
0, w
0, w
1
0
1
0, post
2, 3
0
33–38
C
C. sundainsula
84.5
8–11
7–9
26–37
1
1
0
0
0
0
2–4
1
25–29
A, C
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TABLE 5.—Comparison on color pattern of Cnemaspis muria sp.nov. and congeners of the C. kendallii complex (sensu Grismer et al. 2014) and C. purnamai. ? = data unavailable, and ♂ =
characters present in males only.
muria sp.nov.
baurei
bidongensis
kendallii
mumpuniae
pemanggilensis
peninsularis
purnamai
rajabasa
sundagekko
sundainsula
Dorsal color pattern sexually dimorphic
no
no
no
no
no
no
yes
no
no
no
no
Ventral pattern sexually dimorphic
yes
no
no
no
no
no
yes
?
no
no
yes
Thin, white, nuchal loop
yes
no
no
no
yes
no
no
yes
no
no
no
Black round spots on nape and anterior of
body
no
yes
no
no
no
no
no
no
no
no
no
Dorsal ground color reddish
yes
no
no
no
yes
no
no
yes
no
no
no
Small, light, round spots on flanks
yes
no
no
no
yes
yes
no
yes
no
no
yes
Regenerated tail yellow
no
no
no
no
yes
no
yes
yes
no
no
no
Posterior portion of original tail black
yes
yes
no
no
yes
no
yes
?
no
no
no
Caudal bands present
yes
no
no
no
no
no
no
yes
no
no
no
Gular region orange
no
no
no
no
no
no
no
?
no
no
no
Gular region yellow
yes
no
no
no
no
no
no
?
no
no
yes
Throat orange
no
no
no
no
no
no
no
?
?
no
no
Throat yellow
yes
no
no
no
no
no
no
?
?
no
yes
Pectoral region orange
no
no
no
no
no
no
no
?
?
no
no
Pectoral region yellow
yes (♂)
no
no
no
no
no
no
?
?
no
no
Abdominen orange
no
no
no
no
no
no
no
?
?
no
no
Abdomen yellow
yes (♂)
no
no
no
no
no
no
?
?
no
yes
Ventral surface of forelimbs orange
no
no
no
no
no
no
no
?
?
no
no
Ventral surface of forelimbs yellow
yes (♂)
no
no
no
no
no
no
?
?
no
no
Ventral surface of hindlimbs orange
no
no
no
no
no
no
no
?
?
no
no
Ventral surface of hindlimbs yellow
yes (♂)
no
no
no
no
no
no
?
?
no
no
Subcaudal orange
no
no
no
no
no
no
no
?
?
no
no
Subcaudal yellow
yes (♂)
no
no
no
no
no
no
?
?
no
yes
At least posterior half of subcaudal region
white
yes (♂)
no
no
yes
no
no
no
?
?
no
yes
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FIGURE 8. The habitat type of Cnemaspis muria sp. nov. in Kajar village, Dawe District, Kudus Regency in Gunung Muria.
(A) Large rocks in a small river and (B) Large rock in a coffee plantation. Photos by A. I. S. Martamenggala.
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170 · Zootaxa 4608 (1) © 2019 Magnolia Press
Cnemaspis muria sp. nov. may be easily distinguished from C. purnamai Riyanto, Hamidy, Sidik & Gunalen,
2017, newly described from Belitung Island, by the presence of the nuchal loop bearing a bridge of warts extend-
ing from upper tympanum to the nape in a curved line (versus this configuration lacking) and a greater number of
lamellae under the fourth toe (31–35 versus 22–24); and from C. rajabasa by the presence of an enlarged median
subcaudal row (versus median row not enlarged) and enlarged submetatarsal under first toe (versus lacks enlarged
submetatarsal under first toe); and from C. sundagekko Grismer, Wood, Anuar, Riyanto, Ahmad, Muin, Sumontha,
Grismer, Chan, Quah & Pauwels, 2014, by the presence of single median row of enlarged keeled subcaudals (versus
median row not enlarged), and an enlarged submetatarsal scale on the first toe (versus absent).
