Scatophagus+argus

toc **Figure 1. Spotted scat, //Scatophagus argus// (Linnaeus, 1766). ** **Photo of Spotted scat by ****Chinese Academy of Fishery Sciences, Information Center. ** =Overview =
 * //Scatophagus argus// (Linnaeus, 1766) **
 *  Creative Commons Attribution-Noncommercial 3.0 Unported License. **

The spotted scat (//Scatophagus argus//) is a native fish of Singapore that could be found in both brackish and freshwater environments, including monsoon drains 1]. It has an average total length of 200 mm 2]. It is an important food fish, and brackish water aquarium fish in many countries 3], including Singapore 4]. It is also known by the name “//kim kor//” in Cantonese, and “//Ikan Ketang//” in Malay. //Scatophagus argus// is also known to have been caught by anglers in the coastal waters of Singapore*.

 As this fish has **venomous spines**, it should be noted of and avoided when one enters its habitats especially the mangroves, or attempts to handle similar-looking fishes. This species page thus seeks to provide a comprehensive identification guide (see description and diagnosis) of the spotted scat, and information on the venomous spines possessed by the fish.

 *Spotted scats are reported to have been caught by anglers at the Woodland Jetty. See [] for an example.

=Name =

**Scientific name: ** //Scatophagus argus //(Linnaeus, 1766)  **Common names: **<span style="font-family: 'Times New Roman',serif; font-size: 14pt;">Spotted scat, Butterfish, //Ikan kitang// (Malay), //Kim kor// (Cantonese)

**<span style="font-family: 'Times New Roman',Times,serif; font-size: 14pt;"> Etymology **
<span style="font-family: 'Times New Roman',serif; font-size: 14pt;"> **//<span style="font-family: 'Times New Roman',serif; font-size: 14pt;">Scatophagus: //**<span style="font-family: 'Times New Roman',serif; font-size: 14pt;">means feeding upon dung**:** <span style="font-family: 'Times New Roman',serif; font-size: 14pt;"> Scat= ‘dung’, from Greek stem //skat// <span style="font-family: 'Times New Roman',serif; font-size: 14pt;"> -phagus= ‘eating, feeding on’, from Latin -phagus, from Greek –phagos <span style="font-family: 'Times New Roman',serif; font-size: 14pt;"> **//<span style="font-family: 'Times New Roman',serif; font-size: 14pt;">argus: //**<span style="font-family: 'Times New Roman',serif; font-size: 14pt;">The hundred-eyed giant of Greek mythology. It’s probably so named due to the many spots on its body, resembling many eyes. 5]

**<span style="font-family: 'Times New Roman',Times,serif; font-size: 14pt;"> Synonyms **
<span style="font-family: 'Times New Roman',serif; font-size: 14pt;">// Chaetodon argus // Linnaeus, 1766 <span style="font-family: 'Times New Roman',serif; font-size: 14pt;"> //<span style="font-family: 'Times New Roman',serif; font-size: 12pt;">Scatophagus argus argus //<span style="font-family: 'Times New Roman',serif; font-size: 12pt;"> (Linnaeus, 1766) <span style="font-family: 'Times New Roman',serif; font-size: 14pt;"> //<span style="font-family: 'Times New Roman',serif; font-size: 12pt;">Ephippus argus //<span style="font-family: 'Times New Roman',serif; font-size: 12pt;"> (Linnaeus, 1766) <span style="font-family: 'Times New Roman',serif; font-size: 14pt;"> // Chaetodon pairatalis // Hamilton, 1822 <span style="font-family: 'Times New Roman',serif; font-size: 14pt;"> //<span style="font-family: 'Times New Roman',serif; font-size: 12pt;">Chaetodon atromaculatus //<span style="font-family: 'Times New Roman',serif; font-size: 12pt;"> Bennett, 1830 <span style="font-family: 'Times New Roman',serif; font-size: 14pt;"> //<span style="font-family: 'Times New Roman',serif; font-size: 12pt;">Scatophagus purpurascens //<span style="font-family: 'Times New Roman',serif; font-size: 12pt;"> Cuvier, 1831 <span style="font-family: 'Times New Roman',serif; font-size: 14pt;"> //<span style="font-family: 'Times New Roman',serif; font-size: 12pt;">Scatophagus ornatus //<span style="font-family: 'Times New Roman',serif; font-size: 12pt;"> Cuvier, 1831 <span style="font-family: 'Times New Roman',serif; font-size: 14pt;"> //<span style="font-family: 'Times New Roman',serif; font-size: 12pt;">Scatophagus bougainvillii //<span style="font-family: 'Times New Roman',serif; font-size: 12pt;"> Cuvier, 1831 <span style="font-family: 'Times New Roman',serif; font-size: 14pt;"> // Sargus maculatus // Gronow, 1854 <span style="font-family: 'Times New Roman',serif; font-size: 14pt;"> // Scatophagus argus ocellata // Klunzinger, 1880 <span style="font-family: 'Times New Roman',serif; font-size: 14pt;"> //<span style="font-family: 'Times New Roman',serif; font-size: 12pt;">Scatophagus quadratus //<span style="font-family: 'Times New Roman',serif; font-size: 12pt;"> De Vis, 1882 <span style="font-family: 'Times New Roman',serif; font-size: 14pt;"> //<span style="font-family: 'Times New Roman',serif; font-size: 12pt;">Scatophagus aetatevarians //<span style="font-family: 'Times New Roman',serif; font-size: 12pt;"> De Vis, 1884
 * <span style="font-family: 'Times New Roman',Times,serif; font-size: 11pt;">(Referenced from IUCN) **

=<span style="color: #0fad85; font-family: 'Times New Roman',Times,serif; font-size: 24px;"> <span style="color: #0fad85; font-family: 'Times New Roman',Times,serif; font-size: 130%;">Description =

media type="custom" key="26901506" align="center" <span style="font-family: 'Times New Roman',serif; font-size: 14pt;">Generally, scats have a distinctly quadrangular and highly laterally compressed body which is usually greenish or silvery with dark spots or bars 6]. The figure below shows other descriptive characters of //S. argus.//



<span style="font-family: 'Times New Roman',serif; font-size: 14pt;">**<span style="color: #4f7fbd; font-family: 'Times New Roman',Times,serif; font-size: 12pt;">Figure 2. Diagnostic characteristics of the family Scatophagidae (adapted from Kottelat, 2001 6] with additional labels added). **

<span style="color: #0fad85; font-family: 'Times New Roman',Times,serif; font-size: 14pt;">Size
<span style="font-family: 'Times New Roman',serif; font-size: 14pt;">The average total length of //S. argus// is about 200 mm 2], and they can grow up to a total length of 350 mm 6].

<span style="color: #0fad85; font-family: 'Times New Roman',Times,serif; font-size: 14pt;">Adults and Juveniles
<span style="font-family: 'Times New Roman',serif; font-size: 14pt;">In particular, adults //S. argus// have dark spots that could be faded, and more concentrated at the dorsal side of the body 6](figure 3). Juveniles resemble the adults but have dark spots about the same diameter as the eyes (figure 4, right) or have 5 – 6 dark and broad vertical bars along their body 6] (figure 4, center).



<span style="font-family: 'Times New Roman',serif; font-size: 14pt;">**<span style="color: #4f7fbd; font-family: 'Times New Roman',Times,serif; font-size: 12pt;">Figure 3. Illustration of //S. argus// in the adult stage (Image from FAO in De Bruin et al., 1995) 35]. **

<span style="font-family: 'Times New Roman',serif; font-size: 14pt;">// Scatophagus argus // go through a tholichthys postlarval stage 6]. The tholichthys larval stage is unique to fishes in the family Chaetodontidae (butterflyfishes) and Scatophagidae, and is characterized by having large body plates that enclose the body, particularly the head, forming a hard protective covering 7]. These larvae are small, ranging between a total length of 6mm to 12mm. These protective plates disappear as the larva develops into a juvenile 7].



