Biosaintifika 11 (3) (2019) 325-331 Biosaintifika Journal of Biology & Biology Education

http://journal.unnes.ac.id/nju/index.php/biosaintifika

Phylogenetic Relationships among Ornamental Achanturid Fish from Ujunggenteng and Taman Manalusu, West Java

Afra Nabila, Agus Nuryanto, Romanus Edy Prabowo, Dian Bhagawati, Kusbiyanto

DOI: http://dx.doi.org/10.15294/biosaintifika.v11i3.20671

Faculty of Biology, Universitas Jenderal Soedirman, Indonesia

History Article Abstract Submitted 20 August 2019 Acanthuridae is a marine fish family with some members are known as ornamental Revised 19 September 2019 species. Previous studies has been reported the diversity of Acanthuridae. How- Accepted 9 December 2019 ever, there was no study on the phylogenetic relationship of the Acanthuridae from Ujunggenteng and Taman Manalusu southern coast of West Java. The purpose of Keywords this study was to provide information about the phylogenetic relationship among Acanthuridae; Phylogenetic ornamental fishes under the Acanthuridae family from Ujunggenteng Beach Su- Relationship; Taman Manalu- su; Ujunggenteng; West Java kabumi and Taman Manalusu Beach Garut, West Java. The phylogenetic tree was reconstructed based on morphological characters using maximum parsimony algorithm in software PAUP with 1000 pseudoreplicates and with Halichoeres marginatus selected as an outgroup species. The phylogenetic tree had a consistency index of 0.6429. The value means that the tree was highly reliable due to its low homoplasy. Acanthuridae formed a monophyletic clade compared to the outgroup species by having seven synapomorphic characters. Acanthurid clade was divided into two subclades at generic levels by nine derived characters; three apomorphic characters in Acanthurus and six apomorphic characters in Naso. The tree indicates that Naso brevirostris and Naso lituratus represents basal group, while all species within genus Acanthurus are the most advanced species. This study is the first to provide the data about the evolutionary relationship of ornamental acanthurid fish from the southern coast of West Java. Data on closely related species is one of the essential scientific basis on making policies regarding the sustainable use of those closely related species.

How to Cite Nabila, A., Nuryanto, A., Prabowo, R., Bhagawati, D., & Kusbiyanto, K. (2019). Phylogenetic Relationships among Ornamental Achanturid Fish from Ujunggen-

 Correspondence Author: p-ISSN 2085-191X Jl. dr. Soeparno 63 Purwokerto 53122 e-ISSN 2338-7610 E-mail: [email protected] Afra Nabila et al. / Biosaintifika 11 (3) (2019) 325-331

