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Journal of Threatened Taxa Building evidence for conservaton globally www.threatenedtaxa.org ISSN 0974-7907 (Online) | ISSN 0974-7893 (Print) Review

A review about fish walking on land

Arumugam Kumaraguru, Rosete Celsiya Mary & Vijayaraghavalu Saisaraswathi

26 December 2020 | Vol. 12 | No. 17 | Pages: 17276–17286 DOI: 10.11609/jot.6243.12.17.17276-17286

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Threatened Taxa Journal of Threatened Taxa | www.threatenedtaxa.org | 26 December 2020 | 12(17): 17276–17286 ISSN 0974-7907 (Online) | ISSN 0974-7893 (Print) PLATINUM OPEN ACCESS DOI: htps://doi.org/10.11609/jot.6243.12.17.17276-17286

#6243 | Received 29 May 2020 | Final received 10 November 2020 | Finally accepted 02 December 2020

R e v i A review about fsh walking on land e w Arumugam Kumaraguru 1 , Rosete Celsiya Mary 2 & Vijayaraghavalu Saisaraswathi 3

1 Honorary Wildlife Warden, Thiruvarur Forest Division, Tamil Nadu Forest Department, Tamil Nadu 620021, India. 1 Biodiversity Conservaton Foundaton, #360, Indira Gandhi Street, KK.Nagar, Trichy, Tamil Nadu 620021, India. 2 Nature Club, 2,3 School of Advanced Sciences, Vellore Insttute of Technology University, Vellore, Tamil Nadu 632007, India. 1 [email protected], 2 [email protected], 3 [email protected] (corresponding author) பரெவ%&ம()(ம%+கி.ப/(க0) (Mudskipper) நிலந>/ வாழிக0 வA.ைபCசா/Eத, ேசIJ தி%Kக0, சL.MநிலNக0, அைலஏIற பAதிக0, ஆIJ SகTLவாரU மIJU கழிநிலNக0 ேபா)ற பAதிகைள வாWவXடமாக ெகாZடைவ. ஏற\Aைறய 34 ப)Sக.ப%ட பரெவ%&ம() இனNக0 உலகிc காண.பKகி)றன. பரெவ%&ம()/ம%+கி.ப/க0 Abstract:ஆ\ஸுெட/சிேட Mudskippers ேகாபXேட are amphibious (Oxudercidae species Gobiidae) inhabitng AKUபTைத semi-terrestrial ேச/Eதைவ. ecosystems இEேதா like mudfats,-ேமIA mangroves, பசிபX\ பAதி,marshy ஆ.பXp\காவX)swamps, intertdal regions,ெவ.பமZடல and estuaries. ேமIA Around கடIகைர 34 diversifed மIJU species இEதிய.are found across ெபqNகடலிc the globe. Mudskipper இEத உயXpனNகளr) belongs to the Oxudercidae நிகWs பரவலாக\ family and theகாண.பKகிறL. subfamily is Oxudercinae. பரெவ%&ம() The occurrenceஉயXpயc of Aறிகா%&யாகsU, species is vastly found ஆIJ across the SகTLவாரUகளr) Indo-West Pacifc region, பாLகா.M the tropical கZகாணX.பX) western coast of Africa and in the Indian Ocean. Mudskippers are known for being the biological indicator and also an indicator of estuarine safety Aறிகா%&யாகsU அறிய.பKகி)றன. மனrத/களr) பய)பா%&IA ெபpLU உபேயாகபடாததாc, இைவ monitoring. They are used by people for prey-catching baits. This review paper explains the ecological indicators, taxonomy, species diversity,ெபqUபாtU habitat, behavioural இைரைய patern, பX&\AU respiraton uZ&லிc & kinematcs, பய)பKTத.பKகி)றன. feeding ecology, reproducton, இEத nutriton ஆvs\ content க%Kைர, & its medicinal wIJCxழc value, andAறிகா%&க0, threats to mudskippers. வைகபXpTதc, இனNகளr) ப)SகTத)ைம, வாWவXடU, நடTைத Sைற, wவாசU மIJU இய\கவXயc, உணs xழலியc, இன.ெபq\கU, ஊ%டCசTL தரsக0, அத) மqTLவ மதி.M மIJU அபாயNக0 Keywords:ஆகியவIைற Amphibious வXள\AகிறL. fsh, distributon, ecological indicator, mudskippers, species diversity.

பரெவ%&ம()(ம%+கி.ப/(க0) (Mudskipper) நிலந>/ வாழிக0 வA.ைபCசா/Eத, ேசIJ தி%Kக0, சL.MநிலNக0, அைலஏIற பAதிக0, ஆIJ SகTLவாரU மIJU கழிநிலNக0 ேபா)ற பAதிகைள வாWவXடமாக ெகாZடைவ. ஏற\Aைறய 34 ப)Sக.ப%ட பரெவ%&ம() இனNக0 உலகிc காண.பKகி)றன. பரெவ%&ம()/ம%+கி.ப/க0 ஆ\ஸுெட/சிேட ேகாபXேட (Oxudercidae Gobiidae) AKUபTைத ேச/Eதைவ. இEேதா-ேமIA பசிபX\ பAதி, ஆ.பXp\காவX) ெவ.பமZடல ேமIA கடIகைர மIJU இEதிய. ெபqNகடலிc இEத உயXpனNகளr) நிகWs பரவலாக\ காண.பKகிறL. பரெவ%&ம() உயXpயc Aறிகா%&யாகsU, ஆIJ SகTLவாரUகளr) பாLகா.M கZகாணX.பX) Aறிகா%&யாகsU அறிய.பKகி)றன. மனrத/களr) பய)பா%&IA ெபpLU உபேயாகபடாததாc, இைவ ெபqUபாtU இைரைய பX&\AU uZ&லிc பய)பKTத.பKகி)றன. இEத ஆvs\ க%Kைர, wIJCxழc Aறிகா%&க0, வைகபXpTதc, இனNகளr) ப)SகTத)ைம, வாWவXடU, நடTைத Sைற, wவாசU மIJU இய\கவXயc, உணs xழலியc, இன.ெபq\கU, ஊ%டCசTL தரsக0, அத) மqTLவ மதி.M மIJU அபாயNக0 ஆகியவIைற வXள\AகிறL.

