OT7 MALACOLOGIA, 1988, 29(1): 153-193 BEHAVIOR AND SYSTEMATICS OF CEPHALOPODS FROM LIZARD ISLAND, AUSTRALIA, BASED ON COLOR AND BODY PATTERNS Clyde F. E. Roper1 & F. G. Hochberg2 ABSTRACT Cephalopoda were observed in situ and under laboratory conditions at Lizard Island, Great Barrier Reef, Australia. Observations on habitat, foraging and activity patterns are included. The major chromatic components and body patterns are described for Octopus cyanea, O. ornatus, Hapalochlaena spp., Metasepia pfefferi and Sepia papuensis. Components of body pattern include color, texture, posture and locomotion. A remarkable new type of locomotion, "ambling," is described for M. pfefferi. This is the first description of living M. pfefferi and S. papuensis. On the basis of body patterns, behavior and morphology, the elevation of the subgenus Metasepia to generic status is confirmed. Observations of live Hapalochlaena at Lizard Island and in Sydney and color photographs of live animals from several other localities confirm the existence of a widespread complex consisting of at least three species and support the validity of the genus. Based on observations of live animals and a systematic evaluation of preserved specimens, the presence of Octopus ornatus is reported in Australian waters for the first time. Key words: Octopus; Sepia; Metasepia; Hapalochlaena; cephalopods; color patterns; behav- ior; systematics; field observations; Great Barrier Reef. INTRODUCTION An International Workshop on Molluscs was conducted on Lizard Island, Australia, from 2 to 14 December, 1975 (Ponder, 1979).3 The Workshop was sponsored by the Australian Museum, Sydney, which operates the Lizard Island Research Station located near the northern end of the Great Barrier Reef, Queensland, at 14?40'S, 145?28'E (Fig. 1). Lizard Island is a continental island about 2.9 square km in area composed primarily of granite; it lies about 30 km off the coast and is 17 km from the outermost barrier reefs (Pon- der, 1979). Participants included malacolo- gists from Australia, Great Britain, Hong Kong and the United States. One of us (C.F.E.R.) participated in the Workshop with the objec- tive of surveying the cephalopod fauna around Lizard Island. Cephalopods were col- lected in various habitats and live animals were observed in their natural habitat and in aquaria. A preliminary checklist with collec- tion and habitat data for the 27 species of cephalopods collected at Lizard Island was published separately (Roper & Hochberg, 1987). This paper presents observations on be- havior and body patterning made in the field and in laboratory aquaria on five species of cephalopods: Octopus cyanea Gray, 1849; O. ornatus Gould, 1852; Hapalochlaena cf. maculosa (Hoyle, 1883); Metasepia pfefferi Hoyle, 1885, and Sepia papuensis Hoyle, 1885. One Hapalochlaena cf. fasciata (Hoyle, 1886) was maintained in an aquarium at the Australian Museum, Sydney, subsequent to the Workshop. For the purpose of future identification, a synopsis is provided in which species characters are diagnosed and infor- mation on distributions and life histories are summarized. In addition to color and body patterning, observations are included on for- aging, resource partitioning and activity pat- terns. We regard these as preliminary observa- tions that were made in 1975 in a fortuitous and opportunistic manner prior to the formu- lation of a classification of behavior that char- acterizes more recent studies of color and 'Department of Invertebrate Zoology, Division of Mollusks, National Museum of Natural History, Smithsonian Institution, Washington, D.C. 20560, U.S.A. ^Department of Invertebrate Zoology, Santa Barbara Museum of Natural History, 2559 Puesta del Sol Road, Santa Barbara, California 93105, U.S.A. 3This paper is designated a contribution of the Lizard Island Research Station. (153) 154 ROPER & HOCHBERG North Point J To Yonge Reef 'W/ PRINCE CHARLES ISLAND ? Rocky Point ?# Casuarlna Beach Watsons Bay ? ?? ? ??? N Pidgin Point Research Point Freshwater Beach e Coconut Beach a Blue Lagoon Lizard Head ? + .03 BIRD ISLAND Bommle No. 2 PALFREY ISLAND Q- 1 km. SOUTH ISLAND FIG. 1. Map of Lizard Island, Australia (14? 40'S, 145? 28'E) showing adjacent islands and locations of collecting stations (solid dots). See Roper & Hochberg (1987) for station and habitat data. body patterning in squids, octopuses and cuttlefishes; see especially Holmes (1940), Packard & Sanders (1969, 1971), Moynihan (1975, 1985), Moynihan & Rodaniche (1977, 1982), Packard & Hochberg(1977), Hanlon & Hixon (1980), Boyle & Dubas (1981), Hanlon (1982), and Hanlon & Messenger (1988). Two papers are especially significant in relation to our work. Packard & Hochberg (1977) defined and summarized the hierarchy of anatomical and behavioral systems that lead to the generation of patterns in Octopus and other genera. Hanlon & Messenger (1988) present a comprehensive and detailed study of body patterns and behavior in Sepia officinalis. To the extent possible we have attempted to follow the concepts and termi- nology presented in these two papers, which we consider the standards for modem work. Both papers emphasize that these magnifi- cent and complex animals are capable of providing an almost infinite number of combi- AUSTRALIAN CEPHALOPOD BODY PATTERNS 155 nations and gradations of colors, textures, postures and body patterns. Two of the gen- era we worked with, Hapalochlaena and Metasepia, are so different that we could not always fit our observations into existing termi- nology and hence have introduced several new terms. The majority of detailed research on color and body patterning in cephalopods has been done on species from the Mediterranean and Europe, the Caribbean and Gulf of Mexico, the west coast of North America, Panama, Hawaii, Palau and Guam. Other than photo- graphs of blue-ringed octopuses and occa- sionally of other octopuses and cuttlefishes that have appeared in popular magazine arti- cles, little information is available on the biol- ogy, behavior and body patterns of Australian cephalopods. This paper describes for the first time color and body patterns and other aspects of be- havior of living Hapalochlaena spp., Mefa- sepia pfefferi and Sepia papuensis. It also expands the observations made on Octopus cyanea and O. ornatus in Hawaii (e.g. van Huekelem, 1966, 1973, 1983; Wells & Wells, 1970) and on H. cf. fasciatus in Australia (e.g. Tranter & Augustine, 1973, as H. maculosa). In addition, we describe for the first time "ambling" in M. pfefferi, a newly recognized mode of locomotion for sepiid cephalopods. MATERIALS AND METHODS Observations and collections were made in the intertidal zone (primarily during low tides at night) and on the patch, fringing and barrier reefs by skin and SCUBA diving. Thirty-seven stations were occupied at Lizard Island lor collection and observation during the Work- shop. See Fig. 1 for the location of collecting sites. In the text, stations occupied by the senior author are indicated by the abbrevia- tion CFER. For a complete list of stations and for a checklist of all cephalopods recorded from Lizard Island, see Roper & Hochberg (1987). Station data, diving logs, notes on field and laboratory observations are con- tained in a notebook on file at the National Museum of Natural History, Washington, DC. Cephalopods collected for observation were returned to the Lizard Island Research Station where they were maintained in aquaria supplied with running seawater. Glass-walled aquaria varied from 20-30 liter capacity tanks for small individuals of Hapalochlaena cf. maculosa, Metasepia pfefferi and Sepia papuensis to a large 150-200 liter tank that housed Octopus cyanea and O. ornatus. Aquaria contained fine, light-colored sand, coral rubble or other material to provide a resemblance of habitat for each species. Activity and behavior of all animals were observed both during the day and at night. Results of observations were recorded or sketched in a notebook (day) or tape recorded (night) for later transcription. Photographs, using Kodachrome 64 film, were taken with a Nikon F 35 mm camera, 55 mm macro lens and one or two Braun electronic flash units. Observations on a live Hapalochlaena cf. fasciata were made follow- ing the Workshop at the Australian Museum, Sydney. The animal was kept for two weeks in a 20 liter aquarium where it was studied and photographed. Following observation, cephalopods were fixed in 8% buffered sea water formalin. Voucher specimens of all species discussed in this paper are deposited in the Department of Invertebrate Zoology?Mollusks, National Museum of Natural History, Washington, D.C. or in the Department of Malacology at the Australian Museum, Sydney. The species, number of individuals and museum catalog numbers of these vouchers are: Octopus cyanea (2 specimens), USNM 816646 and 816647; Octopus ornatus (2 specimens), USNM 816649 and 816650; Hapalochlaena cf. maculosa (4 specimens), USNM 730598, 730599 and 816623; Hapalochlaena cf. fasciata (1 specimen), Australian Museum; Metasepia pfefferi (14 specimens), USNM 816620 and 816621; Se- pia papuensis (2 specimens), USNM 816619. While most of the terminology we use is adapted from other works (see especially Packard & Hochberg, 1977, Hanlon & Mes- senger, 1988) some terms need definition here. Chronic patterns are long-term (hours) patterns that allow an undisturbed animal to blend in with the substrate or background (crypsis or concealment). Acute patterns are short-term (only seconds or a few minutes) patterns produced in many cases in response to a disturbance. These patterns are striking or vivid in expression and typically stand out in bold contrast to the background. Acute patterns may take a number of forms, among which we define the following: 1) passing cloud(s)?conspicuous, pulsat- 156 ROPER & HOCHBERG ing flushes of dark color that pass in an amorphous front, unidirectionally across the body (see Packard & Hochberg, 1977). 2) passing wave(s)?distinct sequence of well-defined bands that pass like a set of wave fronts over the dorsal surface of the body. We introduce this term to distinguish this pattern from passing clouds, since the wave sets move both anteriorly and posteri- orly at the same time. 3) play(s) of color?small flushes of dark color that can appear randomly anywhere on the body and radiate out from a point source (see van Heukelem, 1966). 4) flash?instantaneous expansion of chromatophores that highlights or darkens specific components of body patterns such as the ocelli, maculae of Hapalochlaena and mating stripes of Octopus cyanea. When ex- pressed these components often appear to pulsate or flash. 5) flush?instantaneous expansion of chromatophores that can uniformly darken the entire body or when directed toward an interacting animal or disturbance can darken the body unilaterally or just dorsally (see Fig. 61). 