INTRODUCTION Indonesia, namely Prionurus chrysurus (Muse- um Zoologicum Bogoriense, MZB). Bernal and Indonesia’s waters host higher marine or- Rocha (2011) focused on the relationships among namental fishes than the other countries do (Wa- species within Acanthurus from Atlantic, while hyudin, 2011). These marine resources are distri- Ho et al. (2011) emphasized their study in the re- buted to all marine ecosystems all over Indonesia lationships of species under genus Naso. Moreo- (Nijman, 2010). South coast of West Java is one ver, several studies on marine fish had conducted of Indonesian offshore areas with high potenti- in Indonesia (Fadli et al., 2018; Sahetapy et al., als of marine ornamental fish (Wahyudin, 2011; 2018; Hastuty et al., 2014; Sugianti & Mujianto, Djuwendah et al. 2013). Both previous studies 2013). There was no study available on the rela- from Wahyudin (2011) and Djuwendah et al. tionships among species under Family Acanthu- (2013) reported the potential of marine ornamen- ridae from Indonesia, especially on ornamental tal fish in the southern coast of Sukabumi and Acanthuridae from the southern coast of West Garut Regencies. Java. Therefore, it is important to study phylo- Acanthuridae is one of the marine fish fa- genetic relationship among ornamental Acant- milies whose members are referred to ornamental huridae from southern coast of West Java. Data species from Sukabumi (Wahyudin, 2011), whi- on relationships are essential when management le, there were no taxonomic data available from policies are needed for certain species, while no Southern Garut (Djuwendah et al., 2013). The data are available for that target species. Data of member of this family is characterized by having closely related species can be used as preliminary deeply flattened body shape, caudal peduncle spi- information because they generally have similar nes, small and terminal mouth, and small ctenoid characteristics. scales (Allen & Erdman, 2012). Some species The study aimed to provide information have various colorations as well (Allen & Erd- about the evolutionary relationships among or- man, 2012). namental fish species within Acanthuridae from The family of Acanthuridae consists of six Ujunggenteng, and Taman Manalusu offshores genera; Acanthurus, Ctenochaetus, Naso, Paracant- of southern coast of West Java. The results of this hurus, Prionurus, and Zebrasoma. Acanthurus is the study was expected to provide evidence for other most prominent genus in Acanthuridae and wide- taxonomists that morphological characteristics ly distributed across Indo-Pacific regions. Acant- are reliable characters for phylogenetic analysis hurus consisted of 83 described species (Bernal & of fish. Besides, the information on relationships Rocha, 2011), which are divided into six genera among the members of Acanthuridae from sout- (Froese & Pauly, 2019). hern coast of West Java is a vital scientific basis High species diversity in Acanthuridae is for manager or the authority on making policy interested to be studied, with particular empha- for sustainable uses of that fish resources because sis on their relationship. The relationships among closely related species can be managed together species within Acanthuridae can be inferred as since those species have similar characteristics. evolutionary (cladistics) relationships based on morphological (Klingenberg & Gidaszewski, METHODS 2010) and molecular characters (Carpenter et al., 2017; Bernal & Rocha 2011). Cladistics is con- Sampling sites and samples preservation sidered as a better method than phenetics. It is Fish samples were bought from the first because the cladistics can minimize the use of level middlemen in Ujung Genteng Sukabumi homoplastic characters through careful charac- Regency and Taman Manalusu Garut Regency ter selection (Klingenberg & Gidaszewski, 2010). southern coast of West Java (Figure 1). Sampling Homoplasy is a similarity of a particular charac- was conducted in February and March 2018 with ter in two or more lineages (Wake et al., 2011). four-time repetitions. Fresh fish samples were Previous studies had reported the relation- photographed directly before wet preservation. ships among members of Achanthuridae (Car- All samples were preserved in ethanol 70%. penter et al., 2017; Ludt et al., 2015; Sorensen et al., 2013; Bernal et al., 2011; Ho et al., 2011). Parameter measurement Carpenter et al. (2017) studied phylogenetic rela- A total of 30 morphological characters had tionships among Acanthurus from the Philippine. used for phylogenetic analysis consisted of gene- Ludt et al. (2015) studied relationships among ral morphological performances, meristic, and the members of Acanthuridae from all over the morphometric characters. Observation of general World, but only one species was collected from morphological performance had done manually

326 Afra Nabila et al. / Biosaintifika 11 (3) (2019) 325-331 to describe a specific character in each species. was started by character selection from Operatio- Parameter for general morphology includes dots nal Taxonomy Unit (OUT). The observed morpho- in the face and black streak behind the eye. The logical characters were transformed into multi-sta- other general morphological parameters were the te characters which were symbolized by 0, 1, and 2. horizontal line in the lateral of body, white band The symbol 0 referred to as characters of outgroup at caudal peduncle, white dots at the body, and species. The symbols 1 and 2 referred as in group caudal peduncle spines. Meristic characters had characters. Matrices data saved in nexus file type also counted manually to obtain data on soft fin were consisted of taxa block, character block, rays and dorsal and caudal fin spine. Soft fin rays and PAUP block. Phylogenetic tree reconstructi- observation had conducted with the help of mag- on was conducted based on maximum parsimo- nification lens. The morphometric parameters ny algorithm with 1000 bootstrap replications in consisted of a length of anal fin base, and length PAUP 4.0 (Swofford, 2001). Branching polarity of the anal fin. Additional parameters were the was obtained from the out-group comparison. length of dorsal-fin base, length of the dorsal fin, Halichoeres marginatus had referred as outgroup length of pectoral fin base, length of pectoral fin, species due to its monophyly with Acanthuridae. length of ventral fin base, length of the ventral fin, and length of the caudal fin. The remaining RESULTS AND DISCUSSION parameters for general morphology were the ratio between total length and standard length, Evolutionary Relationships the ratio between head length and snout length, Phylogenetic relationships among acant- the ratio between head length and eye diameter, hurid ornamental fish based on 30 characters are ratio between head length and jaw length, ratio presented in Figure 2. between standard length and caudal fin length, and ratio between standard length and ventral fin length. The morphometric characters had measu- red using caliper with the accuracy of 0.01 cm. Taxonomic identification referred to Allen and Erdman (2012) and Froese and Pauly (2019). The parameters for the reliability of the phylogenetic tree were Consistency Index (CI) and character changes.