<span style="font-family: Arial,Helvetica,sans-serif; font-size: 12pt;">**<span style="color: #4f7fbd; font-family: 'Times New Roman',Times,serif; font-size: 12pt;">Figure 4. Left: An illustration of a tholichthys larva of //S. argus// of 12mm. Bony plates encloses the body and forms a protective cover around the head of the larva (Image by and used with permission from Barry & Fast, 19887]; Pending permission from journal). Center: ** **<span style="color: #4f7fbd; font-family: 'Times New Roman',Times,serif; font-size: 12pt;">Illustration of juvenile //S. argus// of 17.1 mm standard length (Image by FAO in Kottelat, 2001 6]). ****<span style="color: #4f7fbd; font-family: 'Times New Roman',Times,serif; font-size: 12pt;"> Right: Illustration of juvenile //S. argus// of 30.2mm standard length (Image by FAO in Kottelat, 2001 ****<span style="color: #4f7fbd; font-family: 'Times New Roman',Times,serif; font-size: 12pt;">6] ****<span style="color: #4f7fbd; font-family: 'Times New Roman',Times,serif; font-size: 12pt;">). **

<span style="color: #0fad85; font-family: 'Times New Roman',Times,serif; font-size: 14pt;">Sex determination
<span style="font-family: 'Times New Roman',serif; font-size: 14pt;">//Scatophagus argus// exhibit slight sexual dimorphism. Males and females can be distinguished by their head profile 7]. In females, the head profile ascends in a gentler slope, whereas males have an abrupt steep sloping of the head above the eyes. In addition, females are of a lighter colour than males 8].



<span style="font-family: Arial,Helvetica,sans-serif; font-size: 12pt;">**<span style="color: #4f7fbd; font-family: 'Times New Roman',Times,serif; font-size: 12pt;">Figure 5. The head profile of male (left) and female (right) //S. argus// (Image by and used with permission from Barry & fast, 1992 8]; Pending permission from journal). **

=<span style="color: #0fad85; font-family: 'Times New Roman',Times,serif; font-size: 130%;">Diagnosis =

<span style="font-family: 'Times New Roman',serif; font-size: 19px; line-height: 28px;">The family Scatophagidae consists of two genera, //<span style="font-family: 'Times New Roman',serif; font-size: 19px; line-height: 28px;">Scatophagus //<span style="font-family: 'Times New Roman',serif; font-size: 19px; line-height: 28px;"> and //<span style="font-family: 'Times New Roman',serif; font-size: 19px; line-height: 28px;">Selenotoca //<span style="font-family: 'Times New Roman',serif; font-size: 19px; line-height: 28px;"> 9].

<span style="color: #0fad85; font-family: 'Times New Roman',Times,serif; font-size: 14pt;">Genus //Scatophagus//
<span style="font-family: 'Times New Roman',serif; font-size: 14pt;">Unlike //Selenotoca//, the gill membrane of //Scatophagus// forms a free fold that covers the isthmus (or throat area of fish) as depicted in figure 6a below 6]. <span style="font-family: 'Times New Roman',serif; font-size: 14pt;"> **<span style="color: #4f81bd; font-family: 'Times New Roman',serif; font-size: 12pt;">Figure 6 ****<span style="color: #4f81bd; font-family: 'Times New Roman',serif; font-size: 12pt;">. Ventral view of a) //Scatophagus argus// and b) and a //Selenotoca// species, //Selenotoca multifasciata//. In //Scatophagus//, the gill membrane forms a fold that covers the isthmus but not in //Selenotoca// (Image from FAO in Kottelat, 2001 ** **<span style="color: #4f7fbd; font-family: 'Times New Roman',Times,serif; font-size: 12pt;">6] **<span style="font-family: 'Times New Roman',serif; font-size: 14pt;">** ) **.

<span style="color: #0fad85; font-family: 'Times New Roman',Times,serif; font-size: 14pt;">Species //Scatophagus argus//
<span style="font-family: 'Times New Roman',serif; font-size: 14pt;"> The genus //Scatophagus// consists of only two species 9], the other being //S. tetracanthus// (Lacepède, 1802).

<span style="font-family: 'Times New Roman',serif; font-size: 14pt;"> //<span style="font-family: 'Times New Roman',serif; font-size: 14pt;">Scatophagus argus //<span style="font-family: 'Times New Roman',serif; font-size: 14pt;"> can mainly be distinguished from //S. tetracanthus// by having a greenish silver coloured body with dark round spots on its upper part of its body 6] while //S. tetracanthus// has a yellow body with seven vertical dark bars across its body 10].


 * <span style="color: #4f7fbd; font-family: 'Times New Roman',Times,serif; font-size: 12pt;">Figure 7. Left: //Scatophagus argus// has greenish silver body with dark spots on its body whereas, right: //S. tetracanthus// has a yellow body with seven dark vertical bars. (Image of //S. argus// by John E. Randall, obtained from Fishbase. Creative Commons Attribution-Noncommercial 3.0 Unported License; Image of //S. tetracanthus// by Liliane Moeremans, used with permission and obtained from Fishbase) **

=<span style="color: #0fad85; font-family: 'Times New Roman',Times,serif; font-size: 130%;">Similar-looking coastal fishes =

<span style="font-family: 'Times New Roman',serif; font-size: 14pt;">**<span style="color: #4f7fbd; font-family: 'Times New Roman',Times,serif; font-size: 12pt; line-height: 1.5;">Figure 8. Both of these fishes can be found in the coastal areas of Singapore. ** **<span style="color: #4f7fbd; font-family: 'Times New Roman',Times,serif; font-size: 12pt; line-height: 1.5;">On first look, both //S. argus// (left) and the spotted green pufferfish (right) may look alike (photos by and used with permission from Ria Tan). **

<span style="font-family: 'Times New Roman',serif; font-size: 14pt;">Other native fishes, the **spotted green pufferfish** (//Tetraodon nigroviridis//) and **Orange-spotted rabbitfish** (//Siganus// //guttatus//) resemble //S. argus// superficially, and co-occur with //S. argus// in coastal waters of Singapore. Thus, these three fishes could be easily misidentified.

<span style="font-family: 'Times New Roman',serif; font-size: 19px; line-height: 28px;">Nevertheless, it is important that **<span style="font-family: 'Times New Roman',serif; font-size: 19px; line-height: 28px;">handling of all three fishes with hands are avoided as they are all venomous- **<span style="font-family: 'Times New Roman',serif; font-size: 19px; line-height: 28px;"> rabbitfishes and //<span style="font-family: 'Times New Roman',serif; font-size: 19px; line-height: 28px;">S. argus //<span style="font-family: 'Times New Roman',serif; font-size: 19px; line-height: 28px;"> have **<span style="font-family: 'Times New Roman',serif; font-size: 19px; line-height: 28px;">venomous spines **<span style="font-family: 'Times New Roman',serif; font-size: 19px; line-height: 28px;"> 11]while the pufferfish may excrete tetrodoxins through their skin which could be readily absorbed through our skin 33].

<span style="font-family: 'Times New Roman',serif; font-size: 19px; line-height: 28px;">The spotted green pufferfish co-occurs with //<span style="font-family: 'Times New Roman',serif; font-size: 19px; line-height: 28px;">S. argus //<span style="font-family: 'Times New Roman',serif; font-size: 19px; line-height: 28px;"> in the mangrove areas of Singapore 36] while the orange-spotted rabbitfish are more commonly found in the coastal areas further from the shore, where seagrass and corals are 11].