The ventral scales in the new species are keeled, whereas the following species have smooth ventral scales:
Cnemaspis andalas, C. biocellata Grismer, Chan, Nurolhuda & Sumontha, 2008, C. flavigaster Chan & Grismer,
2008, C. kumpoli Taylor, 1963, C. monachorum Grismer, Ahmad, Chan, Belabut, Muin, Wood & Ahmad, 2009, C.
minang, and C. tapanuli.
The lack of precloacal pores in the new species distinguish it from all the following species which have precloa-
cal pores: C. andalas, C. affinis (Stoliczka, 1870), C. argus Dring, 1979, C. bayuensis Grismer, Grismer, Wood &
Chan, 2008, C. bidongensis, C. biocellata, C. hangus Grismer, Wood, Anuar, Riyanto, Ahmad, Muin, Sumontha,
Grismer, Chan, Quah & Pauwels, 2014, C. flavigaster, C. dezwani, C. dringi, C. flavolineata (Nicholls, 1949), C.
gismeri Wood, Quah, Anuar & Muin, 2013, C. harimau Chan, Grismer, Shahrul, Quah, Muin, Savage, Grismer, Ah-
mad, Remegio & Greer, 2010, C. karsticola Grismer, Grismer, Wood & Chan, 2008, C. kumpoli, C. leucura, C. limi
Das & Grismer, 2003, C. minang, C. mcguirei Grismer, Grismer, Wood & Chan, 2008, C. modiglianii, C. monocho-
rum, C. narathiwatensis Grismer, Sumontha, Cota, Grismer, Wood, Pauwels & Kunya, 2010, C. nigrida, C. pagai,
C. paripari C. perhentianensis Grismer & Chan, 2008, C. pseudomcguirei Grismer, Ahmad, Chan, Belabut, Muin,
Wood & Ahmad, 2009, C. selamatkanmerapoh Grismer, Wood, Mohamed, Chan, Heinz, Sumarli, Chan & Loredo,
2013, C. stongensis Grismer, Wood, Anuar, Riyanto, Ahmad, Muin, Sumontha, Grismer, Chan, Quah & Pauwels,
2014, C. tapanuli, C. temiah Grismer, Wood, Anuar, Riyanto, Ahmad, Muin, Sumontha, Grismer, Chan, Quah &
Pauwels, 2014 and C. whittenorum.
Cnemaspis muria sp. nov. possesses enlarged submetatarsal scales on the first toe which distinguishes it from
all the following species which lack this character: C. affinis, C. argus C. bayuensis, C. biocellata, C. flavolin-
eata, C. grismeri, C. hangus, C. harimau, C. karsticola, C. limi, C. mashuriae, C. mcguirei, C. narathiwatensis,
C. pemanggilensis, C. perhentianensis, C. selamatkanmerapoh, C. shahruli Grismer, Chan, Quah, Mohd, Savage,
Grismer, Ahmad, Greer & Remegio, 2010, C. stongensis, C. tapanuli, and C. temiah.
The new species possesses caudal tubercles encircling the tail which differentiates it from Cnemaspis affinis,
C. andalas, C. argus, C. bayuensis, C. biocellata, C. dringi, C. flavigaster, C. leucura, C. grismeri, C. hangus, C.
karsticola, C. kumpoli, C. laoensis, C. limi, C. mahsuriae Grismer, Wood, Quah, Anuar, Ngadi & Ahmad, 2015, C.
minang, C. mcguirei, C. monachorum, C. narathiwatensis, C. nigridia, C. pagai, C. paripari, C. perhentianensis,
C. pseudomcguirei, C. roticanai Grismer & Chan, 2010, C. selamatkanmerapoh, C. sundainsula, C. shahruli, C.
stongensis, C. tapanuli, and C. temiah.
In having enlarged median row of subcaudals, Cnemaspis muria sp. nov. can be distinguished from C. aceh,
C. andalas, C. bidongensis, C. biocellata, C. bidongensis, C. dezwaani, C. jacobsoni, C. kumpoli, C. limi, C. mah-
suriae, C. minang, C. modiglianii, C. monachorum, C. mumpuniae, C. nigridia, C. pagai, C. paripari, C. pemang-
gilensis, C. sundagekko, C. sundainsula, C. whittenorum, and C. tapanuli.