பரெவ%&ம()(ம%+கி.ப/(க0) (Mudskipper) நிலந>/ வாழிக0 வA.ைபCசா/Eத, ேசIJ தி%Kக0, சL.MநிலNக0, அைலஏIற பAதிக0, ஆIJ SகTLவாரU மIJU கழிநிலNக0 ேபா)ற பAதிகைள வாWவXடமாக ெகாZடைவ. ஏற\Aைறய 34 ப)Sக.ப%ட பரெவ%&ம() இனNக0 உலகிc காண.பKகி)றன. பரெவ%&ம()/ம%+கி.ப/க0 ஆ\ஸுெட/சிேட ேகாபXேட (Oxudercidae Gobiidae) AKUபTைத ேச/Eதைவ. இEேதா-ேமIA பசிபX\ பAதி, ஆ.பXp\காவX) ெவ.பமZடல ேமIA கடIகைர மIJU இEதிய. ெபqNகடலிc இEத உயXpனNகளr) நிகWs பரவலாக\ காண.பKகிறL. பரெவ%&ம() உயXpயc Aறிகா%&யாகsU, ஆIJ SகTLவாரUகளr) பாLகா.M கZகாணX.பX) Aறிகா%&யாகsU அறிய.பKகி)றன. மனrத/களr) பய)பா%&IA ெபpLU உபேயாகபடாததாc, இைவ ெபqUபாtU இைரைய பX&\AU uZ&லிc Editor: A. Biju Kumar, University of Kerala, Thiruvananthapuram, India. Date of publicaton: 26 December 2020 (online & print) பய)பKTத.பKகி)றன. இEத ஆvs\ க%Kைர, wIJCxழc Aறிகா%&க0, வைகபXpTதc, இனNகளr) ப)SகTத)ைம, வாWவXடU, நடTைத Citaton:Sைற, Kumaraguru, wவாசU மIJU A., R.C. இய\கவXயc,Mary & V. Saisaraswathi உணs xழலியc,(2020). A review இன.ெபq\கU, about fsh walking ஊ%டCசTL on land . தரsக0,Journal of Threatenedஅத) மqTLவ Taxa 12(17): மதி.M 17276–17286 மIJU அபாயNக0. htps://doi. org/10.11609/jot.6243.12.17.ஆகியவIைற வXள\AகிறL.17276-17286

Copyright: © Kumaraguru et al. 2020. Creatve Commons Atributon 4.0 Internatonal License. JoTT allows unrestricted use, reproducton, and distributon of this artcle in any medium by providing adequate credit to the author(s) and the source of publicaton.

Funding: None.

Competng interests: The authors declare no competng interests.

Author details: Dr. A. Kumaraguru is working as conservaton scientst specialized in wildlife biology and genetc studies on tger. Ms. Rosette is currently doing her MSc in biotechnology and her research interests include marine biology, biodiversity and conservaton, phytochemical studies, molecular docking. Dr. V. Sai Saraswathi is an expert in environmental sciences, working for the conservaton of biodiversity, climate change, ethnobotany, pharmacognosy, and phytochemistry.

Author contributon: In this review paper, VS contributed in the literature review collecton, paper editng, proof reading. ER collected literature review, paper drafing. AK prepared the review design, proof reading and editng. All authors were involved in the mansuscript revision.

