Mark D. Norman

Dynamic mimicry in an Indo–Malayan octopus

During research dives in Indonesia (Sulawesi and Bali), we filmed a distinctive long-armed octopus, which is new to science. Diving over 24 h periods revealed that the ‘mimic octopus’ emerges during daylight hours to forage on sand substrates in full view of pelagic fish predators. We observed nine individuals of this species displaying a repertoire of postures and body patterns, several of which are clearly impersonations of venomous animals co-occurring in this habitat. This ‘dynamic mimicry’ avoids the genetic constraints that may limit the diversity of genetically polymorphic mimics but has the same effect of decreasing the frequency with which predators encounter particular mimics. Additionally, our observations suggest that the octopus makes decisions about the most appropriate form of mimicry to use, allowing it to enhance further the benefits of mimicking toxic models by employing mimicry according to the nature of perceived threats.

Female impersonation as an alternative reproductive strategy in giant cuttlefish

Out of all the animals, cephalopods possess an unrivalled ability to change their shape and body patterns. Our observations of giant cuttlefish (Sepia apama) suggest this ability has allowed them to evolve alternative mating strategies in which males can switch between the appearance of a female and that of a male in order to foil the guarding attempts of larger males. At a mass breeding aggregation in South Australia, we repeatedly observed single small males accompanying mating pairs. While doing so, the small male assumed the body shape and patterns of a female. Such males were never attacked by the larger mate–guarding male. On more than 20 occasions, when the larger male was distracted by another male intruder, these small males, previously indistinguishable from a female, were observed to change body pattern and behaviour to that of a male in mating display. These small males then attempted to mate with the female, often with success. This potential for dynamic sexual mimicry may have played a part in driving the evolution of the remarkable powers of colour and shape transformation which characterize the cephalopods.

Sex in giant squid

Recent captures of two female giant squid ( Architeuthis ) off southern Australia have provided the first record of a mated female specimen of these almost mythical deepsea creatures. We found sperm packages (spermatophores) embedded within the skin of both ventral arms of the larger of the two specimens. It seems that male giant squids may use their muscular elongate penis to ‘inject’ sperm packages under pressure directly into the arms of females.

Mitochondrial genome diversity and population structure of the giant squid Architeuthis: genetics sheds new light on one of the most enigmatic marine species

Despite its charismatic appeal to both scientists and the general public, remarkably little is known about the giant squid Architeuthis, one of the largest of the invertebrates. Although specimens of Architeuthis are becoming more readily available owing to the advancement of deep-sea fishing techniques, considerable controversy exists with regard to topics as varied as their taxonomy, biology and even behaviour. In this study, we have characterized the mitochondrial genome (mitogenome) diversity of 43 Architeuthis samples collected from across the range of the species, in order to use genetic information to provide new and otherwise difficult to obtain insights into the life of this animal. The results show no detectable phylogenetic structure at the mitochondrial level and, furthermore, that the level of nucleotide diversity is exceptionally low. These observations are consistent with the hypotheses that there is only one global species of giant squid, Architeuthis dux (Steenstrup, 1857), and that it is highly vagile, possibly dispersing through both a drifting paralarval stage and migration of larger individuals. Demographic history analyses of the genetic data suggest that there has been a recent population expansion or selective sweep, which may explain the low level of genetic diversity.

Revision of the Octopus horridus species-group, including erection of a new subgenus and description of two member species from the Great Barrier Reef, Australia

