Thesis (Ph. D.)--Virginia Polytechnic Institute and State University, 1990. / Vita. Abstract. Includes bibliographical references (leaves 185-200). Also available via the Internet.

Thesis (doctoral)--Universität Leipzig, 1907. / Extracted from Zeitschrift für wissenschaftliche Zoologie, 86 Bd., 3 Heft.

Thesis (Ph. D.)--University of Hawaii at Manoa, 2002. / Includes bibliographical references (leaves 218-235). Also available on microfiche.

Thèse : Sciences Naturelles : Paris, Faculté des sciences : 1866. / Titre provenant de l'écran-titre.

Thesis (PhD) -- AUT University, 2008. / Includes bibliographical references. Also held in print (iv, 259 leaves : ill. ; 30 cm.) in the Archive at the City Campus (T 594.5 BOL)

Cuttlefish are renowned for their ability to quickly alter the colour and texture of their skin, for camouflage and communication. This is due to the presence of thousands of pigment-filled sacs, known as chromatophores, which are distributed across the skin. The chromatophores are innervated by motoneurons, which dilate the chromatophores to create the spots, stripes, and other markings, known as chromatic components. There are 34 recognized chromatic components, and it is an interesting question how cuttlefish coordinate the expression of these components to camouflage and communicate. The digital age has introduced new, powerful algorithms and methods to tease out subtle features in the coloration patterns, by means of image registration, segmentation, and identification, as well as methods for modeling the underlying control systems. These tools offer major new insights into the mechanisms of visual perception. In addition, powerful techniques have recently been developed that have yet to be applied to this complex visual motor control system. These methods have large potential in helping discover what features between the pattern and the environment are necessary to prevent detection. Here I present four laboratory experiments, that for the first time use machine learning models, to investigate cuttlefish pattern formation, implementation, and information. The first two experimental chapters investigate how cuttlefish orchestrate their chromatic components for camouflage patterns, and what strategies they employ on diverse backgrounds. I demonstrate that components are expressed more independently than previously believed, finding that the range of patterns expressed lie on a continuum, allowing us to suggest a revised classification scheme for cuttlefish body patterns. The diversity of patterns seem to imply that a cuttlefish could use its repertoire flexibly to display the maximally cryptic pattern for a given background, however I show that cuttlefish to not in fact select a single (possibly optimal) camouflage pattern, continually alter their appearance on a given background, and that the frequency of change increases in relation to the size of the objects in the environment. My third chapter investigated the language-like properties of cuttlefish communication using human speech recognition models. From our subset of cuttlefish patterns, I discovered cuttlefish utilize a lexicon of 10 patterns, with language-like properties such as: they obeyed Zipf's law, contained around 1.6 bits per display, and interestingly, while 2 patterns were visually similar, they were displayed in separate contexts. By implementing a regression onto the patterns, I introduce a basic dictionary of cuttlefish terms and their meaning. From my investigations into cuttlefish intraspecific signaling, I discovered two previously undocumented patterns, used in agonistic encounters between cuttlefish. My final chapter describes these patterns and the contexts they are displayed.

