Morphometric analysis of male reproductive features of octopodids (Mollusca: Cephalopoda)

Taxonomic accounts of octopodids frequently describe the spermatophore, the penis that releases the spermatophore from the internal organs, and the ligula and calamus that transfer it to a female. To explore relationships among these male features and body size, this study applies principal components analysis to data from 43 species of the family Octopodidae, or benthic octopuses. Covariation in penis and mantle length opposed by covariation in ligula and calamus lengths forms primary shape variation. Secondary shape variation is due to opposing variation between ligula and calamus lengths. Primary shape variation is greatest among shallow-water species. The calami and ligulae of diurnal and crepuscular shallow-water species are short compared to those of nocturnal shallow-water species. Because these structures contain heterogeneous collagen arrays and lack camouflaging chromatophore organs, they are white. Diurnal and crepuscular octopus species may minimize their lengths due to selection imposed by visual predators. Secondary shape variation is greater in deep-sea and high-latitude octopuses. Members of Voss's Eledoninae (except Eledone) and Graneledoninae and two species of Benthoctopus have exceptionally long calami and comparatively short ligulae; these lengths vary among members of the Bathypolypodinae. Variation in spermatophore length is independent of the structures considered.     

The effects of habitat on the vigilance of shorebirds: Is visibility important?

Previous studies of vigilance have concentrated on situations where the prey species has an unimpaired view of its surroundings. Here the effects of reduced visibility caused by objects adjacent to the prey are studied in two species of shorebird. A reduction in visibility causes an increase in the level of vigilance, indicating an increase in vulnerability despite the greater degree of camouflaging. This increase is due to individuals being less able to see both approaching predators and their neighbours. Turnstones, Arenaria interpres, and purple sandpipers, Calidris maritima, show very similar increases in the level of vigilance with decreasing visibility, but achieve these increases by different means: turnstones lengthen the duration of each vigilant scan, while purple sandpipers scan more often. Increasing scanning rate produces a shortened interval between scans, which reduces the risk of being caught unawares by an approaching predator. However, it may also reduce feeding efficiency, and it is suggested that this might be more serious in turnstones due to greater handling times for food items, so causing them to adopt a different strategy to increase vigilance.     

Factors affecting the spatial distribution of natural populations of Octopus joubini robson

Observations were made on the density and distribution of Octopus joubini in its natural environment. The average density of octopuses was 1/33 m² but their distribution was clumped. This distribution correlated significantly with the distribution of molluscan shells in which they hid. In addition, octopuses were attracted to sites that had been enriched with extra empty gastropod shells. Since site stability was not observed and they were not repelled from one another when placed at a high density, octopuses were probably not spaced by social pressure. Small crab species, the normal prey of O. joubini, were at a very high density of 30/m², and food abundance probably did not limit octopus number. The main factor affecting distribution of this population of octopuses may be predator pressure acting through the availability of places to hide.     

Egg brooding by deep-sea octopuses in the North Pacific Ocean

Videotapes made from the submersible Alvin on Baby Bare, a 2600-m-deep North Pacific basalt outcrop, and at two other deep-sea localities document that octopuses of the genera Graneledone and Benthoctopus attach their eggs to hard substrate and apparently brood them through development. The behavior of brooding females was generally similar to that of shallow-water octopuses, but the genera showed apparent differences. In addition to the high density of brooding females observed at Baby Bare, which may relate to the increased availability of exposed hard substrates for egg attachment and of prey, females are suggested to increasingly associate with hard substrates as they mature. The biology of Baby Bare may seem unduly unique because the outcrop is isolated on a sedimented plain and is among the few exposures of hard substrate other than hydrothermal vents that have been explored by submersible. On the sediment-covered ocean floor, the availability of hard substrate may strongly affect the distribution of brooding octopuses. The size and shape of boreholes in 19 of over 400 thyasirid clam shells collected from Baby Bare support the hypothesis that octopuses had preyed upon the clams.     