6) blanch?instantaneous retraction of chromatophores that uniformly pales or whit- ens the entire body as in the deimatic (the "dymantic display" of Packard & Sanders, 1969) and mating patterns (see photographs in van Heukelem, 1970); the opposite of flush. The above acute patterns may be single events (as in a flush or blanch) or multiple (as in passing waves or flashes). Throughout we define continuous, unbro- ken lines that are oriented transversely as bands and those that run longitudinally as stripes. Bars are broken or interrupted bands and streaks are broken or interrupted stripes. Diagonals are short lines that are oriented at angles oblique to bars and streaks. Maculae are spots of dark chromatophores that sur- round the iridescent blue rings of Hapa- lochlaena. Unlike ocelli or eye spots in some Octopus species, the diameters of the maculae are not fixed but are capable of expanding or contracting. Primary papillae are the largest and most conspicuous of the papillae on the body and often are erected for long periods of time. The distribution of primary papillae is fixed morphogenetically and can be used as a diagnostic feature at the genus level. A vari- ety of characteristic shapes may be ex- pressed such as simple conical, compound bifid, compound papillate ridge, flat-truncated flaps, etc. Secondary papillae are smaller and expressed only intermittently. They generally are all simple and conical in shape. Cephalopods, in particular cuttlefishes, swim by means of fin undulations, float when buoyancy is controlled by the cuttlebone, hover when they gently pump water through the funnel and jet when they forcibly pump water through the funnel. Octopuses scuttle when they move across the bottom using their arms. KEY TO ABBREVIATIONS adep, anterodorsal eye papilla admp, anterodorsal mantle papilla ads, arm dark stripe af, ambulatory flap apwl, anterior passing wave, first wave apw2, anterior passing wave, second wave avep, anteroventral eye papilla aws, arm white spot dep, dorsal eye patch dhdf, dorsal head dark field dhws, dorsal head white spot dhwt, dorsal head white triangle dmdf, dorsal mantle dark field dmds, dorsal mantle dark spot dmlf, dorsal mantle light field dmp, dorsal mantle papilla dmwb, dorsal mantle white bar dmws, dorsal mid-mantle white spot flp, finline papilla flws, finline white stripe fws, frontal white spot fwsp, frontal white spot papilla Idmp, laterodorsal mantle papilla Ihlf, lateral head light field Imws, lateral mantle white spot mdmf, mid-dorsal mantle flap mtpr, mantle tip papillate ridge mwbf, mantle white bar flap mwsp, mantle white spot papilla pdep, posterodorsal eye papilla pdmf, posterodorsal mantle flap pdmp, posterodorsal mantle papilla pdmpr, posterodorsal mantle papillate ridge phws, posterior head white spot pmds, posterior mantle dark spot pmws, posterior mantle white spot ppwl, posterior passing wave, first wave ppw2, posterior passing wave, second wave ppw3, posterior passing wave, third wave AUSTRALIAN CEPHALOPOD BODY PATTERNS 157 vepr, ventral eye papillate ridge wws, web white spot I, dorsal arm II and III, lateral arms IV, ventral arm 1, primary 2, secondary OBSERVATIONS, SYNTHESIS AND DISCUSSION 1. Octopus cyanea Gray, 1849. Common name: Cyane's octopus. "Big blue octopus," the common name normally used, is a misnomer resulting from an incor- rect interpretation of the specific name. Gray (1849: 15) named the species "Cyanea,"the capital "C" denoting a patronym referring to Cyane in Greek mythology, a nymph of Persephone who was turned into a fountain. Had Gray intended to refer to the color blue, he would have had to use the word "cyaneus," uncapitalized. Hence, the correct common name should be "Cyane's octopus." This is often called the "day octopus" in Hawaii. A. Synopsis Diagnosis: Body globose, muscular, man- tle length to 100 mm, total length to 1200 mm, total weight to over 5 kg; skin smooth to heavily papillate, with two conspicuous web ocelli; eyes large; arms medium length, subequal, 4-5 times mantle length, thick and muscular; arm formula typically IV.I.II.III; en- larged suckers on all arms of males, espe- cially conspicuous on arms I and II; gills with 9-10 lamellae per demibranch; right arm III hectocotylized, length 75-80% of left arm III; end organ minute, 0.5-1.5% of hectocotylized arm length; ligula bluntly pointed, open with low inrolled edge, groove with faint ridges, calamus small; eggs small, 2.5-3.0 mm long; hatchlings planktonic. Distribution: Widespread in tropical wa- ters of the Indo-Pacific from Hawaii through the Pacific Islands to Australia, through the Indian Ocean to East Africa and the Red Sea. One of the most common shallow-water oc- topuses in New South Wales and Queens- land, Australia. Found in rocky or coral reef habitats from the intertidal zone to 45 m. Life history: Reviewed in van Heukelem (1983). This large octopus is commercially important in Hawaii and elsewhere in the South Pacific. References: Berry, 1914; Le Souef & Al- len, 1933, 1937; Boone, 1938; Dew, 1959; Vevers, 1961, Young, 1962; van Heukelem, 1966, 1970, 1973, 1979, 1983; Maginniss & Wells, 1969; Wells & Wells, 1969, 1970, 1972a, b; Yarnall, 1969; Houck, 1982; and Young, Harm an & Hochberg, in preparation. B. Field observations Two animals of O. cyanea were encoun- tered during the study period; both inhabited dens in cemented coralline rock on the shal- low reef flat directly offshore from the Re- search Station on Casuarina Beach. The reef flat habitat is described under the section on O. ornatus. Animal 1 was captured at night during low tide (CFER-18; refer to Roper & Hochberg (1987) for station data). It was placed in an aquarium and observed for 12 days. Animal 2 was discovered during a day- time dive on the reef flat at high tide (CFER- 27). In situ observations on animal 2 contin- ued for nine days. All in situ observations on animal 2 were made during daytime within three hours be- fore or after high tide. The den, located in a pile of rubble and cemented coralline rock, was elevated about 60 cm above the sur- rounding reef flat. The entrance was littered with a midden of mollusk shells, crustacean parts and coralline pebbles. When examined three days later the den was empty. An intense search along the contour of the reef flat revealed an occupied den about 20 m N of the first den. Although the occupant from den 1 had not been tagged and no unusual mark- ings had been noted, the animal in den 2 was very similar in size and appearance. On the last day of the Workshop, den 1 was still vacant. Den 2 also was empty, although a pile of fresh shells littered the front entrance. Further searching located a third den 30 m along the reef flat N of den 2. It was occupied by a large O. cyanea that appeared to be the same as the individual observed during the preceding week. Octopus cyanea is a transient den dweller that moves periodically from den to den along the reef in search of food or in response to disturbance (van Heukelem, 1966, 1983). Yarnall (1969) and van Heukelem (1966) re- ported that O. cyanea may use its den for up 158 ROPER & HOCHBERG to one month or more. In this regard it resem- bles the behavior of O. vulgaris (Hochberg & Couch, 1971) and O. bimaculatus (Ambrose, 1982) (see Table 3). Octopus cyanea is active during morning and early evening hours and remains quiescent in its den during the day and at night (Houck, 1982). Loch (1980) dis- cussed the hole-drilling technique used pos- sibly by O. cyanea to feed on three species of Cypraea. Mating, egg laying and brooding have been reported by van Heukelem (1970, 1983) and Le Souef & Allan (1933, 1937). Although a large number of dives was made by the Workshop participants in the reef flat area adjacent to the Research Station, only two O. cyanea were observed. In con- trast, over 30 individuals of O. ornatus were observed in the same area during only three excursions onto the reef flat at night during low tide. O. ornatus, like O. macropus, appar- ently is a more free-ranging species that is active at night (van Heukelem, 1966; Houck, 1982) (see Table 3). C. Components of body patterns Octopus cyanea is a robust, medium-sized species similar to O. vulgaris or O. bi- maculatus. The body is heavy set and the arms relatively short and muscular. When occupying its den, O. cyanea assumes one of two body postures that we term the "lookout" (Figs. 2-4) and the "guard" (Fig. 5) postures. In the lookout position the head and eyes are raised out of the entrance of the den with the eyes greatly protruded, so the animal can monitor activities around the den (Figs. 2-4). In this posture mantle, head, web and arms are covered densely with both flat-truncate and conical, cream-colored papillae that give the animal a very rugose appearance that blends well with the surrounding habitat. A single, large compound papilla generally is erected over each eye (Figs. 2, 3); three or four additional simple dorsal eye papillae also may be present. The entire body is covered with a light mottle of reddish purple/maroon on a beige/cream background. Two large ovate ocelli ("eye spots") often are visible on the web just below (anteroventral to) the eyes (Fig. 31). The aboral and lateral surfaces of the arms are banded with dark patches of red interrupted by small to medium-sized white spots. The papillate skin and coloration match the irregularly textured and mottled appear- ance of the substrate. When approached closely or disturbed the animal abandons the lookout posture.flushesadarkmottledmaroon/ red with alternating dark maroon and cream lines or rays radiating around the eyes (Fig. 5) and then retreats into the den. In the "guard" posture the octopus is with- drawn into the den and sits sideways just in- side the entrance. The eye (either right or left) is erect and peers out over the second and third arms that are extended across the en- trance and turned outward so that the largest suckers are exposed and visible (see Fig. 5). In this position the octopus can effectively guard the entrance visually; it also can grasp and test with the exposed suckers anything that approaches the entrance to the den. In the guard position the entire animal is covered with primary cream-colored papillae, both sim- ple, conical as well as flat-truncate papillae. The primary papilla over each eye is leafy and flat and may bear three or four secondary pa- pillae (Fig. 2); primary compound papillae also occur anterior to the eyes, in the mid-line at the junction of the web and head, and one each at the base of the arms. Typical cryptic coloration is a light mottle of red/brown on a cream/beige background. A dark black band extends along the head and through the eye effectively mask- ing it. When disturbed the animal flushes a dark mottle and the eye mask changes to the radiating pattern of light cream and dark red lines mentioned above. Upon repeated distur- bance the animal flushes a uniform dark red. The dark flush starts at the head and radiates down the arms and mantle until the animal is a uniform dark color. When seen in the open or when disturbed enough to leave the den, the animal jets across the bottom. When jetting, the body texture is smooth to granular without conspic- uously erect papillae, and the color is light cream mottled with red patches (Fig. 6). When approached in the aquarium by O. ornatus, the captive O. cyanea exhibited a typical response in which the body reared up and the arms flared out to present the en- larged suckers. This is equivalent to the "fighting display" of Packard & Sanders (1971). The animal changed from a cryptic color pattern to a uniform dark flush. Such posturing effectively drove off the O. ornatus. An inventory of the currently recognized components exhibited by O. cyanea is pre- sented in Table 1. The data are based on our observations of live animals at Lizard Island and on published reports, figures and photo- graphs. AUSTRALIAN CEPHALOPOD BODY PATTERNS 159 FIGS. 2-5. Octopus cyanea. Figs. 2, 3: Frontal view (2) and left lateral view (3); lookout posture with raised flat, truncate dorsal eye papillae. Fig. 4. Frontal/lateral view, left side; lookout posture without raised papillae. Fig. 5. Left lateral view; guard posture with radiating dark and light lines around eye, and exposed suckers. Photographs by W. F. van Heukelem (Honolulu, Hawaii). 