Cyrtodactylus+majulah

=**//Cyrtodactylus majulah//** Grismer, Wood & Lim, 2012= =**Singapore Bent-toed Gecko**=

//Cyrtodactylus majulah//, the Singapore Bent-toed Gecko Photo by Noel Thomas (Used with permission) Singaporeans should be familiar with cicaks*. Many species of geckos can be found in human houses. It can be annoying to clean up the feces that they leave everywhere. Besides that, they do us no harm. However, there are several groups of geckos that do not range into such urban habitats. These lizards can only be found in specific forest types and habitats and are not so commonly encountered by the average Singaporean. One such forest gecko is the Singapore Bent-toed Gecko (Cyrtodactylus majulah). Geckos in this genus have slender, bent toes, which is what gives them their name. This page serves as a resource for interested members of public, educators and even hardcore reptile taxonomy enthusiasts. So, take some time to learn about one of the amazing animals that can be found in Singapore! A Common House Gecko (//Hemidactylus frenatus//), one of the most commonly seen geckos in urban areas Photo by Thomas Brown (CC BY 2.0)
 * Overview **
 * //Cicak// - the Malay word for Gecko

**Name** Scientific Name: //Cyrtodactylus majulah,// Grismer, Wood & Lim, 2012 Common Names: Singapore Bent-toed Gecko, Cicak Hutan Singapura

Etymology
The generic name, Cyrtodactylus, is derived from the Greek words κυρτός (kyrtos, which means bent, or crooked) and δάκτυλος (dáktylos, which means finger), literally coming to mean "bent finger". The specific epithet, majulah, in Bahasa Melayu, means to go forward or to progress. This is both an allusion to the fact that this species was described from Singapore and that Cyrtodactylus taxonomy is rapidly advancing, as many new species are being discovered and described every year [1]. The word Majulah is prominently featured in Singapore's National Anthem.

The State Crest of Singapore, depicting the motto, "Majulah Singapura" on the banner (National Archives of Singapore) Mari kita rakyat Singapura Sama-sama menuju bahagia Cita-cita kita yang mulia Berjaya Singapura Marilah kita bersatu Dengan semangat yang baru Semua kita berseru Majulah Singapura Majulah Singapura
 * Majulah Singapura **

**Native Distribution**

**Distribution of the** // **Cyrtodactylus** // **genus** This genus ranges throughout South, east and Southeast Asia, and even some parts of Northern Australia [2]. A map depicting the global distribution of Genus //Cyrtodactylus// (including //Geckoella//). Adapted from Wood Jr. et al. (2012) (Used in accordance with Fair Use) Cyrtodactylus is the largest genus of geckos, containing at least 233 species (as of October 2017) [3], with many more species still undescribed. Due to their massive trans-regional range across several mainland, peninsular and archipelagic regions, the genus has seen massive diversification [2]. These species also can exhibit an extremely high level of microendemism (where a species is restricted to a very small range) [4]. Despite the unprecedented rate of biodiversity discovery, many areas within the genus' home range have yet to be properly studied [4].

Distribution of //C. majulah//
A map displaying the two localities in which //C. majulah// has been recorded. Diagram adapted from Grismer et al. (2012) (Used in accordance with Fair Use) //Cyrtodactylus majulah// has thus far only been recorded on two islands; on Singapore Island and on Pulau Bintan of the Riau Archipelago [1]. Interestingly, it has not been recorded in Southern Peninsular Malaysia or on any other isands in the Riau Archipelago.

Within Singapore, it was thought to only exist within the Central Catchment Nature Reserve. However, there have been recent records of //C. majulah// in degraded secondary forest growth at Admiralty Park. This is thought to be a remnant population that is likely not connected to the populations in the CCNR [5].