Crypsis, conspicuousness, mimicry and polyphenism as antipredator defences of foraging octopuses on Indo-Pacific coral reefs, with a method of quantifying crypsis from video tapes

We tested the hypothesis that soft-bodied octopuses, which spend most of their lives in dens, remain highly cryptic as their primary defence against predation while they forage. We videotaped foraging octopuses on two widely dispersed Pacific coral reefs and developed a rigorous method to analyse the degree of crypsis from videotapes. Five ranks were assigned (two of‘ highly cryptic’, one of ‘moderately cryptic’, and two of ‘conspicuous’) to assess each octopus's body pattern match to its background, using the criteria of brightness, colour, shape and skin patterning. The data do not support the hypothesis. In Tahiti, octopuses were highly cryptic only 54%, moderately cryptic 24% and conspicuous 22% of the time. In Palau, the respective calculations were 31 %, 19% and 50%. A major feature of their behaviour was their remarkable ability to instantly change their body pattern, or phenotype, by direct neural control of the skin. Six chronic and nine acute categories of body patterns were observed. On average, octopuses changed their phenotype 2.95 times/minute, or 177 times per hour, based upon 7.5 hours of videotaped foraging. This rapid neurally controlled polyphenism was used most often to adjust their appearance as they foraged slowly across highly diverse substrates, thus implementing appropriate mechanisms of crypsis over each (e.g. general background resemblance, deceptive resemblance, disruptive coloration). However, when crawling rapidly, or swimming for short distances, octopuses often engaged a second antipredator lactic that was conspicuous: mimicking fishes or showing bold disruptive patterns that rendered them visibly different from an octopus. Nevertheless, sometimes they were simply conspicuous even when moving slowly, particularly in Palau, where the octopuses were larger, there was a high degree of mating“, and fewer signs of predation were evident. The results suggest that, while foraging, the overall strategy is to use polyphenism to produce ‘apparent rarity’ of any single phenotype (or search image) through mechanisms of crypsis, conspicuousness and mimicry, all of which are guided by keen vision in this marine invertebrate.     

Color matching on natural substrates in cuttlefish, <Emphasis Type="Italic">Sepia officinalis</Emphasis>

The camouflaging abilities of cuttlefish (Sepia officinalis) are remarkable and well known. It is commonly believed that cuttlefish-although color blind-actively match various colors of their immediate surroundings, yet no quantitative data support this notion. We assembled several natural substrates chosen to evoke the three basic types of camouflaged body patterns that cuttlefish express (uniform/stipple, mottle, and disruptive) and measured the spectral reflectance of the camouflaged pattern and the respective background using a fiber optic spectrometer. We demonstrate that the reflectance spectra of cuttlefish skin patterns correlate closely with the spectra of these natural substrates. Since pigmented chromatophores play a key role in cephalopod color change, we also measured the spectral reflectance of individual cuttlefish chromatophores under the microscope, and confirm the results from a previous publication reporting three distinct colors of chromatophores (yellow, orange, and dark brown) on the animals' dorsal side. Taken together, our results show that the color variations in substrate and animal skin can be very similar and that this may facilitate color match on natural substrates in the absence of color vision.     

Autonomous Learning in Maze Solution by Octopus

We tested seven octopuses, Octopus vulgaris, in maze-learning experiments. They tried to reach the goal, so as to get a reward, by using various locomotory actions in the path, and sometimes encountered obstacles. They came to select efficient swimming actions in the path; afterwards less efficient tactile actions (crawling, staying put, and so on: these reduce the speed of movement) gradually increased, while time to detour around the obstacle was reduced. To investigate whether octopuses reduce time spent detouring around obstacles by estimating their actions in the path, we devised a trade-off situation in which octopuses were obliged to use tactile actions even though the set-up also encouraged them to use swimming actions. As a result, we could observe that they reduced the detouring time. In that way, we experimentally constituted a perspective as if octopuses looked around the whole maze and estimated their actions. Such a perspective appeared to be autonomous learning.     