160 ROPER &HOCHBERG FIG. 6. Octopus cyanea. Right lateral view, jetting in forward direction over the bottom in light color phase without erect papillae. Photograph by W. F. van Heukelem (Honolulu, Hawaii). D. Discussion The observations on O. cyanea at Lizard Island basically confirm those of other work- ers (especially van Heukelem, 1966,1983) in that the species is active during crepuscular periods and quiescent during both night and day. A widely-spaced, transient den-dweller, O. cyanea on Lizard Island ranges along the reef flat, occupying one den for a few days to several weeks before moving on to the next. Niche and food resource partitioning are dis- cussed in the following section on O. ornatus. Van Heukelem (1983) stressed that O. cyanea is capable of showing a large variety of color patterns, textures and postures but that a detailed inventory had not been formu- lated. Illustrations and brief notes on color and body patterns are in the literature: sexu- ally mature adults (van Heukelem, 1966, 1983; Wells & Wells, 1972b), brooding fe- males (Le Souef & Allan, 1933,1937), hatch- lings (Le Souef & Allan, 1937; Dew, 1959) and newly settled juveniles (Wells & Wells, 1970). Hawaiian specimens of O. cyanea were described and figured by Hoyle (1885a, 1886, see also Berry, 1914) under the name O. marmoratus, which refers to the color pattern of "ochreous red maculated with pur- ple" and to the series of "intercotyledonary color bands down the surface of the arms." Taki (1964) provided additional notes on color of a species he described as Cal- listoctopus magnocellatus but this is now known to be a synonym of O. cyanea. Taki's species name refers to the presence of a large ocellus that he described as having three parts: a black center 22 mm in diameter, a pale ring and an outer black ring 3-4 mm wide. The total diameter measured 40 mm. In other octopuses the ocellus also is known to contain species specific patterns of chroma- tophores and iridophores (Hochberg, in prep- aration); this character needs further detailed documentation. The best records of the color and patterns of this species are photographs such as those in van Heukelem (1970, 1983), Voss (1971), Roessler (1977) and Travieso (1978). Of all the patterns of O. cyanea, perhaps the most colorful and interesting are the dramatic courtship and mating patterns. Although not AUSTRALIAN CEPHALOPOD BODY PATTERNS 161 observed at Lizard Island these have been photographed and well documented in Hawaii by van Heukelem (1966, 1970, 1983) and by Wells & Wells (1972b). To prepare a complete inventory of pat- terns in O. cyanea, extensive field and labo- ratory observations of young and adult ani- mals are needed. In addition, further compar- isons of patterns need to be made in widely separated geographic populations. 2. Octopus ornatus Gould, 1852. Common name: white striped octopus. This common name is used to emphasize the pattern of conspicuous white markings on the mantle. This is often called the "night octo- pus" in Hawaii. A. Synopsis Diagnosis: Mantle globular to elongate, muscular, mantle length to 120 mm, total length to 1000 mm, total weight to 500 g; skin granular to rough and warty, purplish red, with conspicuous pattern of white markings on the mantle and elongate oval or round white spots on arms; eyes large; arms very long, attenuate at tips, thick and muscular proxi- mally, 6-8 times mantle length; arm formula I.II.III.IV; enlarged suckers on arms I of males; gills with 12-14 lamellae per demibranch; right arm III hectocotylized, length 60-75% of left arm III; end organ medium sized, 4.5-8% of hectocotylized arm length; ligula elongate, pointed, inrolled edges, groove smooth to faintly striated, calamus small; eggs small, 2-4 mm long; hatchlings planktonic. Distribution: Widely distributed in tropical waters; Indo-Pacific from Hawaii through the Pacific islands to Australia, into the Indian Ocean to East Africa; common shallow water species, free ranging through reef flat areas, from intertidal to 15 m. This is the first con- firmed report of the species in Australia. Known currently only from the Great Barrier Reef, northern Queensland. Life history: Life span not known. Large numbers of small eggs are laid. Larvae are planktonic upon hatching. Feeds principally on small crustaceans. Active at night. Free ranging predator, without fixed dens. References: Gould, 1852; Berry, 1914; Boone, 1938; Taki, 1964; van Heukelem, 1966; Yarnall, 1969; Voss, 1981; Houck, 1982; and Young, Harman & Hochberg, in preparation. B. Field observations Thirty individuals of O. ornatus were found in a 20 x 100 m area of the reef flat imme- diately offshore from Casuarina Beach and were observed during three low tide surveys conducted on the reef on moonless nights. At low tide much of the reef flat was ex- posed leaving a network of tide pools that varied in size from small shallow puddles to pools several meters across with depths to 15-20 cm. The bottom of the tide pools consisted of coarse coralline sand and small pieces of coral rubble. The reef flat habitat consisted of scattered heads of dead coral, coralline algae and rubble, and it appeared to be ideal for O. ornatus. The many holes and crevices provided numerous places of protec- tion for resting octopuses, as well as refuge for prey animals, particularly crabs. Three individuals were observed scuttling over open sandy bottom immediately shore- ward of the reef flat, but all others were associated closely with the reef flat pools. One individual hunted by scuttling slowly over the reef flat while the long, sinuous arms were engaged simultaneously in search of prey. The arms were fanned out in all directions, exploring every nook and crevice in the pool. Several arms investigated deep into holes, while others swept under rocks. During the observation period this individual captured two small crabs (unidentified) that immedi- ately were transferred to the web and later devoured. Although frequent dives were made in the reef flat area during the 12-day period of the Workshop, O. ornatus never was observed during the daytime, nor were any found at rest in dens. We assume the animals either hide deep in the reef complex or bury themselves in the sand where they remain quiescent all day. This is in sharp contrast to O. cyanea, a crepuscular den dweller, and is similar to the behavior of O. macropus (Hochberg & Couch, 1971) (see Table 3). C. Components of body patterns. The mantle of O. ornatus is elongate and pointed posteriorly. The arms are extremely long, slender and attenuate. The typical cryp- tic body pattern in the field is a low-intensity, light, brownish-red mottle. The mantle, head, web and arms are covered with light cream to white colored markings (Fig. 31), the vivid- ness of which is controlled by overlying dark 162 ROPER &HOCHBERG TABLE 1. The components of body patterns in Octopus cyanea. A. Light B. Dark C. Other arm white spots2 dark arm bands ocelli dorsal head white stripe dark arm stripes1 dorsal mantle white stripe2 dark arm tips1 white mantle (blanch)1 dark uniform mantle1 white web (blanch) dark anterodorsal mantle patch1 light mantle bands dark head1 dark eye stripe dark eye region1 radiating eye lines II. Textural components A. Primary papillae B. Secondary papillae C. Other (compound, flat) (simple, conical) granular dorsal eye papillae (1/eye) dorsal eye papillae (3 or 4/eye) smooth mantle papillae uniform body papillae arm base papillae (1/arm) III. Postural components radiating arms standing1 curled arms lookout (= alert1) coned arms guard flared arms submissive: raised head & eyes male/male1 enlarged sucker presentation male/female1 sucker shield' flared web IV. Locomotor components and maneuvers sleeping prey drilling resting/sitting inking scuttling ritualized fighting1 (= territorial defense) jetting distance copulation14 escape (body first) courtship strut1 stalking (arms first)1 bobbing1 hunting/prey capture escape (body first) speculative pounce1-3 grooming1 attack jump burying food gathering under web V. Body patterns A. Chronic B. Acute light cryptic mottle (grey, light brown) dark conflict mottle light uniform dark uniform (red or brown flush) sleeping1 light uniform (blanch): deimatic (white w/ocelli1) copulatory (white w/o ocelli1) courtship stripes (male)1" dominant male mantle stripes1 flamboyant1 plays of color1 'Van Heukelem (1966, 1970, 1983) zRoessler (1977) 3Yarnall (1969) "Wells &Wells (1972b) AUSTRALIAN CEPHALOPOD BODY PATTERNS TABLE 2. The components of body patterns in Octopus ornatus. 163 I. Chromatic components A. Light B. Dark C. Other dorsal mantle white stripes or dark uniform bluish-green iridescence streaks dorsal mantle white spots head white spots frontal white spots arm white spots (2 rows/arm) II. Texturai components A. Primary papillae B. Secondary papillae C. Other compound bifid mantle tip papillae Simple uniform body papillae granular simple lateral mantle tip papillae smooth compound bifid dorsal eye papillae (1 or 2) simple frontal white spot papillae (1/white spot) simple arm base papillae (1/arm) III. Postural components radiating arms coned arms elongate pointed mantle IV. Locomotor components and maneuvers resting/sitting scuttling jetting inking hunting w/arms V. Body patterns A. Chronic B. Acute light cryptic mottle dark conflict mottle dark uniform (flush) deimatic chromatophores. White spots on the arms are squarish with rounded corners and arranged in two regular rows; they decrease in size to the tips of the arms. Spots on the web and head are oval and they become elongate interrupted stripes or streaks on the mantle. When a light is shone on the animal at night the white spots, stripes and light patches on the lateral surfaces of the suckers and the cir- cumference of the suckers are visible as a vivid bluish-green iridescence (see comments by Roper & Young in Voss, 1981: 533). Body patterns observed in the field appeared iden- tical to those seen in the aquaria and are described below. All laboratory observations were on animals captured at CFER stations 18 and 20. During the night, inactive or resting animals sit in the typical octopus posture with the head and mantle lying horizontally over the relaxed, radiating arms and web. The mantle is finely granular, while the head and arms are smooth. The posterior tip of the mantle has several papillae. A large, bifid papilla is erect at the very tip of the mantle during jetting, while the rest of the body is smooth. Two or three simple, secondary papillae occur along each side of the lateral tip of the mantle. Above each eye is one or two low, broad, bifid papillae. A