<span style="font-family: 'Times New Roman',serif; font-size: 19px; line-height: 28px;">It seems that both the orange-spotted rabbitfish and //S. argus// could be fished from the same coastal area as accounted by an angler in a blog. <span style="font-family: 'Times New Roman',serif; font-size: 14pt;">Along with other species of rabbitfishes, the <span style="font-family: 'Times New Roman',serif; font-size: 19px;">orange-spotted <span style="font-family: 'Times New Roman',serif; font-size: 14pt;">rabbitfish, like //S. argus//, are also sold in markets whole and eaten during the Chinese Lunar New Year period in Singapore (Kwik, 2014, pers comms).

<span style="font-family: 'Times New Roman',serif; font-size: 19px;">The table below shows how the spotted green pufferfish and rabbitfish can be distinguished from //<span style="font-family: 'Times New Roman',serif; font-size: 19px; line-height: 1.5;">S. argus //<span style="font-family: 'Times New Roman',serif; font-size: 19px;"> by comparing their body shape, <span style="font-family: 'Times New Roman',serif; font-size: 19px; line-height: 28.5px;">body colour, <span style="font-family: 'Times New Roman',serif; font-size: 19px; line-height: 1.5;">features of jaws <span style="font-family: 'Times New Roman',serif; font-size: 19px; line-height: 28.5px;">, and <span style="font-family: 'Times New Roman',serif; font-size: 19px; line-height: 1.5;"> dorsal fin.

<span style="font-family: 'Times New Roman',Times,serif; font-size: 11pt;">Image Sahat Ratmuangkhwang from Fishbase || <span style="font-family: 'Times New Roman',Times,serif; font-size: 11pt;">Image by Kuang Hsiang Liu. Creative Commons Attribution NonCommercial license. Obtained from Fishbase. || <span style="font-family: 'Times New Roman',Times,serif; font-size: 11pt;">Image by Sahat Ratmuangkhwang, obtained from Fishbase || <span style="font-family: 'Times New Roman',Times,serif; font-size: 14pt;">Orange spots on body, large <span style="font-family: 'Times New Roman',Times,serif; font-size: 19px; line-height: 28px;">golden spot below the end of dorsal fin ||
 * <span style="color: #4f7fbd; font-family: 'Times New Roman',Times,serif; font-size: 12pt;">Table 1. Table of comparison of features between //S. argus//, spotted green pufferfish 12; 13] and rabbitfish 14; 11). **
 * || [[image:scat black background edited.JPG width="282" height="260"]]
 * **<span style="font-family: 'Times New Roman',serif; font-size: 14pt;">Features ** || **//<span style="font-family: 'Times New Roman',serif; font-size: 14pt;">S. argus //** || **<span style="font-family: 'Times New Roman',serif; font-size: 14pt;">Spotted green pufferfish ** || **<span style="font-family: 'Times New Roman',serif; font-size: 14pt;">Orange-spotted rabbitfish ** ||
 * **<span style="font-family: 'Times New Roman',serif; font-size: 14pt;">Body shape ** || <span style="font-family: 'Times New Roman',serif; font-size: 14pt;">Laterally flattened and quadrangular || <span style="font-family: 'Times New Roman',serif; font-size: 14pt;">Blunt and round || <span style="font-family: 'Times New Roman',serif; font-size: 14pt;">Laterally flattened, oval-shaped, deep-bodied ||
 * **<span style="font-family: 'Times New Roman',Times,serif; font-size: 19px;">Body colour ** || <span style="font-family: 'Times New Roman',Times,serif; font-size: 19px;">Greenish silver body with dark spots || <span style="font-family: 'Times New Roman',Times,serif; font-size: 19px;">Bright green body and white belly; with dark spots || <span style="font-family: 'Times New Roman',Times,serif; font-size: 14pt;">Bluish colour at upper body, silvery below.
 * **<span style="font-family: 'Times New Roman',serif; font-size: 14pt;">Jaws **<span style="font-family: 'Times New Roman',Times,serif; font-size: 11pt;">(see figures below) || <span style="font-family: 'Times New Roman',serif; font-size: 14pt;">Jaws bear narrow finger-shaped teeth and hidden in mouth || <span style="font-family: 'Times New Roman',serif; font-size: 14pt;">Jaws modified to form beaks || <span style="font-family: 'Times New Roman',serif; font-size: 14pt;">Jaws bear slender, close-set teeth ||
 * **<span style="font-family: 'Times New Roman',serif; font-size: 14pt;">Dorsal fin **<span style="font-family: 'Times New Roman',Times,serif; font-size: 11pt;">(see figures below) || <span style="font-family: 'Times New Roman',serif; font-size: 14pt;">Long and bears spines || <span style="font-family: 'Times New Roman',serif; font-size: 14pt;">Short, bears no spine and located posteriorly on body || <span style="font-family: 'Times New Roman',serif; font-size: 14pt;">14 dorsal fin spine, first dorsal fin spine pointed forward ||

<span style="color: #4f7fbd; font-family: 'Times New Roman',Times,serif; font-size: 12pt; line-height: 0px; overflow: hidden;">
 * <span style="color: #4f7fbd; font-family: 'Times New Roman',Times,serif; font-size: 12pt; line-height: 1.5;">Figure 9. Diagram showing the differences in basic body plans of all three types of fishes (images by ****<span style="color: #4f7fbd; font-family: 'Times New Roman',Times,serif; font-size: 12pt; line-height: 1.5;">FAO in ****<span style="color: #4f7fbd; font-family: 'Times New Roman',Times,serif; font-size: 12pt; line-height: 1.5;">Fischer, 1978 14]). **

=<span style="color: #0fad85; font-family: 'Times New Roman',Times,serif; font-size: 130%;">Native Distribution & Habitat =

media type="custom" key="26901726" align="center" <span style="font-family: Arial,Helvetica,sans-serif; font-size: 12pt;">**<span style="color: #4f7fbd; font-family: 'Times New Roman',Times,serif; font-size: 12pt; line-height: 1.5;">Figure 10. Red and yellow colours highlight the native distribution of //S. argus.// The colours represent the probability of occurrence of //S. argus//, with red representing the highest probability of occurrence (0.8- 1.0) and yellow, the loweest probability of occurrence (0.01- 0.19) (Computer Generated Native Distribution Map for //Scatophagus argus// (Spotted scat) by Fishbase, 2013) 34]. **

<span style="font-family: Arial,Helvetica,sans-serif; font-size: 12pt;">//<span style="font-family: 'Times New Roman',serif; font-size: 14pt;">Scatophagus argus //<span style="font-family: 'Times New Roman',serif; font-size: 14pt;"> has a wide native range which spans across the whole coastal area of the Indian Ocean, and the tropical to warm temperate regions of Indo-West Pacific region, including Australia, Tahiti, and the Southern areas of China, and Taiwan 2, 6]. <span style="font-family: 'Times New Roman',serif; font-size: 14pt;">//Scatophagus argus// is the only //Scatophagus// species that occurs in the Southeast Asian region 9].

