Short Communication Journal of Orthoptera Research 2018, 27(2): 155-158 The fishing mantid: predation on fish as a new adaptive strategy for praying mantids (Insecta: Mantodea) IRROBERTO BATTISTON!, RAJESH PUTTASWAMAIAH2Z, NAYAK MANJUNATH? 1 Musei del Canal di Brenta, Palazzo Perli, via Garibaldi 27, 36020, Valstagna (VI), Italy. 2 Bat Conservation India Trust, #285, 10th Cross, Rashi Residency Layout, Phase 1, Kadabgere, Magadi Road, Bengaluru, Karnataka, Pin: 562130, India. 3 Environmental Conservationist, Ron-582209, Gadag, Karnataka, India. Corresponding author: Roberto Battiston (roberto.battiston@ museivalstagna. it) Academic editor: Matan Shelomi | Received 29 June 2018 | Accepted 10 August 2018 | Published 20 September 2018 http://zoobank.org/51E4B318-1DBE-481 F-9835-8 DEED61FA67C Citation: Battiston R, Puttaswamaiah R, Manjunath N (2018) The fishing mantid: predation on fish as a new adaptive strategy for praying mantids (Insecta: Mantodea). Journal of Orthoptera Research 27(2): 155-158. https://doi.org/10.3897/jor.27.28067 Abstract Observations in unmanipulated, semi-natural conditions were made of a single individual of the praying mantid Hierodula tenuidentata, while hunting and capturing an unusual prey for this kind of insect, guppy fish, Poecilia reticulata. This repetitive fishing behavior, recorded daily, is report- ed here for the first time and discussed in relation to the adaptive behav- ioral plasticity of praying mantids. We speculate regarding learning from experience as a hunting strategy. Key words behavior, evolution, India, learning, vision Introduction Mantids can be considered generalist predators but they are known to feed mostly on other insects, especially on fly-like insects (Prete et al. 1999). Vertebrates as prey for mantids are historically con- sidered “anomalous data” based on anecdotal records, mostly from caged or in some way “induced” encounters between mantids and small birds, lizards, frogs, newts, mice, snakes and turtles (reviewed in Kevan 1985, Prete and Wolfe 1992, Costa-Pereira et al. 2010). Several observations in the literature (i.e. Butler 1949, Hilde- brand 1949, Prescott 1968) describe the predation of birds by mantids. However, McCormick and Polis (1982) point out that these observations are inconclusive since the birds captured by mantids were ultimately freed by concerned ornithologists. It is, however, stated that large praying mantids have the ability to kill small birds like hummingbirds and brown creepers. Recently, passerine birds trapped in mist nets have also been reported as occasional prey for mantids (Copete 2006). A recent overview of mantid predation on birds suggests that this behavior may be fre- quent in mantids, having already been recorded for 12 large-sized mantid species (including the genus Hierodula) in 13 different countries world-wide, preying on over 24 species of birds, with a marked preference for hummingbirds attracted to artificial bird- feeders (Nyffeler et al. 2017). Moreover, even if without scientific validation, it is not difficult to find several pictures and reports on the internet of caged (induced or manipulated) encounters/fights between mantids and a variety of small vertebrates. Materials and methods A single, 5.6 cm large male specimen of Hierodula tenuidentata Saussure, 1869, unequivocally recognizable by a white antenno- merus on the right antenna, was occasionally observed in a private roof garden in Karnataka, India, preying on guppy fish, Poecilia reticulata Peters, 1859 (Fig. 1). The predations were observed every day for five consecutive days from the 7" to the 11" of March 2017 during periodical scans of the habitat from sunset at 6:30 pm to 12:30 am. Therefore, they were unmanipulated and occurred natu- rally without interference. About 40 fish [mostly Poecilia reticulata, few Danio rerio Hamilton, 1882 and one Hypostomus plecostomus (Linnaeus, 1758)] were contained in an artificial pond made in an earthen planter of 58 cm diameter, under natural light conditions. The mantid was able to reach the fish by perching on leaves of water lilies (Nymphaeaceae) and water cabbage (genus Pistia Lin- naeus, 1785) growing on the surface of the pond. The garden is located on top of a building about 5 m from the ground and containing about 15 potted plants. Other natural prey for mantids like wasps, butterflies, spiders, etc., have been fre- quently observed on these plants and in the garden which, even if mostly artificial, can be considered a semi-natural and functional habitat for a mantid like Hierodula, which is well-accustomed to gardens and urban areas (Kurosawa et al. 2003, Leong 2009). Results During the five days, the mantid was observed capturing and devouring a total of nine guppy fish. In seven cases, the mantid started eating from the tail (Fig. 2). On a single occasion, he start- ed from the head and on another, from the top side. On the first four of the five days, the mantid was observed to hunt and devour JOURNAL OF ORTHOPTERA RESEARCH 2018, 27(2) R. BATTISTON, R. PUTTASWAMAIAH AND N. MANJUNATH Fig. 1. The artificial pond with the male of Hierodula tenuidentata eating a