papilla also is present anterior to the eyes in the center of each white spot on the web at the base of each arm. Resting animals have a light to medium dark cryptic mottle with large subdued white spots on the head, web and arms. White markings are not visible on the dorsal mantle. When disturbed 164 ROPER & HOCHBERG TABLE 3. Comparison of factors for coexistence of Octopus cyanea and O. ornatus. Factor/species O. cyanea O. ornatus Body form medium sized, robust small, sleek Arms robust, normal length very long, attenuate Activity period crepuscular, dawn and dusk nocturnal Dwelling "territorial" in fixed den sites, temporary shelters, changed daily, periodic transient "non-territorial" Hunting sites in immediate home range over broad range Prey mollusks (secondarily crabs) crabs Mode of life reclusive, solitary nomadic Relative abundance low high Comparable species 0. vulgaris (Mediterranean, O. macropus (Mediterranean, in other areas Caribbean) Caribbean) O. bimaculatus (California, Mexico) O. alecto (Mexico) the animal first darkens the mottle and then produces a uniform dark flush that covers the entire body. All arms are engaged in locomotion when the animal scuttles rapidly across the bottom at night. During this type of locomotion the mantle is elongate and pointed posteriorly. The entire body is smooth except in the posterior region where several primary papil- lae are erect. The animal shows a darkened mottle in which the white markings are visible. While the animal is at rest during daytime, the coloration is a deep brownish red reticu- lation with subdued white markings. Occa- sionally three small white spots may show on the dorsal mantle, one in the midline just posterior to the junction of the mantle and head and two (the dorsal mantle white spots) lateral to the midline and in the anterior third of the mantle. When disturbed, O. ornatus jets mantle- end first across the bottom. The mantle is elongate and pointed and the arms are brought together in a cone. The body is streamlined with the skin entirely smooth ex- cept for the above-described primary papillae erected on the posterior tip of the mantle. Color is a light to dark mottle to nearly uniform dark with vivid white markings. Table 2 presents the currently recognized components of body patterns of O. ornatus based upon our observations and a compila- tion from published reports and photographs. D. Discussion Octopus ornatus has been studied primarily in Hawaii (van Heukelem, 1966; Houck, 1982). The Lizard Island animals represent the first records of this species in Australian waters, and our observations confirm those made in Hawaii. On Lizard Island, animals are conspicuously abundant at night as they range across the reef flats actively hunting cryptic prey, primarily crabs, with very long, attenuate arms. During the day the animals retreat to temporary shelters of convenience, rather than to any permanent den site, or they may burrow into the sand. The observations on O. ornatus and O. cyanea living sympatrically on a small area of patch reef suggest a system of niche and food resource partitioning. Table 3 presents the currently recognized morphological and eco- logical characteristics that permit these two species to coexist in the same habitat. The existence of ecomorphs has been demon- strated in other groups. We suggest that this phenomenon occurs between O. cyanea and O. ornatus and may occur in a number of sympatric octopod species in various parts of the world. Little information is available on the color, texture and body patterns of O. ornatus. A color plate by J. Drayton appeared in the original description by Gould (1852) along with notes on the color of the living animal. Taki (1964) illustrated his description of Cat- Hstoctopus asakawai (now regarded as a syn- onym of O. ornatus?see Voss, 1981) with a color painting of a preserved specimen and several black and white photographs of live animals. Additional brief references to color and texture are noted in Berry (1914), Boone (1938), van Heukelem (1966) and Voss (1981). The majority of these papers concen- trate on the striking pattern of white spots or stripes but do not provide other color or pattern observations. Van Heukelem (per- sonal communication) reported differences in AUSTRALIAN CEPHALOPOD BODY PATTERNS 165 basic ground color (from deep orange to dark purplish brown to deep reddish brown) that appear to have a geographic or popula- tional basis. This may be an artifact of obser- vation and preservation or an indication that a complex of cryptic species may be involved. Van Heukelem's photograph (1970) of O. ornatus in Hawaii is reproduced here (Fig. 31) for reference to the conspicuous pattern of white markings of this species. O. ornatus closely resembles O. macropus (see Hanlon, this volume, for comparison photograph), a species that has white spots instead of stripes on the mantle. O. macropus has been re- ported in Australia (Lu & Phillips, 1985). Care- ful study and comparison of O. ornatus and O. macropus should be made, since in many parts of the world their distributions appear to overlap. 3. Hapalochlaena cf. maculosa (Hoyle, 1883) Common name: Lesser blue-ringed octo- pus. We introduce this name to draw attention to the very small blue rings (1-2 mm in diameter) on the mantle, head, web and arms. A. Synopsis Diagnosis: Mantle small, elongate, ovoid, pointed posteriorly; mantle length to 30 mm; total length to 80 mm; total weight to 40 g; skin wrinkled or densely covered with papil- lae, with conspicuous pattern of dark maculae and small iridescent blue rings (1-2 mm in diameter) on dorsal mantle, head, web, and arms, ventral head and mantle without rings; eyes large and prominent; arms short, subequal, 1.5 to 2 times mantle length; typical arm formula III.II = IV.I; suckers small, nu- merous, none enlarged; gills with 6 or 7 lamellae per demibranch; right arm III hectocotylized, length 70 to 80% of left arm 111; end organ medium size, 4 to 6% of hectocotylized arm length; ligula smooth; calamus small; egg size unknown. Distribution: This species appears to be widespread, perhaps as a complex of spe- cies, all around Queensland, Victoria, Tasma- nia, South Australia and Western Australia. In tropical waters it occurs close to coral reefs in shallow water to 25 m, on sandy silt bottom, apparently in association with attached green algae. In temperate waters it occurs in rock reefs and rock rubble areas, tide pools, mol- lusk shells, bottles and cans. Life history: Unknown. References: See H. cf. fasciata. B. Field observations Four individuals of Hapalochlaena cf. maculosa were collected during the day in Mrs. Watson's Bay. Animal 1 was discovered in 14 m (CFER-25) and animal 2 in 20-22 m (CFER-28). Both were found on open sand/ silt bottom devoid of rocks but scattered with coral rubble and living solitary corals. Con- spicuous fauna and flora included numerous black holothurians, probably Holothuria atra or H. edulis, scattered patches of attached green algae, Caulerpa cupressoides and Halimeda sp. When first sighted, animal 1 looked like a piece of alga. Animals 3 and 4 were captured at 12 m (CFER-42) on a mixed sand and silt bottom dominated by a dense stand of Halimeda sp., scattered Caulerpa cupressoides, many large black holothurians and a few brown-yellow holothurians (probably species of Holothuria, Pentacta or Stichopus). Both animals were observed in close association with Halimeda plants. Animal 3 was clinging with the tips of its outstretched arms to the upright branches of the alga about 3 cm above the bottom. The skin showed cryptic coloration of mottled ol- ive-green/brown and was finely papillate; this patterning very closely matched the color and texture of the Halimeda. No spots or blue rings were evident. When capture was at- tempted with a small net, the animal escaped and flashed a vivid pattern of dark patches (maculae) surrounding bright, iridescent blue rings. The animal immediately dropped to the bottom and resumed a pale cryptic pattern to match the grey/beige color of the substrate. All rings and patches disappeared. When finally captured, it again flashed its dark chromatophore patches and iridescent blue rings. This behavior also was elicited in the aquarium and is described in detail below. When animal 4 was first approached by the diver, it was hovering in a motionless position immediately above a Halimeda plant. The animal was oriented with the body at a 45? angle to the bottom and the arms drooping downward. A cryptic color pattern and papil- late texture covered the entire body. When the diver was sighted the animal instantly dropped to the bottom, as if in a "free fall," and assumed the characteristic deimatic pat- 166 ROPER & HOCHBERG tern with arms spread out, web expanded and a pale color. Upon capture the animal flashed its vivid blue rings and surrounding dark maculae, then flushed dark reddish brown. The Halimeda plant to which animal 3 was clinging was inhabited by a rich fauna of ep- iphytes that included small gastropods, scal- lops and other bivalves, a small stomatopod and several hermit crabs and swimming crabs. This would seem a rich source of food for a small octopus like Hapalochlaena. In the lab- oratory the animals were maintained on small Ocypode crabs. Hapalochlaena cf. maculosa also has been observed clinging to Caulerpa cupressoides, another green alga that pro- vides habitat for a diversity of epiphytic organ- isms, including the gastropods Engina and Anachis visible in Fig. 14. C. Components of body patterns The Lizard Island Hapalochlaena have a distinct and fixed pattern of small rings 1-2 mm in diameter (Figs. 7, 8) that differ from the rings and lines of the other species. In H. lunulata they are fewer in number and larger, 7-8 mm in diameter (Fig. 13), whereas in H. cf. fasciata the mantle has a pattern of blue lines (streaks and diagonals) instead of rings (Figs. 11, 12). The configuration, dimensions and distribution of these characters often are obscured or distorted in preserved speci- mens, but they are very distinct and promi- nent in living animals when they are dis- turbed. The ring/macula complex is similar in ap- pearance and function to the ocelli of 2- spotted octopuses such as O. bimaculoides (see Packard & Hochberg, 1977). The center and periphery of each ring are raised and densely invested with dark chromatophores. In the region of the ring itself, the skin over- lying the blue iridophores is transparent or translucent and indented. When Hapalo- chlaena cf. maculosa is resting, a cryptic mottle pattern is assumed. In this pattern the blue rings essentially are invisible and the dark maculae are subdued to invisible (Fig. 18). As the chromatophores are expanded and the maculae darken, the vividness of the blue rings intensifies in a fashion similar to the expression of the blue rings in ocelli (Figs. 16, 17, 26-28). When the chromatophores are partially expanded the centers of the rings are lighter than the maculae surrounding the rings, but when the chromatophores are max- imally expanded over the entire body, the centers are very dark and the blue rings may be obscured (Fig. 15). A single, small, conical papilla often is erected in the center of each ring. During the day when the animals are