Habitat
Over the years, the freshwater swamp forest ecosystem in Singapore has diminished drastically. Today, what is left of it is protected by various agencies, including the National Parks Board. It is a unique habitat that is characterised by the occurrence of slow streams that have a lower pH than in other such forests. This is caused by the accumulation and decay of plant matter in these streams, which releases tannic acid. This, in turn, causes the water to be acidic and tea- coloured. The streams and swamps are highly prone to flooding, which results in the forest soil being muddy and unstable. Furthermore, the waterlogged soil tends to be anaerobic as well. These factors provide a challenging but unique niche that many species of flora and fauna have exploited.[6][7] An example of Singaporean Swamp Forest habitat Photo by Law Ing Sind (Used with permission) //Cyrtodactylus majulah// was described from specimens collected at Nee Soon Swamp Forest. This is the largest patch of freshwater swamp forest in Singapore [7]. It spans just 5 km 2 but contains a disproportionately large amount of Singapore's biodiversity within it. It is connected to the Central Catchment Nature Reserve, which is the largest gazetted Nature Reserve in Singapore. Nee Soon is home to 23% of the reptiles, 59% of the amphibians, about 41% of the freshwater fish that are extant to Singapore [8]. Furthermore, countless species of insects and invertebrates may be found residing within this habitat. It is, therefore, an area of immense conservation importance. A detailed map of Nee Soon Swamp Forest, detailing its position in Singapore, topography, and surrounding landmarks. Adapted from Sun //et al.// (2016) (Used in accordance with Fair Use) Nee Soon Swamp Forest is currently a protected area that is inaccessible to the general public. It is protected by the National Parks Board and the Ministry of Defence, which conducts live-firing activities at the firing range. It is bordered by the Seletar Expressway (SLE) and a golfing range [9]. Despite its small size and proximity to urban areas, Nee Soon Swamp Forest continues to be a stronghold for many animals and plants that cannot be found elsewhere on the island, or even the world.

**Biology** **Behaviour** Natural history observations of this species are few, due to its restricted range and cryptic nature. However, an interesting observation of //C. majulah// swimming to escape from immediate threat has been recorded [10]. Similar behaviour is also recorded in the closely related //C. pantiensis// (which also occupies swamp forest habitat), where it has been recorded to "jump into the water and run across its surface to escape capture." [10]

**Feeding Habits** Little is known of the diet preferences of //C. majulah//. However, like its congeners, it likely feeds mostly on insects and other forest invertebrates. [11]

Regenerative Ability
Like other species of geckos, //C. majulah// is able to drop its tail when faced with danger. This is known as autotomy. The tail may subsequently regenerate, but will not have the colouration and patterning of the original tail. A //Cyrtodactylus majulah// with a regenerated tail Photo by Serin Subaraj (Used with permission)

Conservation Status
Since //C//. //majulah// is a relatively recent discovery, it has not been evaluated under the Singapore Red Data Book, which was last published in 2008. However, it is considered a rare species with restricted habitat by Baker & Lim (2012, as //Cyrtodactylus quadrivirgatus//) [12]

// Cyrtodactylus majulah // has a unique combination of morphological characteristics that help to distinguish it from other members of its genus [1]. It has a maximum Snout-Vent Length (SVL) of 68.0mm Illustration drawn by A. Sankar. It possesses 8-9 supralabial scales (top row), and 7-8 infralabial scales (bottom row)  Illustration drawn by A. Sankar // C. majulah // can be distinguished by the strong tuberculation exhibited on its body and limbs. It possesses 39-46 paravertebral tubercules, 43-53 ventral scales, 20-23 subdigital lamellae and 7-11 precloacal pores. It has enlarged precloacal scales and tubercules on the basal portion of the tail [1].
 * Diagnosis **

Many of these terms may seem rather technical and may not make sense if you do not understand their usage. Hopefully, this glossary can help to clarify the usage of these terms in the context of this section.
 * Anatomical Glossary**

Tubercules - A small knob or bump, which projects from the skin of the gecko Paravertebral - Occurring parallel, or adjacent to the vertebra Ventral Scales - The scales on the underside of gecko Subdigital lamellae - Expanded scales under the toe pads Precloacal pores - Pores that are found above the cloaca on the thighs of the gecko, which are used to excrete hormones

**Colouration** //C. majulah// (like many other //Cyrtodactylus// species) exhibits some variation in colouration and patterning. Nevertheless, there are various distinguishing features that can be used for reliable field identification. These hallmark features have been summarised in the picture below. Key colouration and patterning of //C. majulah.// Photo by Noel Thomas (Used with permission), Annotations by A. Sankar Besides the highlighted features, it is also noted that each ventral scale bears one or two tiny black spots. Furthermore, the region underneath the tail is dark brown and displays irregularly shaped beige blotches. [1]

Besides //C. majulah//, several other species of //Cyrtodactylus// have been recorded in Singapore. They may be distinguished based on their characteristics, and the areas in which they are found.
 * Other //Cyrtodactylus// species in Singapore **
 * Species Name and Other Resources || Photo || Identifying features ||
 * //Cyrtodactylus consobrinus//

(Peter's Bent-toed Gecko) 1. EcologyAsia 2. __The Biodiversity of Singapore Database__ 3. __[|NParks Flora & Fauna Web]__ || Photo by Serin Subaraj (Used with permission) || Easily distinguishable from the other //Cyrtodactylus// species in Singapore. Possesses a distinctive black and white/cream banding throughout its body and tail. [13]