Evolution of development type in benthic octopuses: holobenthic or pelago-benthic ancestor?

Octopuses of the family Octopodidae are singular among cephalopods in their reproductive behavior, showing two major reproductive strategies: the first is the production of few and large eggs resulting in well-developed benthic hatchlings (holobenthic life history); the second strategy is the production of numerous small eggs resulting in free-swimming planktonic hatchlings (pelago-benthic life history). Here, we utilize a Bayesian-based phylogenetic comparative method using a robust molecular phylogeny of 59 octopus species to reconstruct the ancestral states of development type in benthic octopuses, through the estimation of the most recent common ancestors and the rate of gain and loss in complexity (i.e., planktonic larvae) during the evolution. We found a high probability that a free-swimming hatchling was the ancestral state in benthic octopuses, and a similar rate of gain and loss of planktonic larvae through evolution. These results suggest that in benthic octopuses the holobenthic strategy has evolved from an ancestral pelago-benthic life history. During evolution, the paralarval stage was reduced to well-developed benthic hatchlings, which supports a “larva-first” hypothesis. We propose that the origin of the holobenthic life history in benthic octopuses is associated with colonization of cold and deep sea waters.     

Microsatellite inheritance and multiple paternity in the deep-sea octopus Graneledone boreopacifica (Mollusca: Cephalopoda)

Abstract. Octopodids are a globally distributed group of marine molluscs. Despite this, our knowledge of their reproductive biology rests heavily on inference, as all phases of copulation, beginning with sperm transfer, occur within the mantle cavity. Male octopuses insert a spermatophore into the female's oviduct, which is predicted to release a sperm-filled sac that either bursts to release sperm for storage or to itself be stored in a gland in the middle of the oviduct. To test whether female octopuses use sperm from multiple males to fertilize their eggs, as may be predicted from anatomy and anecdotal accounts, we apply microsatellite analysis to a partial clutch of Graneledone boreopacifica collected at 1600-m depth to test for multiple paternity. At least two genetically distinct sires contributed sperm to the hatchlings analyzed, demonstrating for the first time multiple paternity in octopodids.     

Pull or Push? Octopuses Solve a Puzzle Problem

Octopuses have large brains and exhibit complex behaviors, but relatively little is known about their cognitive abilities. Here we present data from a five-level learning and problem-solving experiment. Seven octopuses (Octopus vulgaris) were first trained to open an L shaped container to retrieve food (level 0). After learning the initial task all animals followed the same experimental protocol, first they had to retrieve this L shaped container, presented at the same orientation, through a tight fitting hole in a clear Perspex partition (level 1). This required the octopuses to perform both pull and release or push actions. After reaching criterion the animals advanced to the next stage of the test, which would be a different consistent orientation of the object (level 2) at the start of the trial, an opaque barrier (level 3) or a random orientation of the object (level 4). All octopuses were successful in reaching criterion in all levels of the task. At the onset of each new level the performance of the animals dropped, shown as an increase in working times. However, they adapted quickly so that overall working times were not significantly different between levels. Our findings indicate that octopuses show behavioral flexibility by quickly adapting to a change in a task. This can be compared to tests in other species where subjects had to conduct actions comprised of a set of motor actions that cannot be understood by a simple learning rule alone.     

Gonadal development and reproductive strategies of the tropical octopus (Octopus insularis) in northeast Brazil

Octopus insularis inhabits shallow waters along the coast and oceanic islands of northeastern Brazil, where it is the main target of commercial fishing of octopuses. This study aims to investigate the hypothesis that the tropical O. insularis has a distinct gonadal development and reproductive features when compare to its congener O. vulgaris from the subtropical regions. In order to describe its reproductive development, 545 octopuses were collected in the Northeastern Brazil. A good correspondence was observed between the gonad morphology and its histological structure. Oocytes in different development stages were observed in mature females. Most female in early maturity stages had sperm stored in the spermathecae, indicating that females copulate when still immature. There was no correlation between testis weight and the Needham complex, suggesting a protracted period of spermatophore production. Octopus insularis has a general gonadal development pattern similar to O. vulgaris, however, some differences were observed, as maturation at a smaller size, probably associated to a shorter life, and lower fecundity. The distinct reproductive features of O. insularis seem to be related to less variable conditions in the tropical environments. Management should take into account the differences and establish specific rules for the Northeast Brazil octopus fisheries.     