inac- tive and resting on a sandy bottom in the aquarium the eyes are erect. The arms either are held straight out with the tips curled aborally, or they are curled and tucked up under the body. Often the animal sits partially buried in the sandy substrate. Small conical papillae are erected over the entire body. Four large, primary, light-colored, conical pa- pillae are present on the dorsal mantle and two large compound papillae are located on the head above the eyes. A prominent light- colored, papillate ridge is located ventral to each eye. A large, flat, papillate ridge is located in the midline on the posterior dorsal mantle. The posterior tip of the mantle often is drawn out into a point with an elongate, flat, papillate ridge (Figs. 8, 9). Resting, inactive animals all exhibit cryptic color patterns that match the background. The body and arms are either uniform light beige or a beige mottled with light brown that corresponds to the mantle and arm maculae. The iridescent blue rings are only faintly visi- ble on the mantle, but they are slightly more conspicuous on the arms. A broad light beige stripe, the dorsal mantle light stripe, extends along the dorsal midline from the head to the posterior tip of the mantle (Fig. 15). In the mid-region of the dorsal mantle there are two conspicuous, irregularly shaped, white spots at the center of which is a single, large, mantle papilla (Figs. 7, 8, 16). A third white spot is present on the posterior mantle midline. A posterior head white spot is present on the head just posterior to the eyes. In this species it is in the shape of a curved bar or crescent. A frontal white spot is located on the head just anterior to the eyes, and a single conical papilla is present in the center of this spot. All five white spot regions appear to be underlain by dense patches of leucophores (see Figs. 7, 8, 16). At night Hapalochlaena cf. maculosa rests with the arms curled under the body. The skin texture is granular to minutely papillate and the color a darker or more intense mottle. The frontal white spot is especially conspicuous at night in the beam of a flashlight. Animals in aquaria were observed in a few instances to bury themselves partially in AUSTRALIAN CEPHALOPOD BODY PATTERNS 167 the sand. Burying behavior is initiated as the ventral arms push sand aside laterally. As the excavation continues, arms III help to enlarge and deepen the depression. Jets of water from the funnel are not used. Only the ventral half of the mantle, head and arms III and IV are buried. The skin is uniformly papillate and the color is a light cryptic mottle that matches the substrate perfectly. This behavior was not observed in the field; however, field observa- tions were quite limited. Occasionally animals jet about the aquar- ium without being stimulated by the observer. The body is smooth except for the large, primary, mantle papillae, which are erect. The color is a uniform pale beige, and maculae and rings are not visible. In contrast, animals that are disturbed jet away in an escape reaction; the body is entirely smooth with papillae retracted. When animals are dis- turbed the color flushes from an intense yel- lowish beige mottled with dusky brown maculae to a very dark mottle. Next, the dark maculae enlarge and coalesce, coloring the octopus a uniform dark reddish brown. Often ripples of color, in the form of "passing clouds" move over the body surface. A dark stripe extends along each side of the mantle and head and through the eye, then tapers along the lateral edge of the dorsal arms nearly to the tips (Figs. 9,10). The other arms are an intense yellow. The blue rings and associated dark maculae pulsate in syn- chrony, which gives the appearance of flash- ing rings. Ring/macula flashes initially are directed unilaterally toward a disturbance (Fig. 26). If the disturbance continues, the ring/macula flashing pattern becomes bilat- eral as it darkens and intensifies, then spreads wave-like over the entire dorsal and lateral surfaces of the mantle, head, web and arms (Fig. 27). When a disturbance persists or is very intense, young Hapalochlaena cf. maculosa discharge a cloud of thin, reddish brown ink. The ink rapidly dissipates and does not con- geal or "hang" in the water as a pseudo- morph, as is typical in other Octopus spe- cies. Up to 10 clouds of ink were discharged during a period of repeated disturbances. The ink sac degenerates with growth, so that full-grown adults lack the ability to produce ink. Table 4 presents the currently recognized components of body patterns of H. cf. maculosa based on observations and photo- graphs at Lizard Island. D. Discussion A discussion of Hapalochlaena species ap- pears at the end of the section on H. lunulata. 4. Hapalochlaena cf. fasciata (Hoyle, 1886) Common name: Blue-lined octopus. We introduce this name to draw attention to the distinct blue lines on the mantle of this spe- cies. Because of taxonomic confusion, this species frequently has been called the "blue- ringed octopus," but we recommend that use of this common name be restricted henceforth to the true "ringed" species. A. Synopsis Diagnosis: Mantle small, elongate, ovoid, pointed posteriorly, mantle length to 40 mm; total length to 110 mm; total weight 40-50 g; skin papillate, with conspicuous pattern of dark elongate maculae and iridescent blue lines on mantle and dark maculae with small iridescent blue rings on head, web and arms; eyes small; arms short, 1.5 to 2.5 times mantle length, subequal; typical arm formula IV.III.II.I; suckers small, numerous, none en- larged; gills with 4 or 5 lamellae per demi- branch; right arm III hectocotylized, length 95% of left arm III; end organ medium to large, 7-9-12% of hectocotylized arm length; ligula flat and smooth, calamus large; eggs 7-9 mm long. Distribution: Australian endemic. Temper- ate to subtropical, apparently restricted to New South Wales. Widespread in shallow, sheltered coastal waters from the intertidal zone to depths of 10 to 30 m. Typically found along rocky shores where it lives in crevices, rock pools and underwater caves. Also found in bays, living under rocks, in grass beds and in debris such as cans, bottles and empty bivalve shells (scallops, oysters and mus- sels). Life history: Life span 7 to 9 months. Sexually mature at 4 months. In the Sydney area, spawning occurs in March and from September to December. 100 to 200 large eggs (7 to 9 mm) laid by female and brooded loosely in web and arms. Embryonic develop- ment takes about 60 days. Development di- rect; young benthic upon hatching. Food pri- marily crabs, but bivalves also may be eaten. References: Anon., N.D.; McMichael, 1957,1958,1964,1971; Dew, 1959; Hopkins, 1964; Lane & Sutherland, 1967; Sutherland & 168 ROPER & HOCHBERG *L 'a I FIGS. 7-10. Hapalochlaena cf. maculosa. Figs. 7, 8: Stylized dorsal (7) and right lateral (8) views to show size and distribution pattern of blue rings (clear circles) surrounded by dark maculae (black stipple), primary papillae (concentric, spiral and peaked lines) and white spots (encompassed by dashed lines). Fig. 9. Right lateral view, jetting octopus with dark lateral stripe along arms, head and mantle. Fig. 10. Right lateral view of head and eye (anterior to right) with dark lateral stripe. AUSTRALIAN CEPHALOPOD BODY PATTERNS 169 11 fws phws dmws pmws 13 FIGS. 11-13. Hapalochlaena cf. fasciata. Figs. 11,12 and H. lunulata, Fig. 13: Stylized dorsal (11, 13) and right lateral (12) views to show size and distribution pattern of blue lines and rings (clear lines and circles) surrounded by dark maculae (black stipple) and white spots (encompassed by dashed lines). 170 ROPER & HOCHBERG FIG. 14. Hapalochlaena cf. maculosa. Attached to the green alga Caulerpa cupressoides, in Watson's Bay, Lizard Island. Left lateral view; note promi- nently erected papillae on mantle and ventral to the eye. Epiphytic gastropods are Engina and Anachis. Photograph by N. Coleman. Lane, 1969; Deas, 1970; Freeman & Turner, 1970; Croft & Howden, 1972; Cropp, 1972; Friese, 1972; Tranter & Augustine, 1973; Reynolds, 1983; Keith, 1986; Marsh & Slack- Smith, 1986. (Note: Unless illustrations or photographs are provided, identifications in the literature cannot be relied upon with re- gard to H. cf. maculosa and H. cf. fasciata, hence both are combined here.) B. Field observations None. C. Components of body patterns A single individual of H. cf. fasciata (Hoyle, 1886) was observed for 14 days in an aquar- ium at the Australian Museum in Sydney. This species is distinct from those observed at Lizard Island (H. cf. maculosa) and the spe- cies traditionally identified as H. lunulata. We consider the species commonly collected in New South Wales to be H. cf. fasciata. For comparison, we present information on the color and body patterns of the New South Wales specimen (Figs. 19-25). Body propor- tions differ in H. cf. fasciata from those in H. cf. maculosa. The arms are longer and more attenuate and the mantle is slightly larger and more robust, although still distinctly elongate and tapered posteriorly (Figs. 19-21). The normal background coloration is a deep orange/yellow. The dorsal and lateral sur- faces of the mantle are covered with a regular pattern of iridescent blue lines, not rings (Figs. 11, 12, 19-25). The lines are oriented on the body as streaks or diagonals. The ventral surface of the mantle lacks lines or rings. Small blue rings are present on the head, web and arms. Those on the arms coalesce proximally into large irregularly shaped rings or transverse figure-8's and distally into dots of blue (Fig. 11). The body can be covered with small conical and large flat papillae or papillate ridges. The morphogenetic fields or patterns of distribu- tion of papillae appear to be similar to the Lizard Island species, H. cf. maculosa. Rows of small papillae occur in the light areas between the dark lines on the lateral mantle but not within the maculae. In H. cf. fasciata three to four low, connected papillae lie ven- tral to the eye rather than a strong, raised papillate ridge, as in H. cf. maculosa. Behavioral responses manifest in color and body patterns are very similar to the species at Lizard Island. At rest, the cryptic pattern is a uniform light grey/beige (Fig. 22). The iri- descent blue mantle lines and the head, web, and arm blue rings are visible but subdued, the maculae are absent, and the texture is finely granular with minute papillae. With in- creasing disturbance (Figs. 20, 23-25) the animal flushes to a uniform dark charcoal or slate grey and the iridescent blue lines and rings become very intense surrounded by maculae of very dark chromatophores. In this pattern the primary papillae may be erect and prominent. Table 5 presents an inventory of compo- nents of body patterns currently recognized for H. cf. fasciata based upon observations and photographs of a living animal in Sydney and on the literature. D. Discussion A discussion of Hapalochlaena species ap- pears at the end of the section on H. lunulata. AUSTRALIAN CEPHALOPOD BODY PATTERNS TABLE 4. Components of body patterns in Hapalochlaena cf. maculosa. 