Poecilia reticulata. Photo by R. Puttaswamaiah. two fish. The second fish was hunted within 10-30 mins of con- suming the first one. After the fifth day, the mantid disappeared and was not observed again at the pond. Discussion Fish do not move like lizards, locusts, hummingbirds or flies. They swim under the usual hunting field of a mantid and are sepa- rated by the barrier of water. Some other invertebrates like spiders, and especially fishing spiders, can hunt in water, but the origin of this adaptation occurred more than once in their evolution (Nyf- feler and Pusey 2014), and is not well understood. Our observa- tions in this fishing mantid — a unique case at the moment - may indicate the origin of this phenomenon. Despite the limitations, our observations raise three important fields for speculation. First, this case confirms that in their natural habitat, mantids can and do feed on vertebrates, even on fish. In this case, of the approximately 40 individual fish present in the pond, nine of them were eaten by the mantid within a span of five days, showing the potential for a single invertebrate to have a strong impact on the fish community and, since guppies, like many other small fish, are active predators of aquatic insects, indi- rectly on the whole pond ecosystem. Second, although the compound eyes of most mantids are ap- positional, sensitive to movement, and adapted to vision mostly in daylight (Kral 2012), this male was able to see and catch the fish under the water at night and to overcome refractive challenges. The fish were caught near the water's surface, always after sunset, some- times late at night and, in general, in low light [compact fluorescent lamp (CFL)]| conditions. Male mantids are known to be very active during the night (Battiston et al. 2010), but these fishing events sug- gest further visual abilities of mantids that should be investigated. Third, the predation was not just occasional, but repeated. This scenario, from a more speculative perspective, suggests the possibility that the insect learned from the experience where and what to hunt. Mantids are known to use aversive learning to avoid toxic prey (Carle et al. 2018), a first basic learning ability shared by many predators. This case, however, suggests a further step to a more articulated cognitive process, including the ability to learn not only from a single stimulus but from different environmental clues and experiences, for elaborate new hunting strategies. Many mantids, including Hierodula, are known to be sit-and- wait predators (Prete et al. 1999), and there is evidence that at least some species carefully choose their habitat and hunting field (Bat- tiston and Fontana 2010, Watanabe et al. 2013, O’Hanlon et al. 2014). Remembering the prey’s abundance at a particular site in relation to their ease of capture and their nutritional content, could be one important factor of this choice and may indirectly influence individual fitness. This should be investigated in further studies. Acknowledgments We are particularly grateful to Prof. Michael Maxell from the National University (California) for his valuable comments and JOURNAL OF ORTHOPTERA RESEARCH 2018, 27(2) R. BATTISTON, R. PUTTASWAMAIAH AND N. MANJUNATH — 157 aia! Fig. 2. Hierodula tenuidentata eating Poecilia reticulata from the tail while the fish is still alive and breathing in the water. Photo by R. Puttaswamaiah. to Arya Rajesh for his assistance in field observations and photog- raphy. We thank the reviewers for their valuable contribution and the Orthopterists’ Society for the support in this manuscript. Conflict of Interest: The authors declare that they have no con- flict of interest. 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Annals of the Entomological Society of America 106: 447-453. https://doi.org/10.1603/AN12145 JOURNAL OF OrRTHOPTERA RESEARCH 2018, 27(2) 158 R. BATTISTON, R. PUTTASWAMAIAH AND N. MANJUNATH Supplementary material 1 Author: Roberto Battiston, Rajesh Puttaswamaiah, Nayak Manju- nath Data type: JPEG file Explanation note: The mantid eating another fish from the head. Copyright notice: This dataset is made available under the Open Database License (http://opendatacommons.org/licenses/ odbl/1.0/). The Open Database License (ODDbL) is a license agreement intended to allow users to freely share, modify, and use this Dataset while maintaining this same freedom for others, provided that the original source and author(s) are credited. Link: https://doi.org/10.3897/jor.27.28067.suppl1 Supplementary material 2 Author: Roberto Battiston, Rajesh Puttaswamaiah, Nayak Manju- nath Data type: JPEG file Explanation note: The mantid resting under a leaf during the day. Copyright notice: This dataset is made available under the Open Database License (http://opendatacommons.org/licenses/ odbl/1.0/). The Open Database License (ODDbL) is a license agreement intended to allow users to freely share, modify, and use this Dataset while maintaining this same freedom for others, provided that the original source and author(s) are credited. Link: https://doi.org/10.3897/jor.27.28067.suppl2 JOURNAL OF ORTHOPTERA RESEARCH 2018, 27(2)