Marine Protection Induces Morphological Variation in the California Moray, Gymnothorax mordax.

The effectiveness of marine protected areas (MPAs) on the general health and conservation of species, habitats, and community interactions is of great interest to researchers, managers, and recreationalists. However, the ecological and behavioral diversity of vertebrate predators of southern California kelp forests limits our ability to make general conclusions about MPA effectiveness across a variety of species. Identifying and studying species with extreme feeding habits or prey-capture strategies may offer greater insight into predator-prey relationships and reveal the trophic importance of an animal in the larger community. Moray eels (family Muraenidae) have been shown to have morphological and behavioral adaptations that allow them to consume large prey whole, identifying them as important predators. From 2015 to 2018, we studied the health and feeding behavior of a long-lived, elusive, and benthic kelp forest predator, the California moray eel (Gymnothorax mordax). We trapped eels inside and outside of Blue Cavern Onshore State Marine Conservation Area, an MPA on the northwest side of Santa Catalina Island, CA which prohibits the take of any species. Over 4 years, we captured 1736 eels. Overall, we found that morays were longer, older, heavier, had higher body condition, and were found in greater abundance within the MPA. Although fish comprised the majority of their summer diet, morays outside of the MPA were consuming a more diverse set of fish, while kelp bass comprised more than half of the diet for morays inhabiting the MPA. Additionally, we found that morays within the MPA had larger relative vertical gape distances (VGDs) and narrower heads. Our recapture data support the high site fidelity of morays, indicating that their diet and morphology are influenced by their local community. While the majority of morays are thriving in the MPA, as suggested by their robust sizes and longevity, high abundance appears to result in higher frequencies of cannibalism, the presence of an undescribed disease, and lower growth rates. Our results suggest that the MPA affects the life history of morays and may select for an alternative feeding strategy in which eels develop larger VGDs, smaller adductor muscles, and a specialized diet which is presumably influenced by the local environment. In addition, observations of cannibalistic behavior and species-specific disease provide us with important insight into natural factors that may still regulate populations removed from anthropogenic disturbances such as fishing.

Eat whole and less often: ontogenetic shift reveals size specialization on kelp bass by the California moray eel, Gymnothorax mordax.

Despite the importance of predation in many ecosystems, gaps remain in our understanding of nocturnal marine predators. Although the kelp forests of Southern California are some of the most well-studied ecosystems, California morays, Gymnothorax mordax, are predominately nocturnal predators that have remained largely unstudied and their predatory effects on the kelp forest ecosystem are unknown. We use a multi-year data set to examine the dietary breadth of G. mordax and to determine the functional role of this predator. We also quantify bite force to examine the potential performance limitations of morays in exploiting prey. Stomach content analyses and linear selectivity index values indicate that G. mordax specializes on kelp bass, Paralabrax clathratus. Average size of kelp bass consumed varies across years, suggesting that morays respond to fluctuations in prey size availability. The scaling relationship of kelp bass standard length and moray head length reveals an ontogenetic shift, where maximum prey size increases with moray size and small prey are dropped from the diet of larger individuals. Moray bite force exhibited strong positive allometry with moray head size, suggesting that larger morays exhibit greater bite forces for their head and body size. However, we found no relationship between prey size and bite force, suggesting that a disproportional increase in bite force does not facilitate the consumption of disproportionately larger prey. Our results indicate that while G. mordax of Catalina Island is a dietary specialist, it is capable of exhibiting functional shifts in prey size and species based on their abundance.

The beach-spawning California grunion Leuresthes tenuis eats and digests conspecific eggs.