17276 J TT Review about fsh walking on land Kumaraguru et al.

INTRODUCTION afecton towards salinity and temperature variatons (Kanejiya et al. 2017c). They also contribute towards Osche (1962) suggested that mudskippers would the growth of mangrove trees, when the populaton be useful for interpretng the transiton of species of mudskippers is abundant, consequently providing from water to land. There is some evidence about appropriate nutrients supportng the growth of the transiton of fshes to tetrapods, which occurred mangroves (Shenoy et al. 2012). more than 360 million years ago (Ulrich & Elliot 2013). Polgar (2009) atempted the studies of Malayan There are about 40 species classifed under 10 genera. mudskippers as a biomonitor in mangroves concerning Mudskippers predominantly inhabit the mangrove forest species area relatonship (SAR) and revealed that the and mudfats of the Indo-West Pacifc region, the tropical destructon of habitats resulted from reduced species western coast of Africa, and Indian Ocean coastlines. abundance. Mudskippers bioaccumulate pollutants Mudskippers are known to be burrow dwellers preferring through ingeston as they tend to be prey for many the swampy marshes, estuaries, and the intertdal mudfat species. Mudskippers ensure the coastal, regions for their living (Murdy 1989). The feeding intertdal, and estuary’s health (Polgar & Lim 2011). ecology of the Mudskipper Periophthalmus barbarous Studies have shown that mudskippers act as a biological was found to be ‘opportunistc feeders’ meant to be indicator of addressing the severity of oil polluton in the satsfying the food needs with the available resources coastal areas of the Persian Gulf. Biomarker responses of (Chukwu & Deekae 2013). Generally, mudskippers feed Periophthalmus waltoni were recorded and discovered on algae, detritus, diatoms, nematode, polychaetes and that it acts as the ideal candidate for bioindicator in the eggs of fshes, along with mud and sand partcles (Ravi coastal regions and mudfats. Correspondingly, it can be 2013). utlized for achieving sustainable development (Shirani Since long ago, researchers have explored et al. 2012). mudskipper’s distributon patern, species diversity, Similarly, at Kuwait Bay, Periophthalmus waltoni acts behavioural paterns and their locomotory actvites. as a bioindicator in identfying the bioaccumulaton of In 1989, Murdy studied the morphological features metals (Bu-Olayan & Thomas 2008). It tends to be a of mudskippers, which paved the way for taxonomic feeble indicator at the Persian Gulf marine ecosystem, a classifcaton of mudskippers. Also, he hypothesized the biomonitor candidate-Boleophthalmus dussumieri used relatonship of Oxudericnae gobies with other genera in the monitoring of polycyclic aromatc compounds and groups. Mudskipper’s early developmental age (PAH). It might be various other environmental factors & growth, respiratory & circulatory adaptatons, and which are actng in that ecosystem leading B. dussumieri feeding ecology were also studied by many researchers to be helpless (Sinaei & Mashinchian 2014). Metal across the globe (Jaafar & Murdy 2017). toxicity and accumulaton of metal toxicants like Zn, Cd, Due to the medicinal propertes found in Pb, Cu have suggested that mudskippers are an ideal mudskippers, it gains commercial value either through biomonitor. It was observed that their fns and liver are food consumpton or as traditonal medicine by humans the biomonitors and accumulate metal toxins. Due to (Kanejiya et al. 2017a). Sometmes they are even the contaminants of various metal pollutant, it is not used as bait for catching prey (Gadhavi et al. 2017). A recommended for human consumptons (Ikram et al. few of the vernacular names used in diferent places 2010). for referring to mudskippers are Periophthalmus darwini, Periophthalmus sp., and Periophthalmus Taxonomy novaeguineaensis as sakomo, Periophthalmus as Mudskippers belong to the family Oxudercidae and nebesokera, Periophthalmus weberi as paraguamo, the subfamily Oxudercinae. Some of the species are Periophthalmodon freycinet as genora, Boleophthalmus tabulated in Table 1 and Image 1. caeruleomaculatus as ebanea, Boleophthalmus sp. as pot, Scarpellos histophorus as seekakowea, Oxuderces Species Diversity wirzi as canipo (Polgar & Lim 2011). Mudskippers are Globally, 34 diverse species of mudskippers were known as ‘vaet uluvai’ in Tamil (Ragunathan 2010). recognized, and among them, nine were found in Indian coastlines (Murdy et al. 1989). Nearly fve diversifed Ecological Indicator mudskipper species have been efectvely documented Monitoring the intertdal and estuarine health from Gujarat coastlines (Devendra et al. 2016). In the depending on their abundance mudskippers have direct southeastern coast of India, mudskippers were recorded

Journal of Threatened Taxa | www.threatenedtaxa.org | 26 December 2020 | 12(17): 17276–17286 17277 J TT Review about fsh walking on land Kumaraguru et al.

Table 1. Taxonomic rank, genus, and species of mudskippers around the world with reference to various authors belonging to diferent periods.

Tribe (taxonomic rank) Genus Species References Parapocryptes rictuosus, 1 Oxudercini Parapocryptes Valenciennes 1846 Parapocryptes serperaster Apocryptodon madurensis, 2 Oxudercini Apocryptodon Tomiyama 1934 Apocryptodon punctatus Oxuderces dentatus, 4 Oxudercini Oxuderces Koumans 1938 Oxuderces wirzi 5 Periophthalmini Apocryptes Apocryptes bato Hamilton 1822 Pseudapocryptes borneensis, 6 Periophthalmini Pseudapocryptes Bloch & Schneider 1801 Pseudapocryptes lanceolatus 7 Periophthalmini Zappa Zappa confuentus Roberts 1978 Scartelaos cantoris, Scartelaos gigas, 8 Periophthalmini Scartelaos Chu & Wu 1963 Scartelaos histophorus, Scartelaos tenuis Boleophthalmus hirdsongi, Boleophthalmus boddart, Pallas 1772 9 Periophthalmini Boleophthalmus Boleophthalmus caeruleomaculatus, Boleophthalmus dussumieri, Boleophthalmus pectnirostris, Periophthalmodon freycinet, Valenciennes 1846; 10 Periophthalmini Periophthalmodon Periophthalmodon schlosseri, Pallas 1772 Periophthalmodon septemradiatus, Hamilton 1822 Periophthalmus argentlineatus, Periophthalmus harbarus, Periophthalmus chrysospilos, Periophthalmus gracilis, Periophthalmus kalolo, Valenciennes 1846 Periophthalmus Periophthalmus malaccensis, Hamilton 1822 11 Periophthalmini Periophthalmus minutus, Periophthalmus modestus, Periophthalmus novaeguineaensis, Periophthalmus novemradiatus, Periophthalmus waltoni, Periophthalmus weberi