171 I. Chromatic components A. Light light-colored papillae dorsal mantle stripe dorsal mid-mantle white spots (dmws, 2) posterodorsal mantle white spot (pmws) posterior head white spot (phws) frontal white spot (fws) transverse white streak (tws) II. Textural components A. Primary papillae simple dorsal mantle papillae (dmp, 4) compound dorsal eye papillae (adep, pdep, 2/eye) compound ventral eye papillate ridge (vepr, 1/eye) compound flat posterior dorsal mantle papillate ridge (pdmpr, 1) compound flat posterior mantle tip papillate ridge (mtpr, 1) simple dorsal mantle white spot papillae (mwsp, 1/spot) simple frontal white spot papilla (fwsp, 1) extended posterior mantle tip III. Postural components radiating arms curled arms coned arms drooping arms flared web elongate pointed mantle raised head and eyes IV. Locomotor components and maneuvers resting/sitting hovering scuttling jetting inking free tall burying V. Body patterns A. Chronic light cryptic mottle light uniform B. Dark dark maculae (spots on mantle, web and C. Other iridescent blue arms) dark lateral stripe rings B. Secondary papillae C. Other minute simple granular uniform body smooth papillae simple ring papillae (1 in center of each ring) simple intermacular lateral mantle papillae B. Acute dark conflict mottle dark uniform (flush) deimatic (white blanch) flashing maculae and blue rings flamboyant 5. Hapalochlaena lunulata (Quoy & Gaimard, 1832) Common name: Greater blue-ringed octo- pus. We introduce this name to draw attention to the large blue rings (7-8 mm in diameter) on the dorsal mantle, head, web and arms. A. Synopsis Diagnosis: Mantle ovoid, pointed posteri- orly, slightly flattened dorsoventrally; mantle length to 55 mm; total length to over 100 mm; total weight to 80 g; skin soft and semi- gelatinous, mantle, head, web and arms with 172 ROPER &HOCHBERG TABLE 5 The components of body patterns in Hapalochlaena cf. fasciata. I. Chromatic components A. Light dorsal mantle stripe dorsal mid-mantle white spots (dmws, 2) posterodorsal mantle white spots (pmws) posterior head white spot (phws) frontal white spot (fws) II Textural components A. Primary papillae simple dorsal mantle papillae (4) compound dorsal eye papillae (2/eye) compound ventral eye papillate ridge (1/eye) compound flat posterior dorsal mantle papillate ridge compound flat posterior mantle tip papillate ridge simple dorsal mantle white spot papillae (1/spot) simple frontal mantle white spot papilla extended posterior mantle tip III. Postural components radiating arms curled arms coned arms erect head & eyes elongate pointed mantle copulatory embrace1 IV. Locomotor components and maneuvers resting/sitting scuttling jetting inking mating1 V. Body patterns A. Chronic light cryptic mottle light uniform B. Dark dark maculae (spots on web and arms) dark lines (on mantle) dark orange/yellow B. Secondary papillae minute simple uniform body papillae simple intermacular lateral mantle papillae C. Other irridescent blue lines and rings C. Other granular smooth Acute dark conflict mottle dark uniform (flush) flashing maculae, blue lines and rings flamboyant 'Tranter & Augustine (1973). FIGS. 15-22. Hapalochlaena cf, maculosa. Figs. 15-18, dorsal views of resting animals and H. cf. fasciata, FIGS. 19-22, dorsal views. Fig. 15. Dark mottle; blue rings completely subdued; maculae dark, expanded and coalesced, mantle and head white spots and dorsal mantle white stripe expressed but subdued. Fig 16 Light mottle; blue rings and maculae expressed, especially on right side (top) facing disturbance, all mantle and head white spots expressed. Fig. 17. Light uniform; blue rings expressed; maculae subdued, separate. Fig. 18. Dark uniform; blue rings greatly subdued; maculae expanded and coalesced. Fig. 19. Scuttling locomotion while disturbed; blue lines and rings and maculae dark, expressed moderately; mantle, head and frontal white spots evident. Fig. 20. Warning pattern; blue lines and rings expressed vividly; maculae dark, white spots subdued to absent. Fig. 21. Resting after disturbance; blue lines and rings expressed moderately; maculae subdued; white spots subdued. Fig. 22. Resting position; light uniform; blue lines and rings very subdued; maculae absent; no white spots expressed except subdued frontal spot; mantle tip rounded. 174 ROPER & HOCHBERG conspicuous pattern of large iridescent blue rings (7-8 mm in diameter) with broad dark maculae around the outer periphery and clear centers; eyes small; arms short, 1.5 to 2 times mantle length, subequal; typical arm formula IV.III.II.I; suckers few, level with oral surface of arms; gills with 7 to 9 lamellae per demibranch; right arm III hectocotylized, length 80 to 90% of left arm III; end organ medium-sized, 7 to 9% of hectocotylized arm length; ligula flat, smooth, with slightly ele- vated edges; calamus small, open; eggs 2.5 to 3.5 mm long; hatchlings planktonic. Distribution: Widely distributed throughout the Indo-West-Pacific and Indian Oceans. Australia, New Guinea, Philippines, Sri Lanka, Vanuatu Is., Solomon Is., Misal Is., Andaman Is. In Australia the species has been recorded in Queensland, Northern Ter- ritory, and Western Australia. Little informa- tion is available on vertical distribution and habitat. Life history: Life span not known. In the Philippines, spawning occurs in March and April. 60 to 100 small eggs laid by female and attached to substrate in festoons of about 20 eggs each. Embryonic development takes about 25 to 35 days Hatchlings are briefly planktonic prior to becoming benthic. Species feeds actively on crabs and bivalves. References: Adam, 1954; Flecker & Cot- ton, 1955; McMichael, 1957, 1971; Overath, 1973; Overath & Boletzky, 1974; Marsh & Slack-Smith, 1986; Wells & Bryce, 1986. B. Field observations None. C. Components of body patterns Not observed in life. This very distinct spe- cies has been studied by us only from color transparencies of live animals provided by C. Bryce and A. Kertstich (Figs. 29, 30). The drawing (Fig. 13) was developed from a pho- tograph by Kertstich to contrast the general body pattern of H. lunulata with the patterns of the two species we observed alive. Photo- graphs of live animals appear in Wells & Bryce (1986) and in Marsh & Slack-Smith (1986). Discussion of Hapalochlaena spp. The genus Hapalochlaena is a complex of octopuses with small bodies, short arms and shallow webs. The genus is further character- ized by the unique presence of iridescent blue rings or lines set in macula of dark chrom- atophores. Observations were made on live animals of two very distinct species of Hap- alochlaena, one from Lizard Island, the other from near Sydney. When observed alive or from color photographs, these two species are very easy to distinguish, whereas pre- served specimens may be less distinctive. In fact, the study of living animals and of color photographs of living animals has enabled us to recognize species differences and thus to FIGS. 23-31. Hapalochlaena cf. fasciata, Figs. 23-25, dorsal views; H. cf. maculosa, Figs. 26-28, dorsal and left lateral views; H. lunulata, Figs. 29-30, dorsal and left lateral views, and Octopus cyanea/O. ornatus, Fig. 31, lateral views. Fig. 23. [top row, left] Light mottled (grey phase); blue lines and rings expressed; maculae expressed lightly; mantle, head and frontal white spots expressed. Fig. 24. [top row, center] Dark mottle; blue lines and rings expressed vividly; maculae dark, expanded and coalesced; frontal white spot visible, other white spots suppressed. Fig. 25. [top row, right] Light mottle (beige/yellow phase); blue lines and bars expressed; maculae dark and separate; white spots suppressed. Fig. 26. [middle row, left] Moderate warning pattern of Lizard Island animal; blue rings expressed; maculae expanded dark and coalesced on right side facing disturbance (left of image), otherwise subdued; mantle white spots visible. Fig, 27. [middle row, center] Deimatic pattern; blue rings and black maculae expressed vividly; texture finely papillate. Photograph by C. Bryce (Shark Bay, Western Australia). Fig. 28. [middle row, right] Scuttling locomotion; warning pattern; blue rings expressed vividly; black maculae expanded very dark and coalesced. Photograph by C. Bryce (Albany, Western Australia). Fig. 29. [third row, left] Deimatic pattern, blue rings vivid; maculae expressed minimally as black rings outside of blue rings, centers light. Photograph by C. Bryce (Exmouth Gulf, Western Australia). Fig. 30. [bottom row, left] Deimatic pattern; blue rings vivid; dark macular rings expanded around outside of blue rings, centers dark. Photograph by A. Kerstitch (Vanuatu Islands). Fig. 31. [bottom row, right] Octopus cyanea (right) in light uniform pattern (deimatic) with vivid ocellus and smooth texture. Octopus ornatus (left) with white streaks on mantle and white spots on arms against maroon background. Photograph by W. F. van Heukelem (Honolulu, Hawaii). AUSTRALIAN CEPHALOPOD BODY PATTERNS 177 begin resolving problems that have puzzled investigators since the original descriptions of the nominal species over 100 years ago. While a great deal of work remains to be done, we have been able to distinguish at least three species based upon the size and configuration of their iridescent blue mark- ings. Many living and preserved specimens from localities over the entire geographic range of the genus must be examined to make final judgments and decisions. At present, we concur with the identifications provided by 5. S. Berry (in Halstead, 1965) and recognize the following distinct species: 1. Hapalochlaena cf. maculosa (Figs. 7-10) is characterized principally by small blue rings, 1-2 mm in diameter, on the dorsal and lateral (but not ventral) surfaces of the mantle, head, web and arms. This species occurs at Lizard Island and similar small-ringed forms have been recorded from numerous localities in Australian and Indonesian waters. Small- ringed forms may represent a widely distrib- uted species, but more likely a species com- plex is involved, since in at least one area the ventral mantle of specimens examined is par- tially covered with a number of blue rings. This is the maculosa type of blue-ringed oc- topus originally described by Hoyle in 1883. 2. Hapalochlaena cf. fasclata (Figs. 11, 12) is characterized by short, blue lines (streaks and diagonals) on the mantle and by very small blue rings (single circles, figure-8's or complex circular designs) on the head, web and arms. The configuration and distribution of blue lines and rings is comparable in gen- eral terms to that of the lesser blue-ringed octopus. This is the species we studied from waters near Sydney. It has been referred to repeatedly as H. maculosa in the literature. However, all the specimens we have exam- ined from New South Wales have blue lines not rings and, hence, the specific name fasciata Hoyle, 1886 seems applicable. The extent of its distribution is unknown because it has been frequently confused with the "maculosa" species. 