Apparent egg cannibalism was investigated in the beach-spawning California grunion Leuresthes tenuis. Three hypotheses were tested to determine whether L. tenuis regularly consumes and efficiently digests conspecific eggs. First, examination of the gut contents of adults collected at four spawning sites over two seasons showed that the intestines of most fish from all the sites (57-87%, n ≥ 30, each site) contained L. tenuis eggs. The two other hypotheses focused on digestion of the eggs. First, the force required to crush cannibalized eggs was significantly less than that for uncannibalized eggs (fertilized or unfertilized), indicating that ingestion weakens the egg chorions. Second, conspecific eggs fed to fish held in the laboratory visibly degraded as they passed through the gut. The eggs lost c. half of their protein content and about two-thirds of their lipid content as they passed from proximal to distal regions of the gut, indicating that digestion occurred. Digestive enzyme activities of the gut further confirmed that L. tenuis can break down the contents of ingested eggs. Trypsin activity decreased and aminopeptidase activity increased posteriorly along the gut, whereas amylase and lipase activities exhibited less clear patterns by gut region. As far as is known, this study is the first to show that L. tenuis is an egg cannibal.

Effects of prey characteristics on the feeding behaviors of an apex marine predator, the California moray (Gymnothorax mordax).

Moray eels comprise a large radiation of elongate marine predators that are thought to swallow large prey whole but also circumvent gape constraints by manipulating prey into more manageable pieces. Prey manipulation behaviors include shaking, rotation, knotting, and ramming prey against another object to assist in swallowing. Most morays feed on a wide variety of prey that vary in mechanical properties such as stiffness and toughness, which could potentially affect feeding behaviors. There is little diet data informing us of the maximum prey size morays can swallow whole and whether maximum prey size differs between prey types. Our study examines feeding behaviors for the California moray (Gymnothorax mordax) in the laboratory. We recorded morays feeding on freshly thawed fish and cephalopods of varying size. We found that prey size had a strong effect on total feeding time and manipulation duration for both fish and cephalopods. While morays were observed using a diversity of prey manipulation behaviors and the durations for each of these behaviors increased with prey size, prey type had no effect on manipulation behaviors employed. Total manipulation duration, however, comprised a greater proportion of total feeding time for fish compared to cephalopods. As relative prey mass (RPM) increased for cephalopods, morays spent a greater proportion of their total feeding time transporting prey. Transport rate was higher for cephalopod prey but the relationship between RPM and transport rate was negative for both prey types. Despite this decrease in transport rate, we attribute the lower total feeding times for larger cephalopod prey compared to fish to behavioral tactics of morays. Morays used the corners of the aquaria to aid in the transport of larger cephalopod prey. We hypothesize that the deformable tissues of cephalopods and the presumably low coefficient of friction of their thawed mantles and tentacles may be difficult for the recurved teeth on the pharyngeal jaws to pierce and grip during transport.

Scaling of dentition and prey size in the California moray (Gymnothorax mordax).

Scaling patterns of tooth morphology can provide insights on prey capture strategy and dietary patterns as species grow through ontogeny. We report the scaling of dentition and diet and how it relates to body size in the California moray, Gymnothorax mordax. We sampled lengths, widths, and curvature for teeth lining five distinct regions of the oral jaws across 21 G. mordax individuals ranging from 383 to 1110mm total length. Absolute tooth length in relation to moray size shows positive allometry only for the outer maxillary teeth, while teeth lining the inner maxilla display positive allometry in tooth base width. All other regions exhibit isometric growth in both length and width relative to moray size. Similar to previous descriptions of other moray species, the longest teeth in the oral jaws are the median intermaxillary teeth. This series of three teeth are depressible and rooted in the center of the ethmovomer, the bone that forms the roof of the rostrum. We hypothesize that caudal mobility of the median intermaxillary teeth aids in prey transport by enabling the pharyngeal jaws to remove pierced prey without requiring full abduction of the oral jaws. The predominantly isometric tooth growth in G. mordax suggests that the oral teeth grow proportionately as individuals increase in size. Stomach contents from the field suggest that G. mordax is highly piscivorous. While a strong positively allometric relationship between vertical gape and head length supports the expectation that moray increase relative prey size over ontogeny, we found no relationship between prey standard length and moray size. This suggests that while larger individuals are capable of consuming larger prey, individual G. mordax are opportunistic predators that do not specialize on prey of a specific size over ontogeny.