<span style="font-family: 'Times New Roman',serif; font-size: 14pt;">//Scatophagus argus// thrive in both freshwater and brackish waters, and are usually found between surface to a depth of 5m 15]. In Singapore, //S. argus// has been spotted along coastal areas, where mangroves and seagrass occur, as well as under jetties and near monsoon drains 4]. It has been caught or sighted along the [|northern](Johor Straits) and southern coast of Singapore.

media type="youtube" key="3U0R-nV0QrU" width="451" height="347" align="center"


 * <span style="color: #4f7fbd; font-family: 'Times New Roman',Times,serif; font-size: 12pt;">Video 1. //Scatophagus argus// can be caught at the Woodlands Waterfront Jetty. **

<span style="font-family: 'Times New Roman',serif; font-size: 14pt;">Listed below are examples of places in Singapore where //S. argus// was recorded. <span style="font-family: 'Times New Roman',Times,serif; font-size: 14pt;">**At southern coast:** Waters of Kallang, Changi, Tanah Merah <span style="font-family: 'Times New Roman',Times,serif; font-size: 14pt;">**Inland:** Lower Seletar Reservoir Yishun Dam fishing spots, Below Seletar North Link Bridge
 * <span style="font-family: 'Times New Roman',serif; font-size: 14pt; line-height: 1.5;">At northern coast: ** <span style="font-family: 'Times New Roman',serif; font-size: 14pt; line-height: 1.5;">Woodland Waterfront fishing jetty, Chek Jawa 4], Lim Chu Kang, Sungei Buloh Wetland Reserve, Pasir Ris, Pulau Seduku or Frog Island

=<span style="color: #0fad85; font-family: 'Times New Roman',Times,serif; font-size: 130%;">Biology =

<span style="color: #0fad85; font-family: 'Times New Roman',Times,serif; font-size: 14pt;">Diet and feeding habits
media type="custom" key="26901752" align="center" <span style="font-family: Arial,Helvetica,sans-serif; font-size: 12pt;">//<span style="font-family: 'Times New Roman',serif; font-size: 14pt;">Scatophagus argus //<span style="font-family: 'Times New Roman',serif; font-size: 14pt;">is usually gregarious, with adults forming smaller groups than juveniles (Kwik, 2014, pers. comms.). It is also known to feed actively during the day 2]. As mentioned in etymology, “//Scatophagus argus”// means dung-eater. It was so named as many were observed to feed on unwanted waste and offal discarded from ships at harbor 8]. However, its preference for dung matter remains unconfirmed 8]. Instead, its main diet appears to consists of detritus and algae 16]. Juvenile //S. argus// consumed a greater proportion of unicellular algae while adults consumed more multicellular algae, and a greater diversity of food items such as rotifers and sea anemones 16].

<span style="font-family: Arial,Helvetica,sans-serif; font-size: 12pt;"> //<span style="font-family: 'Times New Roman',serif; font-size: 14pt;">Scatophagus argus //<span style="font-family: 'Times New Roman',serif; font-size: 14pt;"> appear to be non-choosy feeders and would consume any food available though there might be a preference for filamentous algae in larger fishes 7,16]. Other animal matter such as copepods, fish scales and fish eggs have also been found in the guts of //S. argus// 16].

<span style="font-family: 'Times New Roman',serif; font-size: 14pt;"> Barry & Fast 7] suggest that //S. argus// is primarily herbivorous and adapted to feed on plant material given its long gut length relative to body length and the presence of “small, sharp teeth used for scrapping and shredding plant material”.

<span style="color: #0fad85; font-family: 'Times New Roman',Times,serif; font-size: 14pt;">Reproduction
media type="custom" key="26901756" align="center" <span style="font-family: Arial,Helvetica,sans-serif; font-size: 12pt;">//<span style="font-family: 'Times New Roman',serif; font-size: 14pt;">Scatophagus argus //<span style="font-family: 'Times New Roman',serif; font-size: 14pt;"> is dioecious (having distinct male and female individuals) and is able to reproduce at 1 year of age 17]. It is estimated that males reach the age of sexual maturity at 115 mm standard length, weighing about 83.5g, while females, at 140 mm standard length and 150g 8].

<span style="font-family: 'Times New Roman',serif; font-size: 14pt;">The reproductive behavior of //S. argus// in the wild does not seem to have been documented thus far. Barry & Fast 7], however, observed the mating behavior of //S. argus// in captivity: Males, when placed in a tank with a female, appear to be aggressive towards each other and attempt to guard the female. The more aggressive male was observed to chase and attack the submissive male which held its dorsal fin erect most of the time. Nudging of the female’s abdomen by the domineering male was seen as well. Both male and female were observed to undulate their bodies in rhythm while holding each other by the lip. This behaviour led to wounded upper lips in both partners. Unfortunately, the spawning behaviour was not observed during the study.

<span style="font-family: 'Times New Roman',serif; font-size: 14pt;">In the life cycle of //S. argus//, it appears that both adults and their offprings have <span style="font-family: 'Times New Roman',serif; font-size: 19px; line-height: 28px;">different salinity requirements- the offsprings live in <span style="font-family: 'Times New Roman',serif; font-size: 14pt; line-height: 1.5;">brackish water 7] while adults have to spawn in waters of higher salinity 18].

<span style="font-family: 'Times New Roman',serif; font-size: 14pt; line-height: 1.5;">The spawning season of //<span style="font-family: 'Times New Roman',serif; font-size: 14pt; line-height: 1.5;">S. argus //<span style="font-family: 'Times New Roman',serif; font-size: 14pt; line-height: 1.5;"> likely correspond to that of the rainy season 7]. A female has an average absolute fecundity (number of eggs laid in a spawning season) of 456,320 eggs 17]. It is vital that spawning occurs in high salinity, such as 25‰ 17], with fertilised eggs that stay afloat in seawater 18]. The eggs have an average diameter of 0.75mm 17] and are round and transparent 7]. In fertilised eggs, oil droplets merge together, conferring buoyancy to the eggs 18] while this does not occur in unfertilised eggs which turn opaque and sinks 7].

<span style="font-family: Arial,Helvetica,sans-serif; font-size: 12pt;"> <span style="font-family: 'Times New Roman',serif; font-size: 14pt;">With a temperature of 27—29 ℃ <span style="font-family: 'Times New Roman',serif; font-size: 14pt;"> and salinity 25 ‰ <span style="font-family: 'Times New Roman',serif; font-size: 14pt;">, fertilized eggs require about 20 to 24 hours to hatch and larvae measures between 1.75 to 1.88 mm in length 17]. First feeding begins after 3 to 4 days when the larvae’s yolk sac disappears 7,17]. The larvae then undergo a tholichthys larval stage 7] (__see figure 4__).

<span style="color: #0fad85; font-family: 'Times New Roman',Times,serif; font-size: 14pt; line-height: 1.5;">Tolerance to salinity changes
<span style="font-family: Arial,Helvetica,sans-serif; font-size: 12pt;"> //<span style="font-family: 'Times New Roman',serif; font-size: 14pt;">Scatophagus argus //<span style="font-family: 'Times New Roman',serif; font-size: 14pt;"> being euryhaline (able to thrive in both salt and freshwater) has been documented to be able to withstand sudden salinity changes between 0‰ to 30‰ 19].

=<span style="color: #0fad85; font-family: 'Times New Roman',Times,serif; font-size: 130%;">Venomous fin spine =

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<span style="color: #0fad85; font-family: 'Times New Roman',Times,serif; font-size: 14pt; line-height: 1.5;">Nature of fin spine
<span style="font-family: Arial,Helvetica,sans-serif; font-size: 12pt;"><span style="font-family: 'Times New Roman',serif; font-size: 14pt; line-height: 1.5;">Like all other Scatophagids, //<span style="font-family: 'Times New Roman',serif; font-size: 14pt; line-height: 1.5;">S. argus //<span style="font-family: 'Times New Roman',serif; font-size: 14pt; line-height: 1.5;"> possess venomous fin spines <span style="font-family: 'Times New Roman',Times,serif; font-size: 19px; line-height: 28px;">for defence purposes 7] <span style="font-family: 'Times New Roman',serif; font-size: 14pt; line-height: 1.5;"> but the pain caused by //<span style="font-family: 'Times New Roman',serif; font-size: 14pt; line-height: 1.5;">S. argus //<span style="font-family: 'Times New Roman',serif; font-size: 14pt; line-height: 1.5;"> is alleged to be more painful 20]. These spines are not used as offense but inflict painful wounds when the fish is handled carelessly- some fishermen who are oblivious of the venomous spines have been wounded by them 21]. Fortunately, the protein-based venom present in the spines is seldom life-threatening 3].