Figure 2. Phylogenetic tree showing evolutionary relationships among acanthurid ornamental fish from Ujunggenteng and Taman Manalusu West Java (number inside the box indicates bootstrap values; letter in the branching point indicates the number of nodes)

The phylogenetic tree in Figure 2 had a consistency index (CI) of 0.6429. The value means that less homoplasy occurred during the analysis. The CI value is close to 1. It indicates that the tree have reliable branching pattern or the evolutionary tree is closely similar to the nature of evolutionary relationships of acanthurid species. Ac- cording to Ucu (2016), the CI value indicates the Figure 1. Sampling sites level of homoplasy. The CI value that is close or similar to 1 prove that the level of homoplasy is Tree reconstruction low or absent and indicates that the tree is highly Phylogenetic relationships among marine reliable. Similar results of a low level of homop- ornamental fish within Acanthuridae was infer- lasy on phylogenetic analysis using morphomet- red from a phylogenetic tree. Tree reconstruction ric characters was also observed by Klingenberg

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& Gidaszewski (2010) on Drosophila melanogaster studies (Nurk and Blattner, 2010). The separation and Klussmann-Fricke et al. (2012) in Scorpion. between Naso and Acanthurus was also reported Homoplasy indicates convergent evolution by Sorenson et al. (2013) which showed in phy- between two or more lineages. In this situation, logenetic tree recosntructed based on molecular two or more different species are highly similar in marker. a particular character. According to Klingenberg The tree in Figure 2 provides information and Gidaszewski (2010), homoplasy results in that Naso brevirostris and Naso lituratus are as a ba- analog characters among lineages. It had further sal group or primitive group of species among all explained by Grandjean et al. (2017) that homop- acanthurid species within a monophyletic group lasy is the appearance of a particular or identical (ingroup). Acanthurus lineatus, A. guttaus, A. nigri- character within two or more lineages which oc- cauda, A. nigrofuscus, A. xanthopterus, A. nigricans, currs independently. Acanthurus sp., A. triostegus, dan A. Maculiceps are It seems that not all nodes are supported referred as a derived group. It means that all spe- by bootstrap values. It could be because the cre- cies in genus Acanthurus is suggested to have more ation of each node was based on different cha- advance characters compared to the basal group. racter number. According to Song et al. (2015), Two groups of sister taxa had formed in bootstrap is value profoundly affected by charac- the phylogenetic tree in Figure 2. The first group ter number. It was also observed that some bran- of sister taxa consisted of Acanthurus guttatus and ches were supported by high bootstrap values. A. nigricauda. The second group of sister taxa was The nodes U, P, O, and M were supported by a formed by Acanthurus triostegus and A. maculiceps. bootstrap value of 90%, 57%, 74%, and 55% res- The four species formed two groups of sister taxa pectively. These high bootstrap supports indicated (A. guttatus-A. nigricauda and A.triostegus-A. ma- that the formation of nodes U, P, O, and M were culiceps) because they shared the most common appropriate and also resulted in a reliable tree to- ancestor plesiomorphic characters (symplesio- pology. High bootstrap value was also observed morphic characters), respectively. According to in many earlier studies in broad range animals, Mitchell et al. (2014), Sister taxa is a group of such as in fish (Nuryanto et al., 2018, 2017), in organisms originated from a single most common crustacea (Hernawati et al., 2013), in Ovis (Rezaei ancestor so that they are closely related one anot- et al., 2010), and in Prenolepis, Hymenoptera (La- her. Moreover, they are separated by having their polla et al., 2010). apomorphich characters. It can be seen in the Figure 2 that Ha- lichoeres marginatus was formed a basal taxon com- Character alteration pared to the ingroup specimens. That condition Acanthuridae is a monophyletic clade of proved that H. marginatus possessed plesiomor- in group samples separated from H. marginatus phic or primitive characters. De la Torre-Barcena as the outgroup species. In this case, Halichoeres et al. (2009) explained that outgroup taxa are marginatus is assumed as plesiomorphic with pri- expected to have primitive characters that can be mitive characters. The members of ornamental used to trace the alteration or evolution of cha- Acanthurid in this study share seven apomorphic racters within ingroup taxa. A detail observation characters; hard dorsal fin rays (5), hard anal fin on the tree (Figure 2) shows that Acanthuridae rays (16), soft anal fins rays (17), soft pectoral fins forms a monophyletic group or clade compared rays (18), soft ventral fins rays (19), number of to the H. marginatus. The monophyly of Acanthu- caudal peduncle spines (22), caudal peduncle spi- ridae indicated that the choice of H. marginatus as ne movement (30). The characters that separate an outgroup species was reliable. According to de Labridae (H. marginatus) and Acanthuridae are la Torre-Barcena et al. (2009), out-group species available in Figure 3. affects reliability of branch topology of a phylo- The subclade Naso (Naso brevirostris and genetic tree. Naso lituratus) and subclade Acanthurus are se- Clade and subclade formation by separa- parated by nine characters (Figure 3). Subclade ting acanthurid from H. marginatus and Acant- Naso possesses six derived characters which were hurus from Naso indicates that the cladistic clas- different from Acanthurus, which have three apo- sification supported conventional or Linnaean morphic characters. The derived characters of classification about the separation of Acanthuri- subclade Naso were the length of anal fin base dae from Labridae (H. marginatus) and Acanthurus (1), anal fin length (2), the number of caudal from Naso. A similar result of cladistic classifica- peduncle spine (22), the first hard dorsal fin ray tion and traditional morphological classification (25), protruded snout (26), and white spots on (Linnaean classification) was observed in other the body (29). The synapomorphic characters