The tropical Indo-West Pacific region contains a distinctive group of small to moderate-sized octopuses referred to in the past as the Octopus horridus species-group. Member species are found primarily on intertidal reef flats. They possess small bodies, long arms and complex skin sculpture and body patterns enabling excellent camouflage. When attacked, these octopuses are capable of autotomising their arms at the base: the writhing severed arm acting as a decoy to predators and aiding escape. Lost arms regenerate within 2–3 months. Attributes of this group of octopuses are described and the subgenus Abdopus, subgen. nov. is here coined to define this group. Historically, many members of this subgenus have been incorrectly identified as Octopus horridus, a distinct large-egg species known only from the Red Sea and the northwest Indian Ocean. Seven species are here recognised as belonging in Abdopus, subgen. nov. Two member species from Great Barrier Reef waters (Octopus aculeatus d’Orbigny, 1834 and Octopus capricornicus, sp. nov.) are described. A number of additional, as yet undescribed, species occur throughout the Indo-West Pacific region. Certain earlier works have linked other octopus genera and species with members of this subgenus on the grounds of long arms and arm autotomy. Significant differences occur in the morphologies of these taxa and the nature of the arm autotomy processes. It is proposed that these groups have evolved independently and that arm autotomy has arisen more than once amongst the octopuses. The restriction of the subgenus Abdopus to the tropical Indo-West Pacific region suggests relatively recent origins and radiation.

The ink sac clouds octopod evolutionary history

Difficulties in elucidating the evolutionary history of the octopods have arisen from problems in identifying informative morphological characters. Recent classifications have divided the largest group, the incirrate octopods, into five groups. These include the pelagic superfamily Argonautoidea and three gelatinous pelagic families (Vitreledonellidae, Bolitaenidae, Amphitretidae). All benthic incirrate octopods have been accommodated in the family Octopodidae, itself divided into four subfamilies, Octopodinae, Eledoninae, Bathypolypodinae and Graneledoninae, which are defined by the presence or absence of an ink sac, and uniserial or biserial sucker arrangements on the arms. We used relaxed clock models in a Bayesian framework and maximum likelihood methods to analyse three nuclear and four mitochondrial genes of representatives from each of the previous subfamilies. Strong evidence indicates that the family Octopodidae is paraphyletic and contains the gelatinous pelagic families. The subfamilies of Octopodidae recognised in earlier works do not reflect evolutionary history. The following clades were supported in all analyses: (1) Eledone/Aphrodoctopus, (2) Callistoctopus/Grimpella/Macroctopus/Scaeurgus, (3) Abdopus/Ameloctopus/Amphioctopus/Cistopus/Hapalochlaena/Octopus, (4) Enteroctopus/Muusoctopus/Vulcanoctopus, (5) Vitreledonella/Japetella, (6) Southern Ocean endemic and deep-sea taxa with uniserial suckers. These clades form the basis for a suite of taxa assigned family taxonomic rank: Amphitretidae, Bathypolypodidae, Eledonidae, Enteroctopodidae, Megaleledonidae and Octopodidae sensu nov. They are placed within the superfamily Octopodoidea.

Redescription of The Southern Dumpling Squid Euprymna Tasmanica and A Revision of The Genus Euprymna (Cephalopoda: Sepiolidae)

Recent attention to members of the sepiolid squid genus Euprymna and symbiotic associations with luminescent bacteria ( Vibrio fischeri strains) has prompted a review of this poorly-resolved group of squids. Twelve nominal species have been placed in this genus of which the majority are ill-defined, known only from their original descriptions and separated on the basis of inadequate characters. As a first step in resolving this group, a temperate Australian species, the Southern dumpling squid, Euprymna tasmanica , is here redescribed in detail. As the genus Euprymna currently stands, most members are only distinguished on the number and position of enlarged suckers in mature males. No diagnostic characters are available to identify females. All nominal species placed in this genus are reviewed and a key to proposed valid species is presented. Six species are considered here to be valid: Euprymna berryi, E. hoylei, E. morsei, E. scolopes, E. tasmanica and an undescribed species treated here as Euprymna sp. 1. Euprymna similis is a synonym of E. morsei of Japan. Due to inadequate original descriptions, and lost or poor type material, two species are considered here to be nomen dubia ( E. schneehageni and E. pusilla ), while the taxonomic status of four additional species remain unresolved ( E. albatrossae, E. bursa, E. phenax and E. stenodactyla ).