A reticulated pattern can also be observed on //C. consobrinus//' head and rostrum. [13]

In Singapore, is found in primary forest, usually restricted to crevices in trees and large rocks formations. [14] || (Panti Bent-toed Gecko) 1. EcologyAsia || Photo by Nick Baker (Photo taken in Gunung Panti) (Used with Permission) || A pair of mirrored, half-moon shaped, blotches can be seen at the base of the lizard's skull. [15]
 * //Cyrtodactylus pantiensis//

The dark brown blotches form four broken longitudinal lines that extend till the base of the tail

Creamy-yellow speckles can be seen in between all the blotches and in the banding on the tail.

Thus far, in Singapore, this species has only been recorded on Pulau Tekong [16]. Like //C. majulah//, this species also appears to be a swamp-forest specialist [1]. || (Marbled Bent-toed Gecko) 1. EcologyAsia || Photo by Serin Subaraj (Used with permission) || This species is highly variable in patterning and colouration throughout its range. It may be difficult to identify from a few pictures.
 * //Cyrtodactylus quadrivirgatus//

Generally, the species is distinguished by the semi-continuous longitudinal lines that extend from behind the eyes and nape till the base of the tail. [17]

Individuals of //C//. //majulah// were considered to belong to //C. quadrivirgatus// until the description of the former in 2012. [1] || (Peninsular Bent-toed Gecko) 1. [|EcologyAsia] _ 2. The Biodiversity of Singapore Database || Photo by Nick Baker (Used with permission) || Possesses a dark stripe behind the eye that ends just after the forelimbs. The two eyestripes on either side are connected over the back by a dark, broken bar. Five to six such broken bars extend across the back until the base of the tail [18].
 * //Cyrtodactylus semenanjungensis//

Similar to //C. majulah//, individuals of this species were classified under //C. quadrivirgatus// until formally being described in 2005. [1]

Like //C. majulah// and //C. pantiensis//, this species is also found around swamp forest habitat [19] ||


 * Type Information **

Holotype
The holotype is the single specimen upon which the description for the entire species is based. The specimen serves as the name-bearer for the species and is usually safely kept in a museum.

Adult male, 67.9 mm SVL (ZRC 2.6950) Singapore: Central Catchment Nature Reserve, Nee Soon Swamp-forest (1°22'48.99N, 103°49'05.39E;51m) Collected by: L. Grismer, E. Quah, K.O. Chan, A. Figueroa & A. Devan-Song, 22 Jun 2011. Stored in: Lee Kong Chian Natural History Museum

**Paratypes** Paratypes are other members of the type series. They are a group of individuals that demonstrate variation within the species. Paratypes are not name-bearing specimens.

Adult male: 68.0 mm SVL (ZRC 2.6951) Adult male: 62.7 mm SVL (ZRC 2.6952) Adult male: 67.2 mm SVL (LSUHC 10458) Adult male: 67.2 mm SVL (ZRC 2.6953) Collection data as per holotype

**Synonyms** Synonyms in the taxonomic sense are not the same as synonyms in the literary sense. Over time, a species may be referred to by a variety of scientific names by various authorities. As per the International Code of Zoological Nomenclature, the oldest available name will take precedence over all other synonyms unless it is invalidated by a reclassification or redescription.

Due to the intricate morphological details that distinguish the various species of //Cyrtodactylus//, there is confusion in many of the records with regards to what species is being recorded. C. //majulah// has been known by a variety of names over the years. It was initially recorded as //Gymnodactylus marmoratus// in the Singapore Botanic Gardens in 1925 by Sworder [20]. Adapted from Sworder (1925) (Used in accordance with Fair Use) Subsequently, until it was properly described as a distinct species, //C. majulah// was grouped under //C. quadrivirgatus// [21]//.// Excerpt from Dring (1979) (Used in accordance with Fair Use) After //C. majulah// was properly described, it was thought that all //C. quadrivirgatus// sightings prior were conspecific with //C. majulah//. (i.e. All the //C. quadrivirgatus// records in Singapore prior to the description were actually //C. majulah//) [1]. However, to add to the confusion, in 2017, an individual of //C. quadrivirgatus// was recorded in Upper Seletar, showing that the two species may actually be sympatric* in Singapore [22].


 * Sympatric - When two species exist in the same geographic area.