most during the post-monsoon season, and their Bakkhali River Estuary, Bangladesh (Rahman et al. 2015). preferred habitats are estuarine lands and mangrove Complete phylogeographic studies of Periophthalmus areas. A group of eight species from the Malay Peninsula distributed along Indo-Pacifc region helped to and 12 species from Sumatra is present along the Straits understand its evolutonary history (Polgar et al. 2014). of Malacca (Takita & Ali 1999). Earlier, there were nine Periophthalmus spilotus, a new species of mudskipper, species of mudskippers recorded in peninsular Malaysia was identfed from Sumatra, Indonesia (Murdy & (Polgar 2009). Recent studies have updated the total Takita 1999). Similarly, a new species Parapocryptes count to 17 in peninsular Malaysia (Khaironizam & serperaster has been recorded in peninsular Malaysia Rashid 2005). Nineteen species have been recorded (Khaironizam & Rashid 2000). Periophthalmus recently in the Ramsar site, Johor, Malaysia (Hui et walailakae has been recorded in southeastern India al. 2019). Some of the mudskipper species found (Mahadevan et al. 2019a). There were about 24 newly in Merauke District, Indonesia are Boleophthalmus recorded species during the recent studies in Indonesian boddart, B. pectnirostris, P. takita, P. argentlineatus, waters (Pormansyah et al. 2019). Scartelaos histophorus, and Oxuderces dentatus (Elviana et al. 2019). Periophthalmus waltoni is maximally Living Habitat distributed along the Persian Gulf though there are many Mudskippers inhabit riparian areas with sof and threats to its populaton density (Sharifan et al. 2018). muddy plains. Also, they inhabit where the salinity Though there is a diverse populaton of Periophthalmus level is found to be low and the place rich with benthic barbarus around southeastern Nigeria, it is afected by invertebrates (Baeck et al. 2008). Numerous species overexploitaton (Abiaobo & Udo 2017). People do not were living on the rocky coastline as much, as they occupy consume Periophthalmus novemradiatus as a result of mudfats, mangroves, and sand fats (Gordon et al. which their growth rate is rapidly increasing along the 1968). Rehabilitatng the coastal region with mangrove

17278 Journal of Threatened Taxa | www.threatenedtaxa.org | 26 December 2020 | 12(17): 17276–17286 J TT Review about fsh walking on land Kumaraguru et al.

Image 1. Blue-spoted Mudskipper Boleophthalmus boddart. © A. Kumaraguru

saplings gives a beter habitat for various species, and completely fresh water region of the Mekong River (Mai it was observed that large-sized mudskippers were et al. 2019). Boleophthalmus pectnirostris has shown found in the breakwater (sheltered area for mangroves) behavioural preferences in choosing their microhabitat (Hashim et al. 2010). Pseudapocryptes elongatus is at their early juvenile stage (Chen et al. 2008). To able to tolerate the salinity, and hence it survives in determine the habitat selecton, genomic studies were the open sea, coastal mangroves and inland habitat performed, and also the comparatve analysis among during diferent stages of their growth (Bucholtz et al. diferent species were conducted (Cai 1996). Diferent 2009). Mudskippers alter the environmental conditons mudskippers inhabit diferent microhabitats and improving the growth of young mangroves as they mix have diferent burrow constructon methods (Clayton the soil with detritus (Ravi et al. 2013). Studies related 1993). Habitat selecton is dependent on its ecological to the microhabitat selecton of Chinese mudskippers interactons (Polgar & Crosa 2009). Some species such as identfed their preferences, such as salinity levels, land Periophthalmodon septemradiatus is found to survive in or water; water and air temperatures; light or dark; habitats with low salinity and far from the sea, whereas and various combinatons among them were conveyed B. boddart survives close to the sea with high salinity (Gordon et al. 1985). The major threats to the mudfats (Khaironiazam & Rashid 2003). Mudskippers preferred are soil erosion, macro algae, terrestrializaton, and thick mudfat areas for carrying out their burrowing lack of estuarine water, human interference, and actvity efortlessly (Kanejiya et al. 2017c). Burrows discharge of efuents (Ravi 2012). Periophthalmodon constructed by Bolephthalmus boddart are classifed septemradiatus species is found to be the frst species as follows: burrows with single apertures represent inhabitng and breeding from a saline environment to a newly constructed one, and the other single and double

Journal of Threatened Taxa | www.threatenedtaxa.org | 26 December 2020 | 12(17): 17276–17286 17279 J TT Review about fsh walking on land Kumaraguru et al.

