California sea lions interfere with striped marlin hunting behaviour in multi-species predator aggregations.

The open ocean offers a suite of ecological conditions promoting the occurrence of multi-species predator aggregations. These mixed predator aggregations typically hunt large groups of relatively small and highly cohesive prey. However, the mechanisms and functions of these mixed predator aggregations are largely unknown. Even basic knowledge of whether the predator species' interactions are mutualistic, commensal or parasitic is typically missing. Moreover, recordings of attack and capture rates of marine multi-species predator aggregations, which are critical in understanding how and why these interactions have evolved, are almost completely non-existent owing to logistical challenges. Using underwater video, we quantified the attack and capture rates of two high-trophic level marine predators, California sea lions (Zalophus californianus) and striped marlin (Kajikia audax) attacking schools of fishes in the Southern California Current System, offshore the Baja California Peninsula. Recording over 5000 individual attacks across 13 fish schools, which varied in species, size and predator composition, we found that sea lions kleptoparasitized striped marlin hunts and reduced the frequency of marlin attacks and captures via interference competition. We discuss our results in the context of the phenotypic differences between the predator species and implications for a better understanding of multi-species predator aggregations. This article is part of the theme issue 'Mixed-species groups and aggregations: shaping ecological and behavioural patterns and processes'.

Mechanisms of prey division in striped marlin, a marine group hunting predator.

Many terrestrial group-hunters cooperate to kill prey but then compete for their share with dominance being a strong predictor of prey division. In contrast, little is known about prey division in group-hunting marine predators that predominately attack small, evasive prey (e.g. fish schools). We identified individual striped marlin (Kajikia audax) hunting in groups. Groups surrounded prey but individuals took turns attacking. We found that competition for prey access led to an unequal division of prey among the predators, with 50% of the most frequently attacking marlin capturing 70-80% of the fish. Neither aggression, body size nor variation in hunting efficiency explained this skewed prey division. We did find that newly arrived groups of marlin gained on average more access to the prey. This raises the possibility that newly arrived marlin were hungrier and more motivated to feed. However, this result does not necessarily explain the unequal prey division among the predators because the skew in prey captures was found at the level of these groups. Dynamic prey division is probably widespread but under-reported in marine group-hunters and the inability of individuals to monopolize prey until satiation likely reduces the importance of social hierarchies for prey division.

Oil gland and oil pores in billfishes: in search of a function.

Billfishes are well known for their distinctive elongated rostra, i.e. bills. The functional significance of billfish rostra has been frequently discussed and the recent discovery of an oil gland (glandula oleofera) at the base of the rostrum in swordfish, Xiphias gladius, has added an interesting facet to this discussion regarding the potential co-evolution of gland and rostra. Here, we investigated the oil gland and oil pores (through which the oil is brought to the skin surface) of four billfish species - swordfish, Atlantic blue marlin (Makaira nigricans), Indo-Pacific sailfish (Istiophorus platypterus) and striped marlin (Kajikia audax) - and provide detailed evidence for the presence of an oil gland in the last three. All four species had a high density of oil pores on the forehead which is consistent with the hypothesis of hydrodynamic benefits of the oil. The extension of the pores onto the front half of the rostrum in sailfish and striped marlin, but not in swordfish or blue marlin, suggests that the oil may have additional functions. One such function could be linked to the antibacterial and anti-inflammatory properties of the oil. However, the available evidence on predatory rostrum use (and hence the likelihood of tissue damage) is only partly consistent with the extension of pores on rostra across species. We conclude that the oil gland probably serves multiple, non-mutually exclusive functions. More detailed information on rostrum use in blue marlin and swordfish is needed to better link behavioural and morphological data with the aim of accomplishing a full comparative analysis.

Linking hunting weaponry to attack strategies in sailfish and striped marlin.

Linking morphological differences in foraging adaptations to prey choice and feeding strategies has provided major evolutionary insights across taxa. Here, we combine behavioural and morphological approaches to explore and compare the role of the rostrum (bill) and micro-teeth in the feeding behaviour of sailfish (Istiophorus platypterus) and striped marlin (Kajikia audax) when attacking schooling sardine prey. Behavioural results from high-speed videos showed that sailfish and striped marlin both regularly made rostrum contact with prey but displayed distinct strategies. Marlin used high-speed dashes, breaking schools apart, often contacting prey incidentally or tapping at isolated prey with their rostra; while sailfish used their rostra more frequently and tended to use a slower, less disruptive approach with more horizontal rostral slashes on cohesive prey schools. Capture success per attack was similar between species, but striped marlin had higher capture rates per minute. The rostra of both species are covered with micro-teeth, and micro-CT imaging showed that species did not differ in average micro-tooth length, but sailfish had a higher density of micro-teeth on the dorsal and ventral sides of their rostra and a higher amount of micro-teeth regrowth, suggesting a greater amount of rostrum use is associated with more investment in micro-teeth. Our analysis shows that the rostra of billfish are used in distinct ways and we discuss our results in the broader context of relationships between morphological and behavioural feeding adaptations across species.