//Gymnodactylus marmoratus// – SWORDER 1925: 63 (not of KUHL) //Gymnodactylus marmoratus// – SMITH 1930: 12 //Cyrtodactylus quadrivirgatus// – DRING 1979: 228 (in part) //Cyrtodactylus quadrivirgatus// – DENZER & MANTHEY 1991: 314 //Cyrtodactylus quadrivirgatus// – LIM & LIM 1992: 121 (not of TAYLOR) //Cyrtodactylus quadrivirgatus// – MANTHEY & GROSSMANN 1997: 227 //Cyrtodactylus quadrivirgatus// – CHAN-ARD et al., 1999: 24 //Cyrtodactylus quadrivirgatus// – BAKER & LIM 2008: 80 //Cyrtodactylus majulah// GRISMER, WOOD & LIM 2012
 * List of Synonyms [3]**

Taxonavigation [23]

 * **Taxonomic Hierachy** ||  ||
 * Kingdom || Animalia ||
 * Phylum || Chordata ||
 * Subphylum || Vertebrata ||
 * Superclass || Tetrapoda ||
 * Superorder || Lepidosauria ||
 * Order || Squamata ||
 * Infraorder || Gekkota ||
 * Family || Gekkonidae ||
 * Subfamily || Gekkoninae ||
 * Genus || //Cyrtodactylus// ||
 * Species || //majulah// ||

Pyron //et al.// (2013) conducted the largest phylogenetic estimate (4161 species) for the Order Squamata (which comprises all lizards and snakes). A total of 12 genes were sequenced, including seven nuclear genes and five mitochondrial genes. This study helped to identify relationships between many clades of Squamates, including Gekkotans. [24] Maximum Likelihood phylogenetic analysis with a 12 gene concatenated matrix. Nodes with support values exceeding 85 are considered to have strong support. Diagram adapted from Pyron //et al.// (2013) (Used in accordance with Fair Use). Annotations by A. Sankar. The phylogenetic tree produces shows that the placement of the two genera //Cyrtodactylus// and //Hemidactylus// next to each other on the tree has a high ML Bootstrap value (97). This indicates that their placement as sister genera has high support in the phylogenetic analysis.
 * Sister Group to //Hemidactylus// **

Swamp Forest Specialists
Grismer //et al.// (2012), while describing //C. majulah//, sequenced a 1505bp fragment of the ND2 mitochondrial gene from various //Cyrtodactylus//, species, including //C. quadrivirgatus//, which was previously thought to be conspecific with //C. majulah//. [1] Maximum Likelihood diagram with Bayesian Posterior Probability (BPP), Maximum Likelihood (ML) and Maximum Parsimony (MP) values being expressed at each node as (BPP/ML/MP). Adapted from Grismer //et al.// (2012) (Used in accordance with Fair Use). Annotations by A. Sankar. The tree clusters the //C. quadrivirgatus// populations together, but does not group //C. majulah// in the same cluster, indicating that they are not conspecific, as was initially thought. Instead, the study grouped //C. majulah// as a basal group to //C. pantiensis// and //C. payacola.// This relationship is well supported by the phylogenetic analysis, with high BPP, and ML/MP bootstrap values (1.0/100/100) for all the nodes, indicating that they are very stable. //C. semenanjungensis// is clustered as basal to these three species. However, BPP and ML/MP bootstrap values (0.86/52/53) indicate that this relationship has somewhat inconsistent support. The ND2 gene is useful in reliably analysing phylogenetic relationships in the //Cyrtodactylus// genus [25]. Altogether, //C. majulah//, //C. pantiensis//, //C. payacola//, and //C. semenanjungensis// represent a clade of swamp-forest dwelling Sundaland species. It is likely that they had common ancestors that also specialised in freshwater swamp forest habitat and subsequently speciated allopatrically. Future biodiversity surveys can potentially target swamp forests in nearby islands to uncover heretofore unknown members of this clade.