openings ones currently exist. In contrast, the burrows comfortable with a terrestrial lifestyle rather than an with multple apertures represent collapsed ones due to aquatc lifestyle. One among them is Periophthalmodon human interventons (Ravi et al. 2004). schlosseri, which has gill arrangements which are highly adaptable for air-breathing, and they spend less tme in Behavioral patern of mudskipper marine habitats (Takeda et al. 1999). Boleophthalmus Populaton density is inversely proportonal to boddart builds mud walls for two signifcant reasons: the growth rate as the availability of food decreases to avoid hostlity between neighbours and as assistance due to an increased populaton. Boleophthalmus for feeding. Diatoms are the most preferred food for pectnirostris (Blue-spoted Mudskipper) is found to mudskippers. They prefer feeding on mud slopes as a have the longest lifespan; seven and six years in males measure of preventng interventon by their neighbours and females, respectvely (Nanami & Takegaki 2005). (Clayton & Wright 1989). Periophthalmus sobrinus A study on the terrestrial life of mudskippers with prefers to live unaccompanied and rarely lives within Periophthalmus sobrinus showed their survival capacity closed groups. There were large spacings between nests out of water is one and a half days. Surprisingly, there and dark places were preferred for foraging (Gordon et was no afecton in metabolic and heart rates as well al. 1968). The growth rates and life duraton of both as the lactc acid concentraton in blood during their the sexes of Pseudapocryptes elongatus obtained from living out of the water (Gordon et al. 1969). Whereas Sundarbans, India is four-plus years for both the sexes Chinese Mudskipper Periophthalmus cantonensis as the maximum age. Their growth index (Φ) is 4.394 survives for two and a half days out of the water and (males) and 4.503 (females) possessing larger caudal fns observed medium sensitveness of metabolism towards (Mahadevan et al. 2019c). temperature. Starvaton for 9.5 days did not afect the The reason for aggressiveness in Periophthalmus excreton of ammonia but afected excreton of urea modestus is the hypothalamic hormone, arginine- (Gordon et al. 1978). vasotocin (VT) (Nao et al. 2013). The foraging behaviours Boleophthalmus dussumieri has separate exit and in Periophthalmus waltoni are not infuenced by entry for males and females. Juveniles pierce deep environmental factors. They hunt in the same area as into mud during high tdes. Adults neglect to build their prey stays inside the burrow for a longer period chimneys surrounding their holes as the consistency of tme (Clayton & Snowden 2000). The growth of of the soil is between sand and clay (Rathod et al. Bolepthalmus boddart has been recorded high during 2019). Mudskippers growth rate is afected when being their juvenile period, decreasing in successive years exposed to polluton at the embryonic stage (Kruitwagen because of maturaton and spending their energy in et al. 2006). There is a signifcant role played by spawning (Ravi & Rajagopal 2007). aquaporins (integral membrane proteins) for adaptng themselves to the terrestrial lifestyle. The selectve Respiraton and kinematcs changes, like pore formaton and substrate selecton, Anatomical characteristcs of mudskipper gills decide have a substantal contributon to their adaptaton to on adaptatons of their habitat and B. boddart is one an amphibious lifestyle (Lorente-Martnez et al. 2018). that shows excellent adaptaton to aquatc lifestyle. The species Periophthalmodon schlosseri is found to be In comparison, terrestrial adaptatons were favoured an ideal species for aquaculture and more mudskipper more at odds with their amphibious lifestyle (Low et al. studies (Quang 2016). Mudskippers like Boleophthalmus 1988). Since mudskippers appear to change the way boddart constructs mud-walls, for territorial exclusion their skin breathes, their epidermis and skin layers have or spacing, territory, and reduces hostlity. This is been studied in detail (Beon et al. 2012). They adapt based on their abundance. These regional behavioural towards terrestrial lifestyle by secretng mucus, and paterns give beter knowledge about the elastc disc their head containing dense capillary network assists concept of territories (Clayton 1987). Mudfats are vital cutaneous respiraton (Jie et al. 2003). Studies related for the survival of mudskippers but global warming is a to gaseous exchange and their demand for oxygen were serious threat as the mudfats are greatly afected due to done in the intertdal regions (Karen 1993). Ammonia high temperature. Spawning season in Scartelaos gigas excreton happens in mudskippers (Periophthalmodon is from May to July. The growth rate is dependent on schlosseri) through their head. Ammonia gets collected diet, water temperature, and mudfat exposure (Park et in their burrow through the acidifcaton process, al. 2002). preventng them from reverse fuctuaton (Randall et al. Some mudskipper species tend to be more 2004). Mudskippers maintain the air phases according

17280 Journal of Threatened Taxa | www.threatenedtaxa.org | 26 December 2020 | 12(17): 17276–17286 J TT Review about fsh walking on land Kumaraguru et al. to the tde. They can breathe both aquatcally and as escapists (Swanson & Gibb 2004). aerially when there is a high tde, while at low tdes they transfer air into their burrow for breathing (Lee Feeding ecology et al. 2005). Comprehensive research was conducted The mudskipper Pseudapocryptes dentatus follows in Periophthalmus magnuspinnatus on cutaneous herbivorous feeding paterns. Their main order is respiraton and its relatonship with skin layers (Park Diatoms-Bacillariophyceae, green algae and blue-green 2002). Several mudskippers reported evaporatve water algae. Levels in size and metabolism are inversely loss along with their behavioural adaptatons (Dabruzzi proportonal (Sarker et al. 1980). Both plants and animals et al. 2011). Signifcant characteristcs of gills found in were documented while studying the Periophthalmus Periophthalmodon schlosseri, contribute to their ability barbarous stomach. This included mainly crabs, fsh to live inland for a longer period of tme (Wilson et al. scales, and insects. We can recognize from this that 1999). The modifcatons and transformatons that occur mud-skippers are opportunistc feeders and are an in the gill respiratory vasculatures and the mudskippers ideal aquaculture choice (Chukwu & Deekae 2013). of the bucco-opercular cavites were examined using the As described, their key food items include -diatoms, technique of corrosion castng. This has helped to defne nematode, polychaetes, fsh eggs, algae, detritus, along their adaptaton to an amphibious lifestyle (Gonzales with partcles of mud and sand (Ravi 2013). It has et al. 2011). Histological studies using the parafn been shown, according to the study conducted with method have studied the ambiguity in the structure Periophthalmodon schlosseri, that there are diferental of the gills and simultaneously compared their aquatc preferences in the selecton of food products between and terrestrial lifestyles (Supriyat et al. 2019). For their male and female. Females preferred small-scale fsh, survival, conditon-specifc biochemical adaptatons namely Oryzias sp., and males over small-scale fddler occurred during hypoxia, where the glycolysis process is crabs because they were highly involved in land actvites modifed to provide energy during muscle movements, (Zulkifi et al. 2012). and lactate is accumulated (Chew & Ip 1992). The periophthalmus sobrinus feeds on small Contradictory circumstances occur concerning their animals. Their food sources are polychaetes, polydora, adaptatons between Periophthalmodon schlosseri and terebellid, nematodes, crustaceans, copepods, tanaids, Boleophthalmus boddart. Around the same tme, P. prawns, schizopod larvae, alpheid shrimp juveniles, Uca schlosseri tends to be adaptable to a terrestrial lifestyle chlorophthalmus, and tny sand crab (Stebbins & Kalk and is an excellent candidate for aquaculture studies 1961). The discerned food items of Periophthalmus related to air-breathing fsh. B. boddart, however, waltoni are crustaceans (high occurrence), snails appear as opposed to the earlier one (Kok et al. 1998). (slightly lower occurrence), 3.4% insects (lower Mudskipper is an anomaly that stores pre-entry air appearance), and fshes (least) (Mhaisen & Al-maliki inside their burrow and has no metabolism afecton. 2013). Boleophthalmus pectnirostris selects their meal They regularly maintain their air stages and adapt them based on the abundance, availability, size of the diatoms to their amphibious lifestyle (Ishimatsu et al. 1998). The and temperature. The size of the species refects over its essental feature of their adaptaton to terrestrial life is feeding apparatus (Yang et al. 2003). the presence of dermal bulges, thick middle cell layer Boleophthalmus boddart likewise feeds on and a vascularized epidermis (Zhang et al. 2000). In low Bacillariophyta (Quang 2015). The anatomical tde conditons, the intertdal fshes, like mudskippers, characteristcs in Periuphthalmus kuelreuteri are tend to have many optons to choose from. Mudskippers examined by light and X-ray cinematography. It pushes either agree to be an aquatc or temporary terrestrial forward with its pelvic fns when the prey gets near, and living being (Karen 1995). Mudskipper’s locomoton brings the jaws near to the target. Opercular bones, aid has been experimentally tested using water on gelatn swallowing into the pharynx to positon the prey. They and glass along with the assistance of a system for found it easy to capture the prey in the land by bitng and digital image processing. Wang et al. (2013) discovered open mouth. It reaches the stomach via an esophagus through their study that mudskipper uses both body and afer passing the pharyngeal jaws (Sponder & Launder pectoral fns for movement in the water and on gelatn, 1981). whereas they use only pectoral fns for land acton. Escapism is a critcal behaviour that is required for their Reproducton survival. A few studies have shown that, during aquatc Studies were conducted in Nigeria’s lagoon swamps, and terrestrial lifestyles, mudskippers alter their positon which concentrated primarily on sex ratos, egg