328 Afra Nabila et al. / Biosaintifika 11 (3) (2019) 325-331 of subclade Acanthurus were the ratio between 21 (vertical line present or absent), and number total length and standard length (9); hard dorsal 29 (white spots present or absent). A. nigricauda is fin rays (1), and caudal peduncle spine (3). The apomorphic on characters number 1 (base anal fin characters differentiated Naso and Acanthurus is length), number 18 (soft pectoral fin rays), num- presented in Figure 3. ber 20 (soft caudal-fin rays), number 24 (black streak behind eye present or absent), and number 28 (white band on caudal peduncle). More detail character changes is presented in Figure 3. The monophyly of all species under Acanthuridae is due to shared apomorphic characters which evolved from plesiomorphic characters in their ancestor. According to Philippe et al. (2011), a monophyletic group formed by all taxa that evolved from one ancestor. It was stated further by de Pinna et al. (2010) and Lapolla et al. (2010) that similar characters shared by several taxa might support the monophyly of those taxa. Similar result of the monophyly of taxa that shared similar characters was reported in previous studies (Hernawati et al., 2013; Dimitrov et al., Figure 3. Cladogram showing apomorphy 2012) changes between acanthurid and Halichoeres mar- This study is the first to provide the data ginatus as an out-group and among acanthurid about the evolutionary relationships of ornamen- species (letter at the branching point indicates tal acanthurid fish from the southern coast of nodes code, number above black, and blue ovals West Java and character changes among species indicate the number of characters; number below during evolutionary processes. Information about black and blue ovals indicates character changes). evolutionary relationships of ornamental acanthurid fish from southern coast West Java provides Sister group A. triostegus and A. maculiceps two benefits. First, it supports the development are separated from the remaining species within of animal systematics subject by providing data genus Acanthurus by two synapomorphic charac- for taxonomist or systematist that morphological ters of the remaining Acanthurus; i.e., the ratio characters are potential characters on phyloge- between caudal fin length and standard length netic classification. Secondly, our result also has (13) and soft pectoral fin rays. Both species are practical implication. For instance, if we have two differentiated by characters number 20 and 21 in sister taxa that have different conservation status, A. triostegus and the present of 8 apomorphic cha- we could use the information of taxa with better racters in A. maculiceps. conservation status as a basis for the conservation N. brevirostrisis is apomorphic on the cha- effort of it related taxa. racters number 3 (length of pectoral fin base), number 5 (length of ventral fin base), number CONCLUSION 13 (ratio between standard length and caudal fin length), number 14 (ratio between standard The obtained phylogenetic tree is a reaso- length and ventral fin length), number 19 (soft nable evolutionary tree based on a low level of ventral fin rays), and character number 20 (soft homoplasy. Acanthuridae formed a monophyle- caudal-fin rays). N. lituratus is apomorphic on tic clade compared to the outgroup species and characters number 12 (ratio between head length is divided into two subclades on the genus level. and jaw length), number 16 (hard anal fin rays), The separation of Naso and Acanthurus in the and number 18 (soft pectoral fin rays). It is sho- phylogenetic tree was following Linnaean clas- wed in Figure 3 that nodes either combined by sification. Naso referred to as basal group and synapomorphic characters or separated by apo- Acanthurus as a derived group. The nodes either morphic characters. separated by specific apomorphic or combined by A. guttatus and A. nigricauda as sister taxa synapomorphic characters. were differentiated by their apomorphic characters. A. guttatus is apomorphic on the character ACKNOWLEDGMENT number 7 (caudal fin length), number 10 (ratio between head length and snout length), number We would like to thank the Directorate

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