 * Literature and References **

[1] - Grismer LL, Wood PL & Lim KKP (2012) //Cyrtodactylus majulah//, a new species of bent-toed gecko (Reptilia: Squamata: Gekkonidae) from Singapore and the Riau Archipelago. The Raffles Bulletin of Zoology 60(2): 487-499 [2] - Wood PL, Heinicke MP, Jackman TR & Bauer AM (2012) Phylogeny of bent-toed geckos (//Cyrtodactylus//) reveals a west to east pattern of diversification. Molecular Phylogenetics and Evolution, 65(3): 992-1003 [3] - Uetz, P. (editor), The Reptile Database, http://www.reptile-database.org, accessed November 15, 2017. [4] - Grismer LL, Wood, PL, Thura M, Zin T, Quah E, Murdoch M, Grismer MS, Aung L, Htet K & Lwin N (2017) Twelve new species of //Cyrtodactylus// Gray (Squamata: Gekkonidae) from isolated limestone habitats in east-central and southern Myanmar demonstrate high localized diversity and unprecedented microendemism. Zoological Journal of the Linnean Society, zlx057 [5] - Law IS (2017). Singapore bent-toed gecko at Admiralty Park. Singapore Biodiversity Records 2017: 77 [6] - Turner IM, Boo CM, Wong YK, Chew PT & Ali BI (1996) Freshwater swamp forest in Singapore, with particular reference to that found around the Nee Soon Firing Ranges. Gardens' Bulletin, Singapore, 48 (1&2): 129-157 [7] - Lee Kong Chian Natural History Museum (2017) Nee Soon Swamp Forest, The Digital Nature Archive of Singapore, https://lkcnhm.nus.edu.sg/dna/places/details/13, accessed November 15, 2017 [8] - Li TJ, Chee KC, Lim WH & Cai YX (2016) The fish fauna of Nee Soon Swamp Forest, Singapore. Raffles Bulletin of Zoology, Supplement No. 32: 56-84 [9] - Sun YB, Dadiyorto W, Kim DE & Liong SY (2016) Development and Application of an Integrated Hydrological Model for Singapore Freshwater Swamp Forest. Procedia Engineering, 154: 1002-1009 [10] - Groenewoud D (2015) Singapore bent-toed gecko swimming underwater. Singapore Biodiversity Records 2015: 42 [11] - National Parks Board (2017) //Cyrtodactylus majulah.// Flora and Fauna Web, https://florafaunaweb.nparks.gov.sg/Special-Pages/animal-detail.aspx?id=6479, accessed November 15, 2017 [12] - Baker N & Lim KKP (2012) Wild Animals of Singapore. Singapore: Draco Publishing and Distribution Pte Ltd. 180pp [13] - Baker N (2017) Peter's Forest Gecko. Ecology Asia, http://www.ecologyasia.com/verts/lizards/peters-forest_gecko.htm, accessed November 15, 2017 [14] - National Parks Board (2017) //Cyrtodactylus consobrinus.// Flora and Fauna Web, https://florafaunaweb.nparks.gov.sg/Special-Pages/animal-detail.aspx?id=253, accessed November 15, 2017 [15] - Baker N (2017) Panti Bent-toed Gecko. Ecology Asia, http://www.ecologyasia.com/verts/lizards/panti-bent-toed-gecko.htm, accessed November 15, 2017 [16] - Lim KKP, Chua MAH & Lim NTL (2016). Freshwater fishes, terrestrial herpetofauna and mammals of Pulau Tekong, Singapore. Nature in Singapore, 9: 165-198 [17] - Baker N (2017) Marbled Bent-toed Gecko. Ecology Asia, http://www.ecologyasia.com/verts/lizards/marbled-bent-toed-gecko.htm, accessed November 15, 2017 [18] - Baker N (2017) Peninsular Forest Gecko. Ecology Asia, http://www.ecologyasia.com/verts/lizards/peninsular-bent-toed-gecko.htm, accessed November 15, 2017 [19] - Grismer LL, Leong TM (2005) New Species of //Cyrtodactylus// (Squamata: Gekkonidae) from Southern Peninsular Malaysia. Journal of Herpetology, 39(4): 584-591 [20] - Sworder GH (1925) The lizards of Singapore Island. Singapore Nature, 5: 61–68. [21] - Dring JCM (1979) Amphibians and reptiles from Northern Trenganggu, Malaysia, with descriptions of two new geckos: //Cnemaspis// and //Cyrtodactylus//. Bulletin of the British Museum (Natural History), 34(5): 228 [22] - Law IS, Law IT & Serin S (2016) Marbled bent-toed gecko at Upper Seletar. Singapore Biodiversity Records 2016: 117-118 [23] - Encyclopedia of Life. http://www.eol.org. Accessed 3 December, 2017 [24] - Pyron R, Burbrink FT & Wiens JJ (2013) A phylogeny and revised classification of Squamata, including 4161 species of lizards and snakes. BMC Evolutionary Biology, 13(1), 93 [25] - Brennan IG, Bauer AM, Ngo VT, Wang Y, Wang W, Zhang Y & Murphy RW (2017) Barcoding utility in a mega-diverse, cross-continental genus: keeping pace with //Cyrtodactylus// geckos. Scientific Reports, 7: 5592 Phylogeny of bent-toed geckos (Cyrtodactylus) reveals a west to east pattern of diversificati  zlx057