Squids in the family Onychoteuthidae Gray, 1847 have been reported from every ocean but the Arctic, are taken frequently in deep-sea fisheries by catch, and are ecologically important in the diets of many marine predators including cetaceans, pinnipeds, sharks, and seabirds. However, the diversity and systematic of the family have remained poorly understood. Of the 60+ nominal species, 12–14 have generally been accepted in recent studies. Challenges to clarity include insufficient species descriptions, original descriptions published in eight languages and often based solely on early life stages, non-designation or subsequent loss of type material, and the existence of several unresolved species complexes. In light of the general systematic disarray of the Onychoteuthidae, a global revision of the family follows, based on ~1500 specimens examined from 19 repositories. Type material has been examined wherever possible; for some species, photographs of type specimens, original illustrations, and/or the original descriptions have provided the only information available. It has not been possible to fully disambiguate taxa in some cases (e.g. Gen. nov. 2), given the limited material and information available, but for all species treated in this revision (25 out of 26 species; no material was available for Kondakovia nigmatullini), descriptions and illustrations are provided to a consistent standard that will enable their reidentification. External and internal morphological characters and states are described for sub adult to adult stages of most species, with external characters reported through ontogeny as permitted by available material. Historically important characters are treated (general external morphology, body proportions, tentacle clubs, photophores, gladius, lower beak, radula), augmented by several more recently recognised characters (palatine teeth, detailed morphology of the tentacular hooks in adults, tentacular suckers in paralarvae, chromatophore patterns). The systematic value of both historical and new morphological characters at the generic and species levels are discussed; at all ontogenetic stages, tentacular club and hook morphology are considered the most valuable characters, although body proportions and gladius also prove useful. Partial disambiguation of the Onychoteuthis banksii complex has been possible in the Pacific and Atlantic Oceans, resulting in the resurrection of Onychoteuthis bergii Lichtenstein, 1818 and Onychoteuthis aequimanus Gabb, 1868, the description of two new species, Onychoteuthis lacrima and Onychoteuthis prolata (in press), and the expansion of one species’ recognised distribution (Onychoteuthis compacta) to include the Atlantic Ocean. The genus Moroteuthis Verrill, 1881 is considered a junior synonym of Onykia Lesueur, 1821, in accordance with the findings of several earlier authors. However, morphological differences in the species ‘Moroteuthis’ ingens necessitate the resurrection of the subgenus Moroteuthopsis Pfeffer, 1908b, with all other Onykia species placed into a new subgenus, Onykia (Onykia). Sexual dimorphism is reported in the beaks of Onykia (Moroteuthopsis) ingens (new comb.), and revised sex-specific equations are given for estimating this species’ biomass based on LRL. Morphological and historical genetic data suggest a more distant relationship between Onykia and the species ‘Moroteuthis’ knipovitchi Filippova, 1972 than was suggested by earlier classifications. This species is therefore considered to represent an undescribed genus, herein referred to as Gen. Nov. 1, which cannot be more fully diagnosed and described at present due to limited material. The generic position of ‘Onykia’ rancureli (Okutani, 1981) is also uncertain; it may be allied to Walvisteuthis virilis Nesis & Nikitina, 1986 (family Walvisteuthidae Nesis & Nikitina, 1986), but confirmation is impossible without examining type material of W. virilis. A second new genus, Gen. Nov. 2, is therefore described for ‘Onykia’ rancureli and several morphological variants reported from the Pacific and Atlantic Oceans. Given that the majority of available onychoteuthid material was collected after 1950, resulting in the descriptions of over half of the generally accepted genera and species since 1960, ongoing collection programmes are necessary to further resolve onychoteuthid systematic.

Graduation date: 1983

The thesis, given in two volumes, deals with., the population ecology of the macrofaunal mollusc species : Macoma balthica (L. ). Cerastoderma edule (L. ). Mya arenaria(L. ), Retusa obtusa (Montagu) and Hydrobia ulvae (Pennant) studied at fourteen stations on Torry Bay, a large intertidal area in the estuarine Firth of Forth, over the period January, 1975 to February, 1977. These species occupy differing ecological niches within the estuarine ecosystem. The environmental factors at the stations, which represent the range of available habitats in the area, are described in Chapter 2. This chapter also outlines the statistical techniques used in ascertaining the relationships between the factors studied. The population dynamics and magnitude of the growth, flesh condition, production and productivity of each species are given in Chapters 3-5. These chapters also discuss the environmental factors affecting the ecology of the species. Multivariate correlation and regression techniques have been used to describe the factors which influence the species' production ecology. The anomalous production ecology of M. balthica found in the Forth Estuary (Chapter 3) was further studied by a field mark and recapture experiment and multivariate laboratory experiments (Chapter 6). Chapter 6 also discusses the mode of feeding of this bivalve in relation to its ecological preferences and describes a small study on heavy metal pollution within M. balthica. The discussion in Chapter 7 gives general conclusions from the studies and their implications in estuarine productivity. Features common to the population ecology of the five species are discussed and a food web detailing the role of the benthic macrofauna on Torry Bay is given.