Morphological assessment of the Octopus vulgaris species complex evaluated in light of molecular‐based phylogenetic inferences

Cryptic species are common in the ocean, particularly among marine invertebrates such as octopuses. Delineating cryptic species is particularly problematic in octopus taxonomy where the plasticity recorded among taxonomic characters often results in low resolution at the species level. This study investigated the morphological relationships among seven phylogenetic clades (identified using cytochrome c oxidase subunit I) of the broadly distributed Octopus vulgaris species complex and close relatives. Morphological analyses in this study were successful in delimiting O. sinensis, Brazilian O. vulgaris and O. vulgaris sensu stricto, which was congruent with the molecular findings of this study. Analyses based on male morphology were successful in distinguishing 14 of 15 total pairwise comparisons and proved to be a more reliable indicator of species‐level relationships in comparison with female morphology. The majority of characters with the greatest discriminatory power were male sexual traits. Significant morphological differences were also recorded among sampling localities of conspecifics, with phenotype showing correlation with local environmental data. The findings of this study support the hypothesis that multiple O. vulgaris‐like species are currently being incorrectly treated under a single species name, O. vulgaris. Octopuses being exported globally under the name O. vulgaris are of extremely high fisheries market value and profile. Our findings have potentially significant implications for the naming and conservation of commercially harvested members of this species complex throughout their ranges.

Allopatric speciation within a cryptic species complex of Australasian octopuses

Despite extensive revisions over recent decades, the taxonomy of benthic octopuses (Family Octopodidae) remains in a considerable flux. Among groups of unresolved status is a species complex of morphologically similar shallow-water octopods from subtropical Australasia, including: Allopatric populations of Octopus tetricus on the eastern and western coasts of Australia, of which the Western Australian form is speculated to be a distinct or sub-species; and Octopus gibbsi from New Zealand, a proposed synonym of Australian forms. This study employed a combination of molecular and morphological techniques to resolve the taxonomic status of the ‘tetricus complex’. Phylogenetic analyses (based on five mitochondrial genes: 12S rRNA, 16S rRNA, COI, COIII and Cytb) and Generalised Mixed Yule Coalescent (GMYC) analysis (based on COI, COIII and Cytb) distinguished eastern and Western Australian O. tetricus as distinct species, while O. gibbsi was found to be synonymous with the east Australian form (BS = >97, PP = 1; GMYC p = 0.01). Discrete morphological differences in mature male octopuses (based on sixteen morphological traits) provided further evidence of cryptic speciation between east (including New Zealand) and west coast populations; although females proved less useful in morphological distinction among members of the tetricus complex. In addition, phylogenetic analyses suggested populations of octopuses currently treated under the name Octopus vulgaris are paraphyletic; providing evidence of cryptic speciation among global populations of O. vulgaris, the most commercially valuable octopus species worldwide.

Multiple fitness benefits of polyandry in a cephalopod.

Background Sex differences in reproductive investment play a crucial role in sexual conflict. One intriguing aspect of sexual conflict is the evolution of female multiple mating (polyandry), particularly in systems where females receive no obvious direct benefits from males, and where mating is highly costly. Here, theory predicts that polyandrous females can increase their reproductive success by taking advantage of the genetic benefits of mating with multiple males. Cephalopods provide a model system for addressing this question, as all species mate multiply. Here we examine differences in reproductive success between monandrous, multiply mated (to the same male) and polyandrous female dumpling squid (Euprymna tasmanica). Methodology/Principal Findings We mated females in the laboratory with two different males (polyandrous; controlling for mating order), or with a single male (monandrous). To control for mating frequency, we mated monandrous females either once (monandrous 1), or with the same male twice (monandrous 2), and measured reproductive success for each of the three treatments (polyandrous, monandrous 1, monandrous 2). Females mated to two different males produced eggs faster and had larger hatchlings relative to egg mass than females mated once with a single male. Conclusions/Significance The benefits of polyandry demonstrated here are the first, to our knowledge, in any cephalopod. These benefits may outweigh the significant costs associated with mating and help to explain how multiple mating has evolved (or is maintained) in this group.