<span style="font-family: 'Times New Roman',serif; font-size: 14pt;"> The venomous spines of //S. argus// is well-described by Cameron & Endean 22]. **All fin spines of //S. argus//** **<span style="font-family: 'Times New Roman',serif; font-size: 19px; line-height: 28px;">contain venom glands- **<span style="font-family: 'Times New Roman',serif; font-size: 14pt; line-height: 1.5;"> the 11 dorsal fin spines, 2 pelvic fin spines, and 4 anal fin spines. Each spine is sharp and pointed, and consists of two lateral grooves that contain venom gland cells. <span style="font-family: 'Times New Roman',serif; font-size: 19px; line-height: 28px;">Smaller fishes were found to have relatively longer glands compared to larger fishes. <span style="font-family: 'Times New Roman',Times,serif; font-size: 14pt;">However, the longest <span style="font-family: 'Times New Roman',Times,serif; font-size: 14pt; line-height: 1.5;">gland found in any of the specimens studied did not correspond to a particular fin spine, that is, the longest venom gland can occur in any of the fin spines in //S.argus.//

<span style="font-family: 'Times New Roman',serif; font-size: 19px;">In the event that the spine punctures the skin, it erects and pressure exerted on the spine causes the venom to be released into the wound 21]. <span style="font-family: Arial,Helvetica,sans-serif; font-size: 12pt;">**<span style="color: #4f7fbd; font-family: 'Times New Roman',Times,serif; font-size: 12pt;">Figure 11. A longitudinal view of a spine of //S. argus// (adapted from Cameron & Endean, 1977 21]). Each spine possesses a pair of lateral groove containing venom glands. "gl" stands for “venom gland” and the dotted lines represent where the dermal scales end on the spine. **

<span style="color: #0fad85; font-family: 'Times New Roman',Times,serif; font-size: 14pt;">Venom's biochemical nature
<span style="font-family: 'Times New Roman',serif; font-size: 14pt;">In a study done by Sivan //et al//. 21] on the biochemical nature of the venom of //S. argus//, it was found that the venom causes haemolysis (lysis of red blood cells) in humans. In addition, when tested on mice, the venom appears to be “cytolytic, oedematic, nociceptive, proteolytic and myotoxic”, leading to tissue damage and pain.

<span style="color: #0fad85; font-family: 'Times New Roman',Times,serif; font-size: 14pt;">Symptoms
<span style="font-family: 'Times New Roman',serif; font-size: 14pt;">The effects of the venom could be felt within 5 to 10 minutes. In addition to unbearable **localized pain** at the wound that can **lasts for an hour** (Ibister, 2004), other symptoms include “**redness**, **swelling** and a **throbbing sensation** that extends to the limbs, followed by **dizziness**” 21]. The severity of the venom varies depending on the amount of venom injected, and the size of the fish 21].

<span style="color: #0fad85; font-family: 'Times New Roman',Times,serif; font-size: 14pt;">Treatment
<span style="font-family: 'Times New Roman',serif; font-size: 14pt;">The wound should be cleaned, and is typically treated by **submerging it into hot water** 21]. This suggests that wounds inflicted by //S. argus// are treated similarly as like any other [|venomous fish stings] 23]. It is important to **sought medical treatment** in case there is any likely infection of the wound and if the pain does not subside 23].

=<span style="color: #0fad85; font-family: 'Times New Roman',Times,serif; font-size: 24px;">Uses =

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<span style="color: #0fad85; font-family: 'Times New Roman',Times,serif; font-size: 14pt;">Aquarium fish
//<span style="font-family: 'Times New Roman',serif; font-size: 14pt;">Scatophagus argus //<span style="font-family: 'Times New Roman',serif; font-size: 14pt;"> is a popular aquarium fish sold in many countries including Singapore 24]. Juveniles are usually caught from the wild for aquariums fish trade 6].

media type="youtube" key="hrydKoip_64" width="448" height="350"media type="youtube" key="rKYZvKS1yXE" width="550" height="356"
 * <span style="color: #4f7fbd; font-family: 'Times New Roman',Times,serif; font-size: 12pt;">Video 2. //Scatophagus argus// in (left) freshwater aquarium and (right) brackish aquarium with the spotted green pufferfish and archer fish as tankmates (Video obtained from Youtube under fair use). **

<span style="color: #0fad85; font-family: 'Times New Roman',Times,serif; font-size: 14pt;">Food fish
//<span style="font-family: 'Times New Roman',serif; font-size: 14pt;">Scatophagus argus //<span style="font-family: 'Times New Roman',serif; font-size: 14pt;"> is also sold fresh or salted in markets as food in the Indo-Pacific islands and Southeast Asian region in countries such as Cambodia 25]. In certain countries, //S. argus// is of minor commercial importance and is not frequently sold in markets 6] while in countries, such as the Philippines, it is highly priced and considered a delicacy 7].

<span style="font-family: 'Times New Roman',serif; font-size: 14pt;">In Singapore, //S. argus// is sold in markets, such as those in Chinatown, particularly during the Chinese Lunar New Year. This is because the spawning period of //S. argus// happens to fall in the early months of the year, and when these fishes aggregate to spawn and females carry riped eggs, they are highly sought-after as food fish ( <span style="font-family: 'Times New Roman',serif; font-size: 19px;">Kwik, 2014, pers comms).

<span style="color: #797979; display: block; font-family: 'PT Serif',Georgia,serif; font-size: small; text-align: center;"> =<span style="color: #0fad85; font-family: 'Times New Roman',Times,serif; font-size: 130%;">Taxonavigation =
 * <span style="color: #4f7fbd; font-family: 'Times New Roman',Times,serif; font-size: 12pt;">Figure 12. Grilled Scatophagus argus prepared as a dish (Image by Hernee Samshudin, obtained from herneenazir.com, Creative Commons Attribution-NonCommercial 3.0 Unported License). **

<span style="font-family: 'Times New Roman',Times,serif; font-size: 14pt;">Note: Each subsequent taxon is a subset of the one before it.
 * <span style="font-family: 'Times New Roman',serif; font-size: 12pt;">(referenced from [|Integrated Taxonomic Information System (ITIS)]) **

<span style="font-family: 'Times New Roman',serif; font-size: 14pt;">Animalia <span style="font-family: 'Times New Roman',serif; font-size: 14pt;">Bilateria <span style="font-family: 'Times New Roman',serif; font-size: 14pt;">Deuterostomia <span style="font-family: 'Times New Roman',serif; font-size: 14pt;">Chordata <span style="font-family: 'Times New Roman',serif; font-size: 14pt;">Vertebrata <span style="font-family: 'Times New Roman',serif; font-size: 14pt;">Gnathostoma <span style="font-family: 'Times New Roman',serif; font-size: 14pt;">Osteichthyes <span style="font-family: 'Times New Roman',serif; font-size: 14pt;">Actinopterygii <span style="font-family: 'Times New Roman',serif; font-size: 14pt;">Neopterygii <span style="font-family: 'Times New Roman',serif; font-size: 14pt;">Teleostei <span style="font-family: 'Times New Roman',serif; font-size: 14pt;">Acanthopterygii <span style="font-family: 'Times New Roman',serif; font-size: 14pt;">Perciformes <span style="font-family: 'Times New Roman',serif; font-size: 14pt;">Acanthuroidei <span style="font-family: 'Times New Roman',serif; font-size: 14pt;">Scatophagidae* <span style="font-family: 'Times New Roman',serif; font-size: 14pt;">//Scatophagus// Cuvier in Cuvier and Valenciennes, 1831 <span style="font-family: 'Times New Roman',serif; font-size: 14pt;">//Scatophagus argus// (Linnaeus, 1766)

<span style="font-family: 'Times New Roman',Times,serif; font-size: 11pt;">* It must be noted that Scatophagidae in Pisces is not a junior homonym of Scatophagidae in Diptera. The spelling of “Scatophagidae” in Diptera is a misspelling of the correct term, “Scathophagidae”, which is based on the genus //Scathophaga// Meigen, 1803: 277 30].

