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Analysis of Interspecific and Intraspecific Mediterranean Octopus Behavior Annette Brennan, Stephanie Douglas, Cara Fuller, Samantha Gartner, 2016

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Analysis of Interspecific and Intraspecific Mediterranean Octopus Behavior Annette Brennan, Stephanie Douglas, Cara Fuller, Samantha Gartner, 2016 Analysis of Interspecific and Intraspecific Mediterranean Octopus Behavior Annette Brennan, Stephanie Douglas, Cara Fuller, Samantha Gartner, 2016 ABSTRACT Species coexistence is theorized to affect the behaviors and adaptations of individuals within the same system. For this study, we conducted two sets of trials looking at the foraging behavior and prey preference of three species of octopus, Octopus sautii, Octopus vulgaris, and Callistoctopus macropus in the harbor of the research station, STARESO, in Calvi, Corsica of September and October 2016. In the foraging behavior trials we documented ten different behaviors, where six were found to have significant relationships (p<0.05) with the species or individual. For the prey preference trials, we documented the octopus’ first choice when given three different prey items. A comparison of individual octopus and their prey choice exhibited a significant relationship (p<0.05). From the results, it was concluded that individual personalities of an octopus have greater influence on behavior than species-specific traits. These individual behaviors may lessen competition and lead to coexistence between the three species studied. INTRODUCTION literature documenting how such differences in Coexistence of many species in otherwise like species may be associated with ecosystems has been a perplexing question to coexistence. Coexistence of octopus has already ecologists for well over 100 years. This is been studied between Octopus vulgaris and because highly diverse systems exist despite the Callistoctopus macropus by D.V. Meisel et al. well-established idea of competitive exclusion, They found that these two species of octopus which has at its core the principle that similar may coexist despite sharing overlapping species will compete for limited resources and temporal and geographical ranges. They that one consequence of the competition may be determined that this possible coexistence could local extinction of one or more of the be due to the two species differing behaviorally competitors. Many hypotheses have been put (Meisel et al., 2006). forward to account for high diversity, given the It is clear that morphological differences counter weight of competitive exclusion, one of between octopus species have allowed them to the earliest and more important being niche succeed in their environment; specialized diversification (Grassle & Sanders, 1973). This features such as webbing, arm length, and color hypothesis focuses on maintenance of diversity distinguish different octopus species. However, through evolution to reduce niche overlap these interspecific characteristics may not have among species thereby reducing competition and as great of an influence as individual behavior. allowing coexistence. A goal of our study was to follow on from these Under the niche diversification ideas and the literature to assess if three species hypothesis, phenotypic traits evolve in a way of octopus, differing substantially in that minimizes resource overlap. Resources in morphology, have variable foraging behavior this context include anything that could be in and prey selection that would be consistent with limited supply leading to competition, for niche diversification. example food and refuge. Often this is Patterns of foraging behavior and prey manifested in morphological traits or species- preference may differ on a species or individual specific behaviors and there is considerable level. Within the STARESO harbor in Corsica, 1 France three species of Mediterranean octopus due to their commercial importance in the may coexist which could lead to competition. international seafood trade and their prevalence The octopods living in close proximity, may in shallow waters. This species is found world- have altered their behaviors or prey preference if wide in both tropical and semitropical coastal the competition between species or individuals water from 0 to 200 meters in depth (Wood & became too great. The primary goal of this study Day, 1998). They feed mostly on invertebrates, is to determine if the difference in foraging using their radula to drill into shelled organisms behavior and prey preference between species is they cannot pry open. O. vulgaris are especially greater than the difference between each opportunistic, often holding their living prey individual’s behavior and how it could relate to within their web to keep their arms free in case their coexistence. We predict that three species, another prey item becomes available to catch Octopus vulgaris, Octopus salutii, and (Wood & Day, 1998). This species can be Callistoctopus macropus exhibit different identified by its skin mottling and is often foraging behaviors and preferred prey which spotted in or near its den made of rocks (Fiorito allows them to coexist within