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diameters, gonodosomatc index and levels of maturaton consumed. (Lawson 2010). Reports on the growth of eggs with The research by Looi et al. (2016) concluded that artfcial fertlizaton and larval reports in the species the bioaccumulaton of mercury in Periophthalmodon Periophthalmus cantonensis were carried out (Tsuhako et schlosseri did not cause any serious efects when al. 2003). The mudskipper’s eggs were laid deep within consumed. From the evaluaton of the nutrient content the burrows where there is a hypoxic environment, and of Pseudapocryptes elongatus, it has been shown to male mudskippers supply the oxygen by depositng contain sufcient protein, carbohydrate, lipid, important oxygen through water. Once the development of the and non-essental amino acids, polyunsaturated faty eggs is complete, they are released from the burrows acid (PFA) with a greater amount than saturated faty and hatched by tdes, making them prepare themselves acid (SFA). For consumpton it is highly recommended during severity (Ishimatsu et al. 2007). A research on because of its nutritve value (Mahadevan et al. 2019b). Apocryptes bato’s (Gobiidae) reproductve biology Mudskippers are widely available in Nigeria and their in the Payra River, southern Bangladesh, helped to prices are comparatvely low, and they are consumed by understand the basics of reproducton in mudskippers the elderly (Edun et al. 2010). Exposure of mudskipper (Ferdous et al. 2018). Studies related to reproductve Boleophthalmus boddart to natural radionuclides (238U, biology, fertlizaton, maximum sizes achieved along with 226Ra and 210Pb and 210Po), the radionuclide concentraton spawning will allow local fshermen to know the exact was found to be below the limit and therefore, did not tme of catching them, rather than disturb them during have a profound impact in Bombay Harbour and coastal spawning seasons. The concept was prominent from the zone (Bangera & Patel 1984). studies of sex maturaton of Boleophthalmus boddart (Quang et al. 2015). The species Periophthalmodon Threats to mudskippers septemradiatus lays eggs year-round. Observing their The efect on mudskipper density was recorded complete duraton during the maturaton stages helped in the mudfats of Hathab coast, Gujarat, considering to understand reproductve biology and to learn about salinity and temperature as independent variables. the methods of conservaton (Dinh et al. 2018). Studies Accordingly, mudskipper abundance and distributon at intertdal swamps of the Imo River estuary dealt about have become a vital indicator for determining intertdal the reproductve biology of Periophthalmus barbarous region health (Kanejiya et al. 2017c). Post-tsunami along with growth, mortality, recruitment patern, studies at Mudasolodai, Tamil Nadu, revealed that rapid gonadosomatc index, and spawning season (Etm et al. changes in soil morphology led to changes that directly 2002). afected the mudskippers’ livelihood. For constructon, they usually prefer clay-rich soil, rather than sandy soil Studies on nutriton content and medicinal value (Ravi 2005). Although metal uptakes were increasingly Nutrient content of three species Periophthalmus high during lower salinity, the levels of salinity in the waltoni, Boleophthalmus dussumieri, and Scartelaos intertdal mudskipper Periophthalmus cantonensis did histophorus were measured, such as starch, protein, not afect the metal (Cd, Se, & Zn) concentraton factors and lipid. Boleophthalmus dussumieri is rich in nutrients (CF) (Ni et al. 2006). Mudskippers face other menaces and it comes from the liver. Because of this adventtous due to metal toxicity. To illustrate, when exposed to efect, people consume them in the Bhavnagar coast, Cr (Vl), Boleophthalmus dentatus causes afecton in Gujarat (Kanejiya et al. 2017a). Mudskippers are known the actvity of Na+ , K+ and ATPase, and further afects to be very rich in proteins and other nutrients. They membrane actvity, also causing metabolic stress (Kundu are either used in traditonal medicines in countries like et al. 1995). Similarly, the same has also been deduced Malaysia or as bait and also utlized for consumpton. in other studies pertaining to Boleophthalmus dentatus During winter, mudskippers are caught and sold at the obtained from the Gulf of Katch (Lakshmi et al. 1991). market by fshermen using net trap methods (Kanejiya The shrimp efuents collected at the mudskipper et al. 2017b). Likewise, there are growing demands habitat in the northern Persian Gulf have proved to for mudskippers at Narmada estuary, Gujarat, and support mudskipper enrichment, which helps them they are favoured by local people. As a result of this, increase their species density and length. Despite the ecosystem can get highly exploited and can impact severe changes and reduced dissolved oxygen, shrimp biodiversity (Bhakta et al. 2018). Though mudskipper efuents create favourable physical conditons for their meat possess nutritonal benefts there are researchers growth; it survived amid these adversites (Kohan et statng about metal toxicity and bioaccumulaton when al. 2018). Mudskippers are known for their versatle