The new species can be separated from Cnemaspis andalas, C. biocellata, C. flavigaster, C. kumpoli, C. laoen-
sis, C. limi, C. minang, C. monachorum, C. sundainsula, and C. tapanuli by having keeled subcaudal scales.
The new species lacks a vertebral stripe which separates it from C. aceh, C. andalas, C. dezwaani, C. flavolin-
eata*, C. jacobsoni, C. narathiwatensis, C. pseudomcguirei*, C. shahruli*, C. tapanuli, C. temiah* and C. whit-
tenorum (species with variable presence of this character indicated by an asterisk *).
Discussion
The discovery of Cnemaspis muria sp. nov. on the island of Java is biogeographically significant, since the occur-
rence of this genus was unexpected for the island. The southernmost and closest known species in distribution is C.
rajabasa, from Lampung, Sumatra. Gunung Muria is separated from Sumatera by ~ 580 km, Kalimantan by ~350
ROCK GECKO FROM JAVA, INDONESIA Zootaxa 4608 (1) © 2019 Magnolia Press · 171
km, and Belitung Island (the type locality of C. purnamai) by ~470 km. The African-Asian Frog genus Chiromantis
also occurs in Java (Riyanto & Kurniati 2014; Wostl et al. 2017), as do other recently described reptiles and am-
phibians (Riyanto et al. 2014; Riyanto et al. 2015; Hartmann et al. 2016; Kieckbusch et al. 2016; Wostl et al. 2017;
Hamidy et al. 2018). These novelties have shown Java to be a more herpetologically interesting place than hereto-
fore believed, even though it was believed to have been relatively well studied during the long history of exploration
during the Dutch colonial period.
The new species was found active in rock microhabitats along rivers and in coffee plantations. With little eco-
logical data we may assume that Cnemaspis muria sp. nov. participates in controlling insect populations, including
pests on coffee plants. As such there are conservation implications for this species. The use of herbicides and insec-
ticides in coffee plantations should be limited, and only applied where there is a serious pest problem. An integrated
pest control management approach should take into account the role of the new species, and that of other lizards, as
natural pest control agents, and not put their existence in jeopardy.
Acknowledgments
AR and MM are indebted to Mr. Budhi Sutrisno and his family for providing housing during fieldwork in Gunung
Muria, and also thank Rubby Alfian, Lianita Rarasandy, and Rega D. Ganiarta for the hospitality, assistance, and
field companionship. AISM is grateful to Ripin (Gaia Consulting) for showing a first picture of the new species,
which lead to further exploration. AISM also thanks to Nur and Roping for assistance during surveys. Eric N Smith
(University of Texas at Arlington, USA), Aaron M Bauer (Villanova University, USA) and an anonymous reviewer
for helpful comments to improve the manuscript. AISM biodiversity fieldwork was funded by Djarum Foundation
through YKAN. AR and MM fieldwork was self-funded. Laboratory work were made possible by a research grant
from the Indonesian government to the Indonesian Institute of Sciences via the “Konstruksi Referensi Sekuen DNA
Barcode Dan Kajian Genetika Populasi Untuk Keperluan Konservasi, CITES, Forensik dan Pengembangan Bank
DNA Fauna Indonesia” project (Grant number 3400.001.051G) lead by YSF, and the “Penguatan Kelembagaan
Dalam Melaksanakan Mandat Sebagai Otoritas Keilmuan Terkait Keanekaragaman Hayati” project (Grant number
3400.002001.051.A) lead by AH.
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APPENDIX 1. Spesimens examined.
Cnemaspis aceh: MZB.Lace.12998 (holotype); C. Andalas: MZB.Lace.12999 (holotype), 13000 (paratype); C. minang: MZB.
Lace.13002 (holotype), 13002 (paratype); C. mumpuniae. MZB.Lace.10167 (holotype); C. muria: MZB.Lace.14564–70,
14572 (paratypes), 14571 (holotye); C. pagai: MZB.Lace.13004 (holotype); C. purnamai: MZB.Lace.14074–75 (paratypes),
14076 (holotype); C. rajabasa: MZB.Lace.6595 (holotype); C. sundainsula: MZB.Lace.9438 (holotype); MZB.Lace.2240
(holotype).