Decision tree classification of behaviors in the nesting process of green turtles (Chelonia mydas) from tri-axial acceleration data

Sea turtles are endangered marine-adapted reptiles that are obligate terrestrial nesters. They have a nesting process that involves a series of behavioral phases: (1) crawling, (2) digging a body pit, (3) digging an egg chamber, (4) depositing eggs, (5) covering the egg chamber, and (6) camouflaging the site. Discrimination among these behavioral phases is necessary for estimating which parts of the nesting process are affected by human activities or characteristics of beaches. We calculated multivariate features from tri-axial accelerations obtained by a data logger to extract key features for discriminating between these behavioral phases of green turtles (Chelonia mydas). The developed decision tree classifier discriminated these behaviors with high accuracy, resulting in 5 behaviors after combining digging a body pit and camouflaging the site (86.207 %). The structure of the tree showed key features for discriminating the 5 behaviors on the basis of differences in the movements of turtles. The decision rules in this study will enable us to discriminate behavioral phases quantitatively without observer interference, thereby providing a basis for estimating turtle energy budget and the influence of human activities on nesting behavior.     

Bipedal locomotion in Octopus vulgaris: A complementary observation and some preliminary considerations

Lacking an external shell and a rigid endoskeleton, octopuses exhibit a remarkable flexibility in their movements. Bipedal locomotion is perhaps the most iconic example in this regard. Until recently, this peculiar mode of locomotion had been observed only in two species of tropical octopuses: Amphioctopus marginatus and Abdopus aculeatus. Yet, recent evidence indicates that bipedal walking is also part of the behavioral repertoire of the common octopus, Octopus vulgaris. Here we report a further observation of a defense behavior that encompasses both postural and locomotory elements of bipedal locomotion in this cephalopod. By highlighting differences and similarities with the other recently published report, we provide preliminary considerations with regard to bipedal locomotion in the common octopus.     

Nutrient absorption,storage and remobilization in octopus vulgaris

Average absorption and conversion to ¹⁴CO₂for free leucine included in a meal were 96% and 30% after 24 h. The values for glucose were 98% and 48% and for palmitate 46% and 12.5%. Muscle was the major repository of leucine (38% of the total ingested) and glucose (44%), but the digestive gland contained most of the palmitate (20%).

During normal feeding ¹⁴CO₂ production from octopuses given leucine dropped to low stable levels after only 2 days, those given glucose required 4 to 5, but palmitate apparently did not enter a stable reserve. Fasting increased the release of ¹⁴CO₂ from octopuses given palmitate and leucine, but glucose animals showed little change. A 5‐fold increase in ¹⁴CO₂ production during forced exercise after fasting by octopuses given glucose may indicate increased carbohydrate catabolism. Reduced ¹⁴CO₂production in exercise after other substrates is consistent with this, suggesting muscle carbohydrate reserves may be exclusively for locomotion.

A comparison of nutrient uptake, catabolism and growth suggests that lipids are the limiting nutrient for octopuses on a crab diet.     