3. Hapalochlaena lunulata (Fig. 13) is dis- tinguished by relatively large blue rings up to 7 or 8 mm in diameter on the dorsal and lateral surfaces of the mantle, head, web and arms. This species also seems to be widely distributed in Australian and central Indo- Pacific waters. A complex of species or subspecies may be involved that can be resolved only by further critical study. What could be the function of the brilliant, iridescent blue displays of Hapalochlaena? Judging from the behavioral responses to disturbance or threat, these seem to be a form of warning coloration with which these small octopuses signal their unpleasant taste or poisonous bite. Interestingly, there are several other vividly marked small octopuses (e.g. Octopus zonatus, O. chlerchlae), but it is not known whether they also are poisonous. Surprisingly, reference to color and body patterns in the Hapalochlaena complex is very limited. The majority of reports detail the blue rings and dark spots or maculae on the body and arms. Although several photo- graphs show animals with distinct lines on the mantle, the animals still are referred to as the "blue-ringed octopus." On the basis of photo- graphs in published reports we can identify the following: H. cf. maculosa?Reynolds, 1983; Marsh & Slack-Smith, 1986; H. cf. fasciata? Sutherland & Lane, 1969; Deas, 1970; Hal- FIGS. 32-42. Sepia papuensis, Figs. 32-35, right lateral and dorsal views; Metasepia pfefferi, Figs. 36-41, right lateral and dorsal views, and Metasepia tullbergi, Fig. 42, right lateral view. Fig. 32. [top row, left] Bipod position; dark mottle on dorsal mantle, light mottle on ventral mantle; dorsal mantle dark spot prominent; lateral mantle white spots expressed against light background. Fig. 33. [second row, left] Bipod position, dark mottle on dorsal and ventral mantle; head, mantle and (inline white spots conspicuous. Fig. 34. [top row, center] Light mottle approaching light uniform; unilateral dorsal mantle dark spot directed toward source of disturbance; dorsal mantle white spots expressed. Fig. 35. [top row, right] Dark mottle. Fig. 36. [third row, left] Tripod posture; raised arms I; diagonal mantle white bars, (inline white stripe, lateral mantle white spots expressed; head and arm light components expressed; yellow and magenta absent. Fig. 37. [fourth row, left] Ambling locomotion on ambulatory flaps and arms IV; right arm IV down, left arm IV raised to take next step; flamboyant body pattern; yellow and magenta expressed. Fig. 38. [middle row, center] Dark uniform with diagonal mantle white bars expressed; yellow and magenta absent. Fig. 39. [middle row, right] Passing wave pattern, especially on left side of mantle; first anterior wave in mid-cycle, second anterior wave originating at anterior mantle margin; posterior wave originating at posterolateral margins of mantle. Fig. 40. [bottom row, center] Prone position with flanged fins and flattened arms; yellow and magenta expressed. Fig. 41. [bottom row, right] Head-on view of flamboyant arm pattern; suckers enveloped by yellow protective membranes. Fig. 42. [bottom row, left] Flamboyant pattern of color, texture and posture; note ambulatory flap. Photograph of Japanese animal by T. Koyama, courtesy of T. Okutani. 178 ROPER &HOCHBERG TABLE 6. The components of body patterns in Metasepia pfefferi. I Chromatic components A. Light arm IV white spots (aws IV) arm III white spots (aws III) arm I white spots (aws I) web white spots (wws) dorsal head white spots (dhws) dorsal head white triangles (dhwt) lateral head light field (Ihlf) dorsal mantle white bars (dmwb) dorsal mantle light field (dmlf) (inline white stripe (flws) lateral mantle white spots (Imws, ventral to fin) ventral mantle white spots II. Textural components A. Primary papillae and flaps anteroventral eye papilla (avep, 1/eye) anterodorsal eye papilla (adep, 1/eye) posterodorsal eye papilla (pdep, 1/eye) (inline papillae (flp 1, 5-7/side) laterodorsal mantle papillae (Idmp, 2 pairs) anterodorsal mantle papilla (admp, 3) posterodorsal mantle papillae (pdmp, 1 pair) posterodorsal mantle flaps (pdmf, 1 pair) mid-dorsal mantle flaps (mdmf, 1 pair) mantle white bar flaps (mwbf, 1 pair) III. Postural components arms I raised arms IV lowered (tips on substrate) flamboyant arm splay drooping arms (while hovering) raised head flanged fin (folded down) flattened/flared arms bent arms IV (while ambling) bipod (body off bottom) tripod (posterior mantle on bottom) prone (body flat on bottom) IV. Locomotor components and maneuvers resting floating hovering swimming jetting ambling inking V. Body patterns A. Chronic light cryptic mottle light uniform B. Dark dorsal head dark field (dhdf) lateral mantle dark fields posterior mantle dark spot (pmds) dorsal mantle dark spots (dmds) (eye spots of deimatic pattern) arm IV dark stripe (ads) B. Secondary papillae dorsal head papillae dorsal mantle papillae finline papillae (flp2, 5/side) lateral mantle white spot papillae B. Acute dark conflict mottle dark uniform (flush) flamboyant passing wave (apw, ppw, 1,2) deimatic AUSTRALIAN CEPHALOPOD BODY PATTERNS 179 stead & Danielson, 1970; Voss, 1971; Cropp 1972; Tranter & Augustine, 1973; Keith, 1986 H. lunuIata?Robson, 1929 (pi. 4, fig. 1) Flecker & Cotton, 1955; Anon., 1968; Marsh & Slack-Smith, 1986; Wells & Bryce, 1986. Brief descriptions of color in adults and hatchlings of H. cf. fasciata were provided by Dew (1959) and Tranter & Augustine (1973), who found that the characteristic blue lines and rings of H. cf. fasciata appeared 6 weeks after hatching, although the macula pattern developed at an age of 3 or 4 weeks. The postures and activities associated with mating were described by Tranter & Augustine (1973) but patterns of color specific to court- ship and mating were not mentioned. Species of the subfamily Octopodinae are all very similar morphologically and anatomi- cally. Hence, few supraspecific taxa have been erected to help subdivide an enormous number of seemingly uniform species. The fixed and characteristic markings observed in Hapalochlaena lead us to conclude that this species complex should be recognized as distinct at the generic level. Morphogenetic patterns, especially of color and texture, are fixed, conservative charac- ters within families and genera. Although more research is needed, this phenomenon will add a whole new suite of genetically stable characters upon which to base system- atic analyses, diagnoses of species and gen- era, etc. The elucidation of these morpho- genetic patterns, through study of living animals or color photographs of living ani- mals, will provide valuable information for understanding both the systematics and be- havior of cephalopods, and for identification of species in the field. 6. Metasepia pfefferi Hoyle, 1885 Common name: Pfeffer's flamboyant cuttle- fish. We name this species after the German teuthologist Georg Pfeffer. "Flamboyant" em- phasizes the striking, flowery body patterns of the genus Metasepia. A. Synopsis Diagnosis: Mantle very broad, oval; mantle length to about 80 mm; total length to about 160 mm; dorsal mantle with 3 pairs large, pri- mary flat, flap-like papillae and 1 pair promi- nent mantle white bars; head with primary pa- pillae over eyes; arms broad, blade-like; fins broad, transparent; tentacular club short, with swimming keel twice as long as club; dorsal and ventral protective membranes separate on tentacular stalk; club suckers very few in number, in about 5 transverse rows, with 3 or 4 median suckers enormously enlarged; cuttlebone broad, rhomboidal, shorter than mantle; dorsal surface completely chitinized, without median rib or spine. Distribution: Tropical Australian waters: Arafura Sea (type locality), Queensland (Cap- ricorn group, Moreton Bay?Alan Jones, per- sonal communication) to Western Australia (to about 33?S). Shallow-water species on sand/silt bottom to about 50 m. Life history: Unknown. This paper is the first report of observations on living animals. References: Hoyle, 1885b, 1886; Adam & Rees, 1966; Adam, 1979; Lu & Phillips, 1985; Wells & Bryce, 1986. B. Field observations Two juvenile M. pfefferi were captured in 10 to 13 m at separate locations during daytime dives in Watson's Bay (CFER-22 and CFER- 42). In both localities, the flat bottom was composed of a mixture of sand and silt, devoid of rocks and rubble. The bottom was inhabited by a large number of black holothur- ians, Holothuria sp., a few brown-yellow holothurians, and dense patches of attached green algae: a calcareous species of Halimeda and scattered Caulerpa cupres- soides. In its natural habitat, M. pfefferi looks like anything but a cuttlefish. When first ob- served and prior to capture, animal 1 was variously identified by divers as a small frilly crab, a crayfish, a pufferfish or a piece of alga slowly moving or drifting along the bottom. Even when the divers knew what to look for during subsequent dives, recognition of ani- mal 2 was difficult. When first observed it was hovering in a stationary position 10 cm above the bottom, motionless except for gentle man- tle undulations. Upon sighting the diver, it swam to the bottom and assumed a cryptic pattern that matched the beige/grey color of the sandy silt substrate. When pursued and captured, the animal changed to a vivid black and yellow color pattern and a distinctly papil- late texture. Metasepia pfefferi's ability to conceal itself is due both to the vivid and exaggerated color and textural patterns as well as the unusual configuration of the arms (described below). Observations of behavior and body patterns in aquaria were consistent with those observed in the field. 180 ROPER & HOCHBERG dmds dmwb ppw3 ppw2 ppwl af lmws FIGS. 43?45. Metasepia pfefferi. Field drawings to show texture, flamboyant configuration of arms and color patterns on mantle during passing wave pattern. Fig. 43. Head-on view; arm configuration observed while in bipod, tripod and prone postures. Fig. 44. Dorsal view. Fig. 45. Right lateral view; prone posture; note ambulatory flaps (shown shorter than normal). Drawings by B. Morton of Lizard Island animal in aquarium. AUSTRALIAN CEPHALOPOD BODY PATTERNS 181 FIG. 46. Metasepia pfefferi: Stylized dorsal view showing location of large, primary papillae and flaps and smaller secondary papillae. C. Locomotion In addition to floating, hovering, swimming and jetting, M. pfefferi moves along the bot- tom by means of a remarkable new type of locomotion we term "ambling." This mode of locomotion is aided by a pair of muscular flaps along the margin of the posterior third of 182 ROPER & HOCHBERG c f-J; g \ ft :'.V' ? E ; to / r- ?* x> ? g i? ? * a *J o ? "" Z *? E (5 .? in a c I c I t Q. o o o o o ?D 0) N C/3 a. ra I * * 1 * a. a. a ? m "" TJ a. ? a. ** c C Q. O rj ?o c ?o o J5 5 o ^= o 5 0) ?