17282 Journal of Threatened Taxa | www.threatenedtaxa.org | 26 December 2020 | 12(17): 17276–17286 J TT Review about fsh walking on land Kumaraguru et al. behaviour and amphibious character. They have less Applied Ichthyology 29(2): 410–415. understanding of the public and have lost interest. The Bhakta, D., W.A. Meetei, G. Vaisakh, S. Kamble, T.N. Chanu & S.K. Das (2018). Mudskipper Fishery and Indigenous Fishing most striking characteristc of them is their survival Devices in Narmada Estuary, Gujarat. Journal of the Indian ability and adaptaton to diferent environmental Society of Coastal Agricultural Research 36(2): 74–78. Bloch, M.E. & J.G. Schneider (1801). Systema Ichthyologiae Iconibus changes. Surprisingly, mudskippers feed on the available CX Illustratum. Post Obitum Auctoris Opus Inchoatum Absolvit, foodstufs, rather than relying on specifc foods. Correxit, Interpolavit Jo. Gotlob Schneider. Jo. Gotlob Schneider, Further research can be done with them as they defne Berlin, 584pp. Bucholtz, R.H., A.S. Meilvang, T. Cedhagen, J.T. Christensen & D.J. themselves as an ideal candidate for aquaculture. Gills’ Macintosh (2009). Biological Observatons on the Mudskipper anatomical features are specifcally built to suit both the Pseudapocryptes elongatus in the Mekong Delta, Vietnam. Journal terrestrial and aquatc lifestyles. They got adapted from of World Aquaculture Society 40(6): 711–723. Bu-Olayan, A.H. & B.V. Thomas (2008). Trace metals toxicity and the inital stages of egg producton to adjust to severites bioaccumulaton in Mudskipper Periophthalmus waltoni Koumans, such as high tde, and airfow maintenance. 1941 (Gobiidae: Perciformes). Turkish Journal of Fisheries and Aquatc Sciences 8: 215–218. Cai, Z. (1996). Populaton structure and reproductve characteristcs of mudskipper Boleophthalmus pectnirostris, in Shenzhen Bay, China. CONCLUSION Acta Ecologica Sinica 16 (1): 77–82. 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of Mudskipper (Periophthalmus gracilis) Gills. In: Proceeding AcAAJ&oi=fnd&pg=PA1&dq=Histoire+Naturelle+des+Poiss Internatonal Conference on Science and Engineering, Universitas ons&ots=75SD1EbV-3&sig=BGqsMA5Og4ze3gQrCbOykDNl- Islam Negeri Sunan Kalijaga, Indonesia 2: 177–179. 48&redir_esc=y#v=onepage&q=Histoire%20Naturelle%20des%20 Swanson, B.O. & A.C. Gibb (2004). Kinematcs of aquatc and terrestrial Poissons&f=false escape responses in mudskippers. Journal of Experimental Biology Wang, L., M. Xu, B. Liu, T. Jiang, S. Zhang & J. Yang (2013). Experimental 207: 4037–4044. Study on Morphology and Kinematcs of Mudskipper in Amphibious Takeda, T., A. Ishimatsu, S. Oikawa, T. Kanda, Y. Hishida & K.H. Khoo Environments. In: IEEE Internatonal Conference on Robotcs and (1999). Mudskipper Periophthalmodon schlosseri can repay Oxygen Biomimetcs (ROBIO), Shenzhen, China. Date of conference 12–13 debt in air but not in water. Journal of Experimental Zoology 284(3): December 2013. htps://doi.org/10.1109/ROBIO.2013.6739610 265–270. Wilson, J.M., T.W.K. Kok, D.J. Randall, W.A. Vogl & K.Y. Ip (1999). Takita, T. & A.B. Ali (1999). Distributon and habitat requirements Fine structure of the gill epithelium of the terrestrial Mudskipper, of oxudercine gobies (Gobiidae: Oxudercinae) along the Straits of Periophthalmodon schlosseri. Cell and Tissue Research 298(2): 345– Malacca. Ichthyological Research 46: 131–138. 356. Tomiyama, I. (1934). Four new species of gobies from Japan. Journal of Yang, K.Y., S.Y. Lee & G.A. Williams (2003). Selectve feeding by the the Tokyo Imperial University 3(3): 325–334. mudskipper (Boleophthalmus pectnirostris) on the microalgal Tsuhako, Y., A. Ishimatsu, T. Takeda, K.K. Huat & K. Tachihara (2003). assemblage of a tropical mudfat. Marine Biology 143: 245–256. The eggs and larvae of the giant mudskipper, Periophthalmodon htps://doi.org/10.1007/s00227-003-1067-y schlosseri, collected from a mudfat in Penang, Malaysia. Zhang, J., T. Taniguchi, T. Takita & A.B. Ali (2000). On the epidermal Ichthyological Research 50(2): 178–181. structure of Boleophthalmus and Scartelaos mudskippers with Ulrich, K. & J.M. Elliot (2013). Do mudskippers and lungfshes reference to their adaptaton to terrestrial life. Ichthyological elucidate the early evoluton of four-limbed vertebrates?. Evoluton: Research 47: 359–366. htps://doi.org/10.1007/BF02674263 Educaton and Outreach 6(8). htps://doi.org/10.1186/1936-6434- Zulkifi, S.Z., F.M. Yusuf, A. Ismail & N. Miyazaki (2012). Food 6-8. preference of the giant mudskipper Periophthalmodon schlosseri Valenciennes, A. (1846). Natural History of Fish. France. 19. (Teleostei: Gobiidae). Knowledge and Management of Aquatc https://books.google.co.in/books?hl=en&lr=&id=4tBcAAA Ecosystem 405: 10. htps://doi.org/10.1051/kmae/2012013