the STARESO & Gherardi, 1999). harbor. Callistoctopus macropus are known as the Red Octopus or White Spotted Octopus. The MATERIALS AND METHODS Mediterranean variety of this species is Background exclusive to the Mediterranean Sea and Eastern We conducted our study using the three Atlantic Ocean (Taki, 1964). They can be species of octopods that occur in shallow waters recognized by their long, ovate red to brown of the Mediterranean (Morton & Britton, 2000). mantle with distinct white spots along the mantle Octopus salutii is commonly known as and down the arms. They have a shallow web the Long Armed Octopus or Spider Octopus depth and very muscular arms that are 5 to 8 (Gofas, 2004). O. salutii is often confused with times the mantle length (Taki, 1964). C. the Common Octopus, though upon close macropus are often a shallow species, but can be examination O. salutii has a deep web depth and found between 0 and 200 meters in depth longer arms compared to their mantle length (Rocha & Cheikh, 2015). They feed on small (Gofas, 2004, Quetglas, et al., 2005). This fish and invertebrates, and tend to be generalists species occurs throughout the Mediterranean Sea (Meisel et al., 2006). and the East Atlantic Ocean, including in the We caught one O. salutii, three O. Gulf of Biscay and Cadiz between 25 and 800 vulgaris, and two C. macropus (Table 1) during meters in depth (Quetglas et al., 2005). In the September and October 2016 at Station de Mediterranean, this species primarily inhabits Recherche Océanographiques et sous-marines the lower continental shelf, as well as the upper (STARESO) in Corsica, France (42°34’N, slope. The juvenile and adult diet often consists 08°43’E). Divers caught the individuals in the of crustaceans, fishes and cephalopods (Quetglas harbor of STARESO, which is about 100m et al., 2005). O. salutii are rarely caught and across, at depths ranging from 1.8-3.6m. even more rarely studied due to their usual depth. For this reason, little is known about their juvenile biology, as they are infrequently caught, especially with a mantle length less than 9 cm. Octopus vulgaris, or the Common Octopus, is an extensively researched species 2 These containers were 51mm petri dishes, modified with multiple 4mm diameter holes in order to allow the scent of the prey to travel through the aquarium. We taped two petri dishes together using duct tape to either seal a prey item inside or keep empty as a control. In order to limit the octopus’ ability to visually visually learn, we used an opaque plexiglass sheet before and between trials to ensure that the octopus could not see the food and control being randomly placed on the Table 1: A list of the individuals and their opposite side of the tank. We measured and cut corresponding species. the plexiglass to guarantee a correct fit and Field Methods prevent the octopus from being able to slip past Opportunistic SCUBA and skin diving the partition while preparing the trials. surveys were conducted during the day and night in STARESO harbor to find an octopus of any Foraging Trials species. If an octopus was found, the divers In order to test if the various species of would catch the octopus in a mesh bag and bring octopus foraged differently, ten behaviors it into the lab. exhibited by the individuals during trials were In order to gather prey for both foraging documented. Beginning at 21:15 CET, we and food preference trials, SCUBA and skin turned off all fluorescent lights, turned on red divers flipped rocks in the STARESO harbor to lights to simulate night light, and inserted the search for Red Abalone (Haliotis tuberculata), opaque partition to divide the aquarium into two Rock Crab (Pachygrapsus marmoratus), and a equal sides. In addition, we prepared the small ark clam (Barbatia barbata). We kept the containers for the night, filling one with an prey in an aquarium with an open circuit, and abalone and leaving one empty as a control. At gathered fresh prey multiple times each week. 22:00 we would return to the lab, having given 45 minutes to acclimate to the new surroundings Materials and put the containers on the side of the tank After capturing an octopus in the harbor opposite the octopus. The containers would then at STARESO, we placed it in a large aquarium be covered with rocks to simulate abalone (250 or 300L) with nonuniform rocks and a hiding. After removing the partition, we gave the cinder block inside, giving it the opportunity to octopus 15 minutes to participate in the trial. create a den. Before beginning trials, we left the During this, we documented the starting location octopus in the aquarium for two days to of the octopus and piles, any behaviors acclimate, turning on and off lights to simulate exhibited, and whether the octopus successfully daytime. During this time the octopus were not preyed on the abalone. fed to guarantee hunger for trials. During the trials, we kept the prey in The behaviors documented are as follows: closed containers that could easily be removed if 1. Moved whole body around tank the octopus began to feed. This prevented the 2. Moved arms through water column prey from moving throughout the tank, and 3.
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