> o ?a ?o 1 .CO a> ?S d ! W 184 ROPER &HOCHBERG AUSTRALIAN CEPHALOPOD BODY PATTERNS 185 the ventro-lateral mantle (Figs. 37, 45, 58, 59). The flaps, which look like a pair of elongate, ventrally-directed fins, border the edge of the ventral "suction" disk. They are grey to beige in overall color with two indis- tinct pale yellow vertical bands. We term these structures "ambulatory flaps," to de- note their function in this unusual mode of locomotion. The ambulatory flaps are erect most of the time, whether the animals are resting, swim- ming, hovering or ambling. At rest the animals sit tripod-fashion (Fig. 36) on the erected ambulatory flaps and on the ventrally directed ventral arms. The tips of the ventral arms typically are flattened and curled medially giving the appearance that the animals are resting on their "forearms" (Fig. 41). In this position the head and mantle opening are elevated well above the bottom. The animals amble in a "slow walking" motion along the bottom using the ventral arms as "legs." The ambulatory flaps partici- pate by sliding or shuffling forward alternately in sequence with the ventral arms (Figs. 37, 58, 59). This locomotion pattern is similar to the gait of a quadruped vertebrate in that the right ventral arm in front and the left ambula- tory flap in the rear move forward, followed by the left front arm, right rear flap. The move- ment is slow and deliberate, resembling an amble or a shuttle. By reversing the sequence the animals are able to move backwards. When approached from the head end, the animals amble backward rather than jetting away as is typical of most other cuttlefishes. Finally, M. pfefferi can amble sideways when threatened from the side. Side stepping also is quite slow and deliberate. In moving to the right, for example, the body is supported on the right ventral arm and the right ambu- latory flap. The left ventral arm and flap are swung toward the midline and set down in place. The weight then is shifted to the left arm and flap and the "appendages" on the right side are swung out to the right and placed on the substrate. Many details of the fascinating new "am- bling" behavior of M. pfefferi still remain to be worked out, but these original observations leave no doubt that this type of locomotion is normal for this species. In fact, it seems to be preferred, as the animals were seldom ob- served to swim. D. Components of body patterns Figs. 43 to 50 and Table 6 indicate the location and terminology of the various color, textural and postural components discussed below. When resting or sitting on the bottom, the animals assume one of three basic postures. (1) As an animal moves into the resting position it hovers just above the bottom (Fig. 60), lowers the ventral arms., to a vertical position, then settles into a "bipod" position with the weight supported on the tips of the two lowered ventral arms; it may remain in the bipod position for some time. (2) In the "tri- pod" position, the body is supported by the ventral arms and the posterior mantle, which rests on the ambulatory flaps in contact with the bottom (Figs. 36, 37, 57). The head and anterior mantle are elevated, arms II and III are splayed apart and arms I are raised dorsally and held together with the tips curled ventrally (Figs. 36, 37, 57). Arms IV may be either straight or curled at the tips. When the arms are all splayed out, and variously flat- tened and curled, the resemblance to a plant is striking (Fig. 41). (3) In the "prone" resting position, the entire ventral surface of the mantle is in contact with the bottom, and the ventral arms are horizontal with the bottom. In this position the head and anterior mantle are not elevated. In these resting positions, the skin is sculp- tured with a complex pattern of primary and secondary papillae characteristic of the ge- nus. Primary papillae and flaps are large and tonically erect (Figs. 36, 37). They may be colored solid white or yellow or tipped in white or yellow. The primary papillae include the dorsal and anteroventral eye papillae, the five to seven lateral mantle papillae on each side FIGS. 51-60. Metasepia pfefferi, dorsal and right lateral views. Figs. 51-54. Passing waves, various stages; note papillate and chromatic pattern on head and arms. Fig. 55. Dark chromatic phase with subdued dorsal head white triangles and dorsal mantle white stripe; flanged fins. Fig. 56. Flamboyant color, texture and posture; dorsal mantle dark spots prominent. Fig. 57. Tripod position, flamboyant arm splay. Fig. 58. Ambling locomotion, both arms IV on bottom. Fig. 59. Ambling locomotion, left arm IV set, right arm IV raised to take step. Note ambulatory flaps in figs. 57-59. Fig. 60. Moving off bottom with drooping arms IV; ambulatory flaps reduced, edges covered with sand. 186 ROPER &HOCHBERG just dorsal to the insertion of the fin, two pairs of flat dorsal mantle flaps, a field of seven conical papillae on the anterior dorsal mantle and a pair of cup-shaped flaps located in the center of the mantle white bars. A discrete series of smaller conical secondary papillae is located in parallel with the primary papillae especially along the finline (Fig. 39). The overall color of undisturbed animals is a uniform pale grey-white that matches the sandy silt background. The protective mem- branes on the arms are closed over the sucker rows and are striped in pale yellow (Fig. 41). When animals are disturbed slightly the head and dorsal mantle change to a highly contrasting black and white body pat- tern with pale yellow along the finline (e.g. Figs. 39, 47, 51-54). The dorsal head white triangles and the broad dorsal midline white stripe may be masked with black chro- matophores (Figs. 38,55). Yellow or magenta are not expressed except along the protective membranes. "Passing waves" may move over the dorsal mantle (see details below). The fins lack chromatophores and always appear transparent. When animals are dis- turbed repeatedly the lateral stripe above the fins, the finline stripe, turns bright yellow and the yellow of the protective membranes and the oral surfaces of the arms is intensified (Figs. 39, 40, 56). The dorsal mantle is black and partitioned distinctly by the white midline stripe and the intense mantle white bars; mantle dark spots may be expressed (Fig. 56). The lateral man- tle, ventral to the fins, is black with yellow/ orange spots outlined in darker orange (Fig. 36). The ventral mantle is a uniform pale grey with a single white spot posteriorly. The dor- sal head white spots are intense white; the dorsal head white triangles may be yellow or white and the dorsal eye patches are white mottled with black (Figs. 39, 40). The arms often are flattened. The aboral edges of arms l-lll are magenta, the mid-regions are white and the oral edges bright yellow. The tips are pale yellow. Arms IV are dark brown/purple fringed and mottled with white. A distinct patch of yellow occurs on the aboral edge of arms III. This appears to be related to the flamboyant pattern described in juveniles of other genera of cuttlefishes and octopuses. Often while the disturbed animals are in this color and body configuration, a series of "passing waves" washes over the surface of the dorsal mantle (Figs. 39, 49-54). This is seen as a band-like wave of white that moves through the dark field of the mantle dorsal to the fins. The white midline stripe is not af- fected. As one transverse wave originates at the anterior mantle margin and moves poste- riorly, another wave originates on the sides of the posterior mantle and moves anteriorly until the two waves meet and disappear in the region of the mantle white bars. A second set of waves typically is generated before the first set is extinguished (Figs. 39, 50), so it is possible to see for an instant four passing waves, two moving posteriorly and two mov- ing anteriorly. When the fins are used to hover in a stationary position the body is oriented hori- zontally to the bottom and the arms either are splayed out as described above or held to- gether in a ventrally drooping position (Fig. 60). At times the arms are extremely flattened dorso-ventrally and flared out laterally (Figs. 40, 47). In this posture the color pattern may be either a high contrast black and white or a pattern of black, white, yellow and magenta. Metasepia pfefferi typically rests on or hov- ers just above the bottom. When disturbed the animals generally "amble" away and were only rarely observed to swim or jet through the water when prodded repeatedly with a rod or finger. When they do swim or jet the primary papillae and dorsal mantle flaps are erect over the entire body and the color is either a pale uniform beige/white or a vivid black, white, yellow and magenta as de- scribed above. Some variations on these basic patterns are seen in the photographs. It is important to stress that our observations were brief and hence represent only a preliminary inventory of the chromatic, textural and postural com- ponents shown by this species. To develop a more complete body pattern inventory that can be used for comparison with other genera and species of cuttlefishes, additional effort in the laboratory and field is needed. E. Discussion The studies on M. pfefferi at Lizard Island represent the first and only detailed observa- tions of live animals published since its origi- nal description in 1885. A photograph of a live animal is published in Wells & Bryce (1986). Working from the framework provided by Hanlon & Messenger (1988) for young Sepia officinalis, we have been able to recognize over 50 chromatic, textural, postural and locomotor components based upon observa- AUSTRALIAN CEPHALOPOD BODY PATTERNS TABLE 7. The components of body patterns in Sepia papuensis. 187 I. Chromatic components A. Light B. Dark dorsal mantle white spots dorsal mantle dark spots (eye spots dorsal mantle white stripe of deimatic pattern) finline white spots dark uniform (inline white stripe dark mottle lateral mantle white spots (ventral to fin) lateral mantle dark fields dorsal head white patch dorsal head dark field dorsal head white bar lateral head white stripe II. Textural components A. Primary papillae B. Secondary papillae C. Other mantle white spot papillae (1/spot) dorsal head papillae smooth dorsal eye papillae (2/eye) anterodorsal mantle papillae finline papillae (outer row) dorsal mantle flaps (1 pair) III. Postural components bipod (body off bottom) tripod (posterior ventral mantle on bottom) prone (body flat on bottom) arms I raised arms IV lowered splayed arms flanged fin (folded down) IV. Locomotor components and maneuvers resting floating hovering swimming jetting burying inking V. Body patterns A. Chronic light cryptic mottle dark cryptic mottle light uniform dorsal mantle papillae finline papillae (inner row) lateral mantle white spot papillae arm papillae dorsal eye lid papillae (1/eye) B. Acute dark conflict mottle dark dorsal mantle (flush) dark uniform (flush) deimatic (blanch) passing wave tions and photographs on two juvenile speci- mens studied separately for only a few days. Metasepia pfefferi thus revealed an extremely rich repertoire of components. Observations of interacting and sexually mature adults will add still further to the already impressive behavioral repertoire of this species. Traditionally, most authors have stated that the species Sepia pfefferi and its congener from Japanese waters, S. tullbergi Appellof (1886), belong in the subgenus Metasepia erected by Hoyle (1885b) to contain pfefferi (e.g. Adam & Rees, 1966; Adam, 1979; Natsukari, 1979; Lu & Phillips, 1985). How- ever, some authors, such as Iredale (1954), Okutani (1973) and Okutani & Habe (1975) have interpreted Metasepia as a genus. The primary characters that distinguish Metasepia from Sepia are: (1) the short, round, dorsoventrally thickened body; (2) the very unusual morphology of the cuttlebone that is rhomboidal in shape, very broad, covered dorsally with a chitinous layer, and lacks the posterior spine and a dorsal midline rib and 188 ROPER & HOCHBERG t: j= O) O C c = ifi 0) 0) _ 10 > > to (a c-o 2 a. O C O X d) (8 -O 0] is c j.- - .2> c = > ~~ ifS - Q/g " % 3 ,ra 5. o w > I- .s>iS J. L.U- a5 T (o ? ? <= r- a>