Mimicry and foraging behaviour of two tropical sand-flat octopus species off North Sulawesi, Indonesia

The so-called 'mimic octopuses' of tropical Indonesia are reputed to mimic up to 13 species of other local marine organisms. We tested for mimicry by allowing individuals of two species of octopus to habituate to divers, then observing and filming two species continuously as they foraged daily in the same open, featureless volcanic sand habitat. Mimicry of a local, abundant flounder occurred commonly during 5 days of natural foraging: nearly 500 episodes were analysed. Both octopus species mimicked the shape, swimming actions, speed, duration, and sometimes the coloration of swimming flounders. During flounder mimicry, octopuses were actively moving and conspicuous, whereas immediately before and after flounder mimicry, they were camouflaged and motionless (sitting or very slowly crawling). Furthermore, when motionless, octopuses assumed body patterns and postures that resembled small sponges, tube-worm tubes, or colonial tunicates, which were among the few objects in the open sand habitat. The key finding was that octopuses used flounder mimicry only when their movement would give away camouflage in this open habitat. In all cases, octopuses used mimicry as a primary defense. Several interactions with fishes and stomatopods were filmed and typical secondary defense behaviours, not mimicry, were used by the octopuses. Foraging occurred twice per day and two tactile feeding tactics were used. Dens and food were not limiting; thus, we observed a highly unusual circumstance of a guild of small, long-armed octopus species that shared the same habitat, den sources, food, activity period, and some behaviours.  © 2008 The Linnean Society of London, Biological Journal of the Linnean Society , 2008, 93 , 23–38.     

Home occupancy by male Octopus vulgaris in a large seawater tank

Pairs of individually recognizable male Octopus vulgaris were observed in a large seawater tank containing two suitable homes (brick pots or plastic buckets). None of the animals established exclusive occupancy of one home and for much of the time both animals were associated together at the same site. Usually one of the two homes was preferred and its occupant was most likely to be the larger animal, or the earlier resident if they were of equal size. Large animals were observed to take food forcefully from smaller octopus. An arm alignment interaction is described which, it is suggested, may be a means by which two octopuses establish their relative sizes.     

Interspecific feeding associations of groupers (Teleostei: Serranidae) with octopuses and moray eels in the Gulf of Eilat (Agaba)

Synopsis Temporary interspecific feeding associations between groupers (Serranidae) and moray eels or octopuses in the Red Sea were observed 41 times during over 400 scuba dives. Such behavior is a feeding tactic adopted by groupers in which small organisms, escaping the primary (= nuclear) predators (eel or octopus), are caught by the groupers. Eels and octopuses, due to their body structure or shape, gain access to crevices that most fishes cannot enter. This feeding tactic increases the diversity of food resources used by their associates by making normally inaccessible food items available to them. Serranids from many parts of the world exploit normally unavailable food resources in this fashion. It appears that this is a learned behavior which increases food diversity and feeding efficiency.     

Interspecific evaluation of octopus escape behavior

The well-known ability of octopuses to escape enclosures is a behavior that can be fatal and, therefore, is an animal welfare issue. This study obtained survey data from 38 participants-primarily scientists and public aquarists who work with octopuses-on 25 described species of octopus. The study demonstrates that the likeliness to escape is species specific (p = .001). The study gives husbandry techniques to keep captive octopuses contained. This first interspecific study of octopus escape behavior allows readers to make informed species-specific husbandry choices.     

Octopus vulgaris uses visual information to determine the location of its arm

Octopuses are intelligent, soft-bodied animals with keen senses that perform reliably in a variety of visual and tactile learning tasks. However, researchers have found them disappointing in that they consistently fail in operant tasks that require them to combine central nervous system reward information with visual and peripheral knowledge of the location of their arms. Wells claimed that in order to filter and integrate an abundance of multisensory inputs that might inform the animal of the position of a single arm, octopuses would need an exceptional computing mechanism, and "There is no evidence that such a system exists in Octopus, or in any other soft bodied animal." Recent electrophysiological experiments, which found no clear somatotopic organization in the higher motor centers, support this claim. We developed a three-choice maze that required an octopus to use a single arm to reach a visually marked goal compartment. Using this operant task, we show for the first time that Octopus vulgaris is capable of guiding a single arm in a complex movement to a location. Thus, we claim that octopuses can combine peripheral arm location information with visual input to control goal-directed complex movements.