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PLATINUM The Journal of Threatened Taxa (JoTT) is dedicated to building evidence for conservaton globally by publishing peer-reviewed artcles online every month at a reasonably rapid rate at www.threatenedtaxa.org. OPEN ACCESS All artcles published in JoTT are registered under Creatve Commons Atributon 4.0 Internatonal License unless otherwise mentoned. JoTT allows allows unrestricted use, reproducton, and distributon of artcles in any medium by providing adequate credit to the author(s) and the source of publicaton.

ISSN 0974-7907 (Online) | ISSN 0974-7893 (Print)

December 2020 | Vol. 12 | No. 17 | Pages: 17263–17386 Date of Publicaton: 26 December 2020 (Online & Print) www.threatenedtaxa.org DOI: 10.11609/jot.2020.12.17.17263-17386

Artcle Buterfies (Lepidoptera: Rhopalocera) of the undivided Midnapore District, West Bengal, India: a preliminary report Genetc and reproductve characterizaton of distylous Primula reinii – Anirban Mahata, Niladri Prasad Mishra & Sharat Kumar Palita, in the Hakone volcano, Japan: implicatons for conservaton of the Pp. 17347–17360 rare and endangered plant – Masaya Yamamoto, Honami Sugawara, Kazuhiro Fukushima, Occurrence of Corica soborna Hamilton, 1822 (Clupeiformes: Hiroaki Setoguchi & Kaoruko Kurata, Pp. 17263–17275 Clupeidae) in the Godavari basin, India – Kante Krishna Prasad, Mohammad Younus & Chelmala Srinivasulu, Pp. 17361–17365 Review

A review about fsh walking on land Notes – Arumugam Kumaraguru, Rosete Celsiya Mary & Vijayaraghavalu Saisaraswathi, Pp. 17276–17286 Strobilanthes afnis (Acanthaceae): a new additon to the fora of Manipur, India – Sanjeet Kumar & Rajkumari Supriya Devi, Pp. 17366–17369 Communicatons A new species of the genus Opius Wesmael, 1835 (Hymenoptera: Diversity, distributon and conservaton status of the Adder’s-tongue Braconidae: Opiinae) from Kashmir Himalaya, India ferns in Goa, India – Zaheer Ahmed, Ahmad Samiuddin, Altaf Hussain Mir & – Sachin M. Patl & Kishore Rajput, Pp. 17287–17298 Mohammad Shamim, Pp. 17370–17373

An inventory of the natve fowering plants in East Siang District of Larvae of the blow fy Caiusa testacea (Diptera: Calliphoridae) as egg Arunachal Pradesh, India predators of Polypedates cruciger Blyth, 1852 (Amphibia: Anura: – Momang Taram, Dipankar Borah, Hui Tag & Ritesh Kumar Rhacophoridae) Choudhary, Pp. 17299–17322 – W.G.D. Chathuranga, K. Kariyawasam, Anslem de Silva & W.A. Priyanka P. de Silva, Pp. 17374–17379 Crepuscular huntng of swiflets (Family: Apodidae) by Besra (Family: Accipitridae) in the urban areas of the Andaman Islands, India Blank Swif Caltoris kumara moorei (Evans, 1926) (Lepidoptera: – Amruta Dhamorikar, Dhanusha Kawalkar, Prathamesh Gurjarpadhye Hesperiidae) in Dehradun Valley, Utarakhand, India: a new record & Shirish Manchi, Pp. 17323–17329 for the western Himalaya A study on diversity of mammalian species using camera traps – Arun Pratap Singh, Pp. 17380–17382 and associated vegetaton in Mizoram University Campus, Aizawl, Mizoram First photographic record of the Asiatc Brush-tailed Porcupine – J.H. Zothanpuii, Sushanto Gouda, Abinash Parida & G.S. Solanki, Atherurus macrourus (Linnaeus, 1758) (Mammalia: Rodenta: Pp. 17330–17339 Hystricidae) from the Barak Valley region of Assam, India – Rejoice Gassah & Vijay Anand Ismavel, Pp. 17383–17384

Short Communicatons Book Review Distributon of Syzygium travancoricum Gamble (Myrtaceae), a Critcally Endangered tree species from Kerala part of Western A look over on Red Sanders Ghats, India – S. Suresh Ramanan, Pp. 17385–17386 – V.B. Sreekumar, K.A. Sreejith, M.S. Sanil, M.K. Harinarayanan, M.P. Prejith & R.V. Varma, Pp. 17340–17346

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