Zeiformes

(Dories)

Class Actinopterygii

Order Zeiformes

Number of families 6

Evolution and systematics

Fossils identified as zeiform fishes are confined to marine habitats and are relatively young, ranging in age from Oligocene to Holocene deposits. They have been found in Europe, the West Indies, South Africa, and Indonesia. The phylogenetic relationships of the Zeiformes were explicated by Johnson and Patterson in their 1993 publication on the phylogenetic relationships of the percomorph fishes. They recognized Zeiformes as a monophyletic taxon characterized by the following shared derived characters. There is a distinctive configuration of upright columnar processes on the dorsal elements of the gill arch skeleton. Baudelot's ligament originates immediately under the vagus foramen of the exoccipitals. The distal part of the proximal middle radials of the dorsal fin pterygiophores is expanded laterally. The palatines have a specialized, mobile articulation with the ectopterygoid, which is truncated dorsally. The metapterygoid is extremely reduced, and a continuous median cartilage extends below the frontals and between the ethmoid cartilage and pterosphenoids.

Based on what they term "admittedly tenuous evidence," Johnson and Patterson considered the Zeiformes to be the sister taxon of a group making up the order Beryciformes and a huge conglomeration of spiny-rayed fishes known as the "percomorpha," including the Perciformes, Pleuronectiformes (flounders, soles, etc.), Tetraodontiformes (triggerfishes and pufferfishes, among others), Scorpaeniformes (scorpionfishes, gurnards, flatheads, and so on), Dactylopteriformes (helmet gurnards), Synbranchiformes (swamp eels, spiny eels, and others), Elassomatidae (pygmy sunfish), Gasterosteriformes (pipefish, trumpetfish, etc.), Mugiloidei (mullets), and Atherinomorpha (Atheriniformes, Beloniformes, and Cyprinodontiformes).

The order Zeiformes comprises six families: Zeidae, Parazenidae, Zeniontidae, Oreosomatidae, Grammicolepidae, and Caproidae. In their 1966 seminal paper on the phylogenetic relationships of teleost fishes, P. H. Greenwood and collaborators included the boarfishes (family Caproidae) in the Zeiformes, but they presented no evidence to support this assignment. Heemstra excluded the Caproidae from the Zeiformes in his 1980 taxonomic revision of the zeid fishes of South Africa, and in the book Smiths' Sea Fishes, Heemstra included the Caproidae in the order Perciformes. For the purposes of this publication, the Caproidae are included in the Zeiformes.

Much work remains to be done on the systematics of the Zeiformes, and the classification adopted here is tentative. The composition, definitions, and distinction of the families Zeidae, Zeniontidae, and Parazenidae are unsettled. The genus Cyttomimus (Gilbert, 1905) appears to be related closely to Capromimus (Gill, 1893). Although these two genera seem to be placed correctly in the Zeiformes, their affinity with any of the families recognized here is unclear. The genus Macrurocyttus (Fowler, 1933) may belong in the Zeniontidae, but the head of Macrurocyttus acanthopodus (Fowler, 1933) looks very different from those of the two species of Zenion that are known.

Physical characteristics

The body is deep, compressed, and oblong to disk-shaped. The upper jaw is more or less protrusible, and there are minute, slender, conical teeth in the jaws and vomer. Adults range in size from the dwarf dory (Zenion hololepis) at 4 in (10 cm) to the 3-ft (90 cm), 12-lb (5.3 kg) South African Cape dory (Zeus capensis). Most species are silvery, bronzy, brown, or reddish. The John Dory is silvery or bronzy, with indistinct longitudinal dark stripes from head to tail and a conspicuous white or yellow-edged black ocellus in the middle of the body. The juvenile buckler dory (Zenopsis conchifer) is silvery and covered with scattered, vaguely defined black spots. Dories can change from silvery to dark brown or gray in seconds. Males and females are colored similarly.

Zeiform fishes can be recognized by the following combination of characters: five to ten dorsal fin spines; zero to four anal fin spines; pelvic fins with one spine and five to seven soft rays or no spine and six to 10 soft rays; a caudal fin with 11, 13, or 15 principal (segmented) rays, of which 9, 11, or 13 rays are branched; and unbranched dorsal, anal, and pectoral fin rays. The orbitosphenoid bone is absent, and there is no subocular shelf or supramaxilla. There are seven or eight branchiostegal rays, 3.5 gills (no slit behind the last hemibranch), 25–46 vertebrae, and a gas bladder.

Distribution

The order is represented in the western Atlantic from Canada to Argentina, including the Gulf of Mexico and the Caribbean Sea. In the eastern Atlantic, zeiforms are known from the North Sea to South Africa, including the Mediterranean, Black Sea, Azores, Madeira, Canary Islands, Cape Verde Islands, and Saint Helena. In the Indian Ocean, zeiforms occur along the east coast of Africa to the Gulf of Aden, Oman, the Persian Gulf, Pakistan, India, Sri Lanka, Maldives, and Madagascar and eastward to the Andaman Sea, Burma, Thailand, Cambodia, Indonesia, Vietnam, Australia, New Zealand, New Guinea, the Philippines, China, Taiwan, Japan, and Korea. In the eastern Pacific, zeiforms range from Canada to Chile.

Habitat

Most zeiform fishes are demersal, living near the bottom of the continental shelf or upper continental slope region. They range in depth from 110 to 5,084 ft (35–1,550 m). The adult John dory (Zeus faber), and probably adults of some other zeiform species, frequently occurs in midwater or near the surface. Some species have a pelagic prejuvenile stage that lives near the surface in the open ocean. Adults are found over soft (sandy or muddy) or hard (rocky) substrata.

Behavior

Little is