14Batch spawning, intermittent spawning and multiple spawning represent common 15 reproductive strategies among cephalopods. These flexible spawning strategies are also common 16 in fishes, and are thought to be a female trait that is programmed depending on environmental 17 parameters. The ommastrephid squid Todarodes pacificus, being a terminal spawner, is 18 considered to have a single spawning event, extruding one large egg mass and dying soon 19 thereafter. Females that are interrupted by males exhibiting mating behavior, while extruding the 20 egg mass, instead spawn multiple egg masses over the course of 2-3 days instead of dying soon 21 after spawning the first egg mass. We demonstrate that male mating behavior causes "forced" 22 intermittent spawning by females (i.e., more than one spawning event). We hypothesize that in T. 23 pacificus, some males use this strategy to mate with females unable to repel advances while 24 spawning, and thus providing the male with the opportunity to contribute sperm and enhance 25 gene flow. 26Keywords: batch spawning, spawning interruption, multiple paternity, egg mass, oceanic squid 27
INTRODUCTION 28Coleoid cephalopods are considered semelparous (i.e., no gonadal resting phase) 29 (Mangold 1987), with the exception of the recently reported iteroparous vampire squid (Hoving 30 et al. 2015). Although the spawning pattern of cephalopods is monocyclic (single spawning 31 season), semelparity occurs in species that spawn eggs in single or multiple events, with their 32 reproductive strategies being considered very flexible (Pecl 2001). The terms "batch spawning," 33 "intermittent spawning," and "multiple spawning" are common in studies of cephalopod 34 reproductive biology, particularly among species of the family Ommastrephidae, with these 35 3 terms being used interchangeably (Rocha et al. 2001). This family contains a few species that 36 spawn (lay egg masses) intermittently, with no somatic growth between spawning events 37 (Nigmatullin & Laptikhovsky 1994, Nigmatullin 2011. For instance, Todarodes pacificus, an 38 ommastrephid squid, is a semelparous cephalopod mollusk that lays eggs by embedding oocytes 39 inside a large egg mass (Sakurai et al. 2013). Being an intermittent terminal spawner, the 40 female's feeding ceases prior to spawning (Bower & Sakurai 1996) in order to allocate all 41 available energy to spawning, during which the female undergoes strenuous muscular 42 contractions, involving the mantle, arms, and tentacles, while extruding the egg mass (Hamabe 43 1962). 44The multiple spawning events of semelparous species are assumed to be a programmed 45 reproductive strategy (Rocha et al. 2001); however, the reason why all spawning events by these 46 individuals occur in the final days or weeks of their life cycle remains unknown (lifespan ≈ 1 y) 47 (Rocha et al. 2001). For T. pacificus, the reported residual fecundity and potential fecundity are 48 >150,000 and 320,000-470,000 (Soeda 1956) respectively. Thus, it is possible that the multiple ...