Squids in the family Onychoteuthidae Gray, 1847 have been reported from every ocean but the Arctic, are taken frequently in deep-sea fisheries by catch, and are ecologically important in the diets of many marine predators including cetaceans, pinnipeds, sharks, and seabirds. However, the diversity and systematic of the family have remained poorly understood. Of the 60+ nominal species, 12–14 have generally been accepted in recent studies. Challenges to clarity include insufficient species descriptions, original descriptions published in eight languages and often based solely on early life stages, non-designation or subsequent loss of type material, and the existence of several unresolved species complexes. In light of the general systematic disarray of the Onychoteuthidae, a global revision of the family follows, based on ~1500 specimens examined from 19 repositories. Type material has been examined wherever possible; for some species, photographs of type specimens, original illustrations, and/or the original descriptions have provided the only information available. It has not been possible to fully disambiguate taxa in some cases (e.g. Gen. nov. 2), given the limited material and information available, but for all species treated in this revision (25 out of 26 species; no material was available for Kondakovia nigmatullini), descriptions and illustrations are provided to a consistent standard that will enable their reidentification. External and internal morphological characters and states are described for sub adult to adult stages of most species, with external characters reported through ontogeny as permitted by available material. Historically important characters are treated (general external morphology, body proportions, tentacle clubs, photophores, gladius, lower beak, radula), augmented by several more recently recognised characters (palatine teeth, detailed morphology of the tentacular hooks in adults, tentacular suckers in paralarvae, chromatophore patterns). The systematic value of both historical and new morphological characters at the generic and species levels are discussed; at all ontogenetic stages, tentacular club and hook morphology are considered the most valuable characters, although body proportions and gladius also prove useful. Partial disambiguation of the Onychoteuthis banksii complex has been possible in the Pacific and Atlantic Oceans, resulting in the resurrection of Onychoteuthis bergii Lichtenstein, 1818 and Onychoteuthis aequimanus Gabb, 1868, the description of two new species, Onychoteuthis lacrima and Onychoteuthis prolata (in press), and the expansion of one species’ recognised distribution (Onychoteuthis compacta) to include the Atlantic Ocean. The genus Moroteuthis Verrill, 1881 is considered a junior synonym of Onykia Lesueur, 1821, in accordance with the findings of several earlier authors. However, morphological differences in the species ‘Moroteuthis’ ingens necessitate the resurrection of the subgenus Moroteuthopsis Pfeffer, 1908b, with all other Onykia species placed into a new subgenus, Onykia (Onykia). Sexual dimorphism is reported in the beaks of Onykia (Moroteuthopsis) ingens (new comb.), and revised sex-specific equations are given for estimating this species’ biomass based on LRL. Morphological and historical genetic data suggest a more distant relationship between Onykia and the species ‘Moroteuthis’ knipovitchi Filippova, 1972 than was suggested by earlier classifications. This species is therefore considered to represent an undescribed genus, herein referred to as Gen. Nov. 1, which cannot be more fully diagnosed and described at present due to limited material. The generic position of ‘Onykia’ rancureli (Okutani, 1981) is also uncertain; it may be allied to Walvisteuthis virilis Nesis & Nikitina, 1986 (family Walvisteuthidae Nesis & Nikitina, 1986), but confirmation is impossible without examining type material of W. virilis. A second new genus, Gen. Nov. 2, is therefore described for ‘Onykia’ rancureli and several morphological variants reported from the Pacific and Atlantic Oceans. Given that the majority of available onychoteuthid material was collected after 1950, resulting in the descriptions of over half of the generally accepted genera and species since 1960, ongoing collection programmes are necessary to further resolve onychoteuthid systematic.