=<span style="color: #0fad85; font-family: 'Times New Roman',Times,serif; font-size: 130%;">Phylogeny =

media type="custom" key="26902798" align="center" <span style="font-family: 'Times New Roman',serif; font-size: 14pt;">Currently, it seems that the phylogenetic relationships within the suborder Acanthuroidei have yet to be resolved, with different families proposed to be the sister family of Scatophagidae 26, 27, 28].

<span style="font-family: 'Times New Roman',serif; font-size: 14pt;">Thus far, however, it seems that the Chaetodontidae (butterflyfishes) is closely related to Scatophagidae (27, 28). <span style="font-family: 'Times New Roman',serif; font-size: 14pt;">According to the study done by Fessler & Westneat 28], Scatophagidae appears to be the sister group to Pomacanthidae (marine angelfish), and that this sister-family pair (Scatophagidae and Pomacanthidae) is the clade most closely related to Chaetodontidae (butterflyfishes) (see figure 9 below).

<span style="font-family: 'Times New Roman',serif; font-size: 14pt;">Within Scatophagidae, however, the phylogenetics has yet to be studied 29].




 * <span style="color: #4f7fbd; font-family: 'Times New Roman',Times,serif; font-size: 12pt;">Figure 13. Phylogenetic tree derived for Chaetodontidae by Fessler & Westneat 28] through Bayesian analysis of mtDNA (12S, 16S, and ND3) and nuclear DNA (RAG2, and Tmo-4c4). It could be seen that the sister-family clade of Scatophagidae and Pomacanthidae is sister to the Chaetodontidae. Scatophagidae is boxed in red. (Figure adapted and used with permission from authors Fessler & Westneat, 2007; Permission Pending from journal). **

=<span style="color: #0fad85; font-family: 'Times New Roman',Times,serif; font-size: 130%;">Original description and Type information =

<span style="font-family: 'Times New Roman',serif; font-size: 14pt;">The type locality of //S. argus// is India 30]. <span style="font-family: 'Times New Roman',serif; font-size: 14pt;">//Scatophagus argus// was first named and first described as //Chaetodon argus// by Linnaeus in 1766. <span style="font-family: 'Times New Roman',serif; font-size: 14pt;">The original description of //S. argus// by Linnaeus in 1766 as //<span style="font-family: 'Times New Roman',Times,serif; font-size: 14pt;">Chaetodo ////<span style="font-family: 'Times New Roman',Times,serif; font-size: 14pt;">n argus // <span style="font-family: 'Times New Roman',serif; font-size: 14pt;">was made based on a information and/or drawing given to him. According to Kottelat 30], the description was done based on “information and/or drawing of a specimen in Schlosser's collection communicated by Brünnich (Boddaert, 1770: 16–17) 31]. This specimen is described in detail and figured by Boddaerts (1770)”. It is likely that the holotype does exist however, its exact location is unknown 30].

<span style="font-family: 'Times New Roman',Times,serif; font-size: 14pt;">The original description (figure below) of //S. argus// as //Chaetodon argus// by Linnaeus can be found on page 464 of his book, <span style="font-family: 'Times New Roman',Times,serif; font-size: 14pt;">Linnaeus, C. (1766). Systema naturae sive regna tria naturae, secundum classes, ordines, genera, species, cum characteribus, differentiis, synonymis, locis. Laurentii Salvii, Holmiae. 12th ed. v. 1 (pt 1): 1-532.


 * <span style="color: #4f7fbd; font-family: 'Times New Roman',Times,serif; font-size: 12pt;">Figure 14. A digitised page of Linnaeus' book showing the original description of //S. argus// as //Chaetodon argus// (image from Biodiversity Heritage Library, contributed by Harvard University Botany libraries). **

=<span style="color: #0fad85; font-family: 'Times New Roman',Times,serif; font-size: 130%;">Conservation status and threats =

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//<span style="font-family: 'Times New Roman',serif; font-size: 14pt;">Scatophagus argus //<span style="font-family: 'Times New Roman',serif; font-size: 14pt;"> does not seem to face any risk of extinction. It is not threatened nor endangered in Singapore 4] and is also listed as “Least Concern” on IUCN Red List of Threatened Species 32].

<span style="font-family: 'Times New Roman',serif; font-size: 14pt;">However, potential localized threats might exist given that //S. argus// occur in coastal areas, and is thus subjected to threats such as pollution in estuarine areas, and coastal habitat loss through development activities such as land reclamation (32, 4). In Singapore, coastal developments and over-fishing may affect local populations 4].

=<span style="color: #0fad85; font-family: 'Times New Roman',Times,serif; font-size: 130%;">Acknowledgement =

<span style="font-family: 'Times New Roman',serif; font-size: 12pt;">I would like to express my deepest appreciation to all those who provided me the possibility to complete this species page: <span style="font-family: 'Times New Roman',serif; font-size: 12pt;">Professor Rudolph Meier for his guidance on how a proper species page should be done, and for the opportunity to create a species page for a Singaporean species that would contribute to the growing knowledge of wildlife in Singapore. <span style="font-family: 'Times New Roman',serif; font-size: 12pt;">The many kind professors who granted me permission to re-use the images of their published papers. <span style="font-family: 'Times New Roman',serif; font-size: 12pt;">Ms Lee Bee Yan for her guidance on taxonomy and phylogenetics, and Mr Low Bi Wei and Dr Jeffrey Kwik for sharing information on the spotted scat.

<span style="font-family: 'Times New Roman',Times,serif; font-size: 12pt;">Kwik, Jeffrey Teik Being, Post-doctoral Research Fellow, Freshwater and Invasion Biology Lab, Department of Biological Sciences, National University of Singapore
 * <span style="font-family: 'Times New Roman',Times,serif; font-size: 14pt;">Personal Communications **

=<span style="font-family: 'Times New Roman',Times,serif; font-size: 14pt;">References = <span style="font-family: 'Times New Roman',Times,serif; font-size: 14pt;">1) <span style="font-family: 'Times New Roman',serif; font-size: 12pt;">Tan, L. W. H. & P. K. L. Ng, 1988. A Guide to Seashore Life. Singapore Science Centre, 160pp.

<span style="font-family: 'Times New Roman',serif; font-size: 12pt;">2) Allen, G.R., 1984. Scatophagidae. In: Fischer, W. & G. Bianchi (eds.), //FAO species identification sheets for fishery purposes. Western Indian Ocean (Fishing Area 51).// volume 4. Food and Agriculture Organisation, Rome. 4 pp.

<span style="font-family: 'Times New Roman',serif; font-size: 12pt;">3) <span style="font-family: 'Times New Roman',serif; font-size: 16px;">Sivan, G & C. Radhakrishnan, 2007. Studies on Venom of the fish //<span style="font-family: 'Times New Roman',serif; font-size: 16px;">Scatophagus argus //<span style="font-family: 'Times New Roman',serif; font-size: 16px;"> from Cochin estuary- a biochemical approach, PhD Dissertation, Cochin University of Science and Technology.

<span style="font-family: 'Times New Roman',serif; font-size: 12pt;">4) Tan, R., 2013a. Spotted scat, //Scatophagus argus//. Family Scatophagidae. Wild Factsheet, //Wild Singapore//. URL: [] (accessed on 1 November 2014).

<span style="font-family: 'Times New Roman',serif; font-size: 16px; line-height: 24px;">5) Romero, P **<span style="font-family: 'Times New Roman',serif; font-size: 16px; line-height: 24px;">. **<span style="font-family: 'Times New Roman',serif; font-size: 16px; line-height: 24px;">, 2002. An etymological dictionary of taxonomy. Madrid, unpublished. Accessible at: [] (Accessed on September 2014)

<span style="font-family: 'Times New Roman',serif; font-size: 12pt;">6) Kottelat, M., 2001. Scatophagidae. Scats. In: K.E. Carpenter & V. Niem (eds.), //FAO species identification guide for fishery purposes. The living marine resources of the Western Central Pacific//. Vol. 6. Bony fishes part 4 (Labridae to Latimeriidae), estuarine crocodiles. Food and Agriculture Organisation, Rome. Pp. 3623-3626. Link]

<span style="font-family: 'Times New Roman',serif; font-size: 12pt;">7) <span style="font-family: 'Times New Roman',serif; font-size: 16px; line-height: 24px;">Barry, T. P. & Fast, A. W, 1988. Introductions. In: Fast, A. W (ed.), //<span style="font-family: 'Times New Roman',serif; font-size: 16px; line-height: 24px;">Spawning induction and pond culture of the spotted scat (Scatophagus argus Linnaeus) in the Philippines. //<span style="font-family: 'Times New Roman',serif; font-size: 16px; line-height: 24px;"> Mariculture Research and Training Center, Hawaii Institute of Marine Biology, University of Hawaii at Manoa. Pp. 1-3.

8) <span style="font-family: 'Times New Roman',serif; font-size: 12pt; line-height: 1.5;">Barry, T. P. & A. W. Fast., 1992. Biology of the spotted scat ( //<span style="font-family: 'Times New Roman',serif; font-size: 12pt; line-height: 1.5;">Scatophagus argus //<span style="font-family: 'Times New Roman',serif; font-size: 12pt; line-height: 1.5;">) in the Philippines. //<span style="font-family: 'Times New Roman',serif; font-size: 12pt; line-height: 1.5;">Asian Fisheries Science //<span style="font-family: 'Times New Roman',serif; font-size: 12pt; line-height: 1.5;">, **<span style="font-family: 'Times New Roman',serif; font-size: 12pt; line-height: 1.5;">5 **<span style="font-family: 'Times New Roman',serif; font-size: 12pt; line-height: 1.5;"> (2): 163-179.

<span style="font-family: 'Times New Roman',serif; font-size: 16px; line-height: 24px;">9) Parenti, P., 2004. Family Scatophagidae Bleeker 1876 — scats. //<span style="font-family: 'Times New Roman',serif; font-size: 16px; line-height: 24px;">California Academy of Science //<span style="font-family: 'Times New Roman',serif; font-size: 16px; line-height: 24px;">, Annotated Checklists of Fishes No. 36, 5 pp. URL: [] (Accessed on 8 November 2014).

<span style="font-family: 'Times New Roman',serif; font-size: 12pt;">10) Mostarda, E. & R. Anam, 2012. //Field identification guide to the living marine resources of Kenya//. Food and Agriculture Organization, Rome. 357 pp.

<span style="font-family: 'Times New Roman',serif; font-size: 12pt;">11) Tan, R., 2013b. Rabbitfishes, Siganidae. Wild Factsheet, //Wild Singapore//. URL: http://www.wildsingapore.com/wildfacts/vertebrates/fish/siganidae/siganidae.htm (accessed on 12 November 2014).

<span style="font-family: 'Times New Roman',serif; font-size: 12pt;">12) Shipp, R. L., 1978. Tetraodontidae. In: Fischer, W. (ed.), //FAO species identification sheets for fishery purposes. West Atlantic (Fishing Area 31).// volume 5. Food and Agriculture Organisation, Rome, Rome. Pp 1988- 1998.

<span style="font-family: 'Times New Roman',Times,serif; font-size: 16px; line-height: 24px;">13) Dekkers, W. J, 1975. Review of the Asiatic freshwater puffers of the genus Tetraodon Linnaeus, 1758 (Pisces, Tetraodontiformes, Tetraodontidae). Bijdragen tot de Dierkunde. **<span style="font-family: 'Times New Roman',Times,serif; font-size: 16px; line-height: 24px;">45 **<span style="font-family: 'Times New Roman',Times,serif; font-size: 16px; line-height: 24px;"> (1): 87-142.

<span style="font-family: 'Times New Roman',serif; font-size: 12pt;">14) Fischer, W. (ed.), 1978. //FAO species identification sheets for fishery purposes. West Atlantic (Fishing Area 31).// volume 5. Food and Agriculture Organisation, Rome, Rome. Pp 1988- 1998.

<span style="font-family: 'Times New Roman',serif; font-size: 16px; line-height: 24px;">15) <span style="font-family: 'Times New Roman',serif; font-size: 16px; line-height: 24px;"> Allen, G.R. & M.V. Erdmann, 2012. Reef fishes of the East Indies. Tropical Reef Research, Perth, Australia. Volumes I-III. University of Hawai’i Press, Honolulu. 1292 pp.

<span style="font-family: 'Times New Roman',serif; font-size: 16px; line-height: 24px;">16) Sivan, G. & C. Radhakrishnan, 2011. Food, feeding habits and biochemical composition of //Scatophagus argus//. //<span style="font-family: 'Times New Roman',serif; font-size: 16px; line-height: 24px;">Turkish Journal of Fisheries and Aquatic Sciences //<span style="font-family: 'Times New Roman',serif; font-size: 16px; line-height: 24px;">, **<span style="font-family: 'Times New Roman',serif; font-size: 16px; line-height: 24px;">11 **<span style="font-family: 'Times New Roman',serif; font-size: 16px; line-height: 24px;">: 603-608.

<span style="font-family: 'Times New Roman',serif; font-size: 16px; line-height: 24px;">17) Cai, Z. P., Y. Wang, J. W. Hu, J. B. Zhang & Y. G. Lin, 2010. Reproductive biology of //<span style="font-family: 'Times New Roman',serif; font-size: 16px; line-height: 24px;">Scatophagus argus //<span style="font-family: 'Times New Roman',serif; font-size: 16px; line-height: 24px;"> and artificial induction of Spawning. //<span style="font-family: 'Times New Roman',serif; font-size: 16px; line-height: 24px;">Journal of Tropical Oceanography //<span style="font-family: 'Times New Roman',serif; font-size: 16px; line-height: 24px;">, **<span style="font-family: 'Times New Roman',serif; font-size: 16px; line-height: 24px;">29 **<span style="font-family: 'Times New Roman',serif; font-size: 16px; line-height: 24px;"> (5): 180- 185.

<span style="font-family: 'Times New Roman',serif; font-size: 16px; line-height: 24px;">18) Winfree, R. A., 1986. Progress in the population of valuable aquarium fishes: First recorded breeding of //<span style="font-family: 'Times New Roman',serif; font-size: 16px; line-height: 24px;">Scatophagus argus //<span style="font-family: 'Times New Roman',serif; font-size: 16px; line-height: 24px;"> Presented at the 17th Annual meeting of World Mariculture Society, 17-23 January, Reno, NV, USA.

<span style="font-family: 'Times New Roman',serif; font-size: 16px; line-height: 24px;">19) Ghaziloua, A., Chenarya, F., Morovvatib, H., Zolgarneinea, H., 2011. Time course of saltwater adaptation in spotted scat ( //<span style="font-family: 'Times New Roman',serif; font-size: 16px; line-height: 24px;">Scatophagus argus //<span style="font-family: 'Times New Roman',serif; font-size: 16px; line-height: 24px;">) (Pisces): a histomorphometric approach. //<span style="font-family: 'Times New Roman',serif; font-size: 16px; line-height: 24px;">Italian Journal of Zoology //<span style="font-family: 'Times New Roman',serif; font-size: 16px; line-height: 24px;">. **<span style="font-family: 'Times New Roman',serif; font-size: 16px; line-height: 24px;">78 **<span style="font-family: 'Times New Roman',serif; font-size: 16px; line-height: 24px;"> (1): 82–89.

<span style="font-family: 'Times New Roman',serif; font-size: 16px; line-height: 24px;">20) Marshall, T.C., 1964. //<span style="font-family: 'Times New Roman',serif; font-size: 16px; line-height: 24px;">Fish of the Great Barrier Reef and Coastal Waters of Queensland //<span style="font-family: 'Times New Roman',serif; font-size: 16px; line-height: 24px;">. Livingston publishing Co., Sydney. 566 pp.

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<span style="font-family: 'Times New Roman',serif; font-size: 12pt;">22) Cameron, A. M. & R. Endean., 1977. Venom glands in Scatophagid fish. //Toxicon//, **8** (2): 171–178.

<span style="font-family: 'Times New Roman',serif; font-size: 12pt; line-height: 1.5;">23) Isbister, G. K., 2004 //<span style="font-family: 'Times New Roman',serif; font-size: 12pt; line-height: 1.5;">Marine envenomation and poisoning. //<span style="font-family: 'Times New Roman',serif; font-size: 12pt; line-height: 1.5;"> In: Dart, R. C. (ed.), //<span style="font-family: 'Times New Roman',serif; font-size: 12pt; line-height: 1.5;">Medical toxicology. //<span style="font-family: 'Times New Roman',serif; font-size: 12pt; line-height: 1.5;"> 3rd ed. Williams & Wilkins, Lippincott. Pp. 1621- 1644.

<span style="font-family: 'Times New Roman',serif; font-size: 16px; line-height: 24px;">24) Lim, K. K. & Low J. K., 1998. //<span style="font-family: 'Times New Roman',serif; font-size: 16px; line-height: 24px;">A guide to common marine fishes of Singapore //<span style="font-family: 'Times New Roman',serif; font-size: 16px; line-height: 24px;">. Singapore Science Centre, Singapore. 163 pp.

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<span style="font-family: 'Times New Roman',serif; font-size: 12pt;">27) Bellwood, D.R. & L. van Herwerden & N. Konow, 2004. Evolution and biogeography of marine angelfishes (Pisces: Pomacanthidae). //Molecular Phylogenetics and Evolution//, **33** (1): 140–155.

<span style="font-family: 'Times New Roman',serif; font-size: 16px; line-height: 24px;">28) Fessler, J. L. & M. W. Westneat, 2007. Molecular phylogenetics of the butterflyfishes (Chaetodontidae): Taxonomy and biogeography of a global coral reef fish family. //<span style="font-family: 'Times New Roman',serif; font-size: 16px; line-height: 24px;">Molecular Phylogenetics and Evolution //<span style="font-family: 'Times New Roman',serif; font-size: 16px; line-height: 24px;">, **<span style="font-family: 'Times New Roman',serif; font-size: 16px; line-height: 24px;">45 **<span style="font-family: 'Times New Roman',serif; font-size: 16px; line-height: 24px;"> (1): 50-68.

<span style="font-family: 'Times New Roman',serif; font-size: 12pt;"><span style="font-family: 'Times New Roman',Times,serif;">29) Chen, Jianhua, J. Chen, Y. Li, M. He, B. Yan, & X. Meng, 2013. Complete mitochondrial genome of the spotted scat //Scatophagus argus// (Teleostei, Scatophagidae). //<span style="font-family: 'Times New Roman',Times,serif;">Mitochondrial DNA, //<span style="font-family: 'Times New Roman',Times,serif;"> Early Online //<span style="font-family: 'Times New Roman',Times,serif;">: //<span style="font-family: 'Times New Roman',Times,serif;"> 1-2.

<span style="font-family: 'Times New Roman',serif; font-size: 16px; line-height: 24px;">30) Kottelat, M., 2013. The fishes of the inland waters of Southeast Asia: a catalogue and core bibliography of the fishes known to occur in freshwaters, mangroves and estuaries. //<span style="font-family: 'Times New Roman',serif; font-size: 16px; line-height: 24px;">Raffles Bulletin of Zoology //<span style="font-family: 'Times New Roman',serif; font-size: 16px; line-height: 24px;">, Supplement 27: 1–663.

<span style="font-family: 'Times New Roman',serif; font-size: 12pt;">31) Boddaert, P., 1770. Epistola ad virum Johannem Burmannum […] de chaetodonte argo descripto atque accuratissima icone illustrato ex museo viri celeberrimi Johannis Alberti Schlosseri. Brief […] aan den weledelen hooggeleerden Heere Johannes Burmannus […] behelzende eene nauuwkeurige beschrijving, en naar het leven vervaardigde afbeelding, van den gevlakten klipvisch, uit de verzameling van wylen den weledelen zeer geleerden Heere Johannes Albertus Schlosser in zijn weled. Van Tongerlo, Amsterdam, 43 pp., pl. 2.

<span style="font-family: 'Times New Roman',serif; font-size: 12pt;">32) Collen, B., N. Richman, A. Beresford, A. Chenery & M. Ram (Sampled Red List Index Coordinating Team), 2010. //Scatophagus argus//. The IUCN Red List of Threatened Species. Version 2014.2. URL: [] . (Accessed on 8 November 2014).

<span style="font-family: 'Times New Roman',serif; font-size: 16px; line-height: 24px;">33) Mahmud, Y., O. Arakawa, A. Ichinose, M. B. Tanu, T. Takatani, K. Tsuruda, K. Kawatsu, Y. Hamano & T. Noguchi, 2003. Intracellular visualization of tetrodotoxin (TTX) in the skin of a puffer //<span style="font-family: 'Times New Roman',serif; font-size: 16px; line-height: 24px;">Tetraodon nigroviridis //<span style="font-family: 'Times New Roman',serif; font-size: 16px; line-height: 24px;"> by immunoenzymatic technique. //<span style="font-family: 'Times New Roman',serif; font-size: 16px; line-height: 24px;">Toxicon //<span style="font-family: 'Times New Roman',serif; font-size: 16px; line-height: 24px;">, **<span style="font-family: 'Times New Roman',serif; font-size: 16px; line-height: 24px;">41 **<span style="font-family: 'Times New Roman',serif; font-size: 16px; line-height: 24px;">(5): 605-611.

<span style="font-family: 'Times New Roman',serif; font-size: 12pt;">34) Computer Generated Native Distribution Map for //Scatophagus argus// (Spotted scat) (modelled future range map based on IPCC A2 emissions scenario). URL: [|www.aquamaps.org], version of Aug. 2013. Web. (Accessed 9 November 2014).

<span style="font-family: 'Times New Roman',serif; font-size: 12pt;">35) De Bruin, G.H.P., B.C. Russell & A. Bogusch, 1995. FAO species identification field guide for fishery purposes. The marine fishery resources of Sri Lanka. Food and Agriculture Organisation, Rome. 400 p.

<span style="font-family: 'Times New Roman',serif; font-size: 12pt; line-height: 1.5;">36) <span style="font-family: 'Times New Roman',serif; font-size: 16px; line-height: 24px;">Ng, P.K.L. & N. Sivasothi (eds.), 1999. //<span style="font-family: 'Times New Roman',serif; font-size: 16px; line-height: 24px;">A guide to the mangroves of Singapore I: The Ecosystem and plant diversity //<span style="font-family: 'Times New Roman',serif; font-size: 16px; line-height: 24px;">. Singapore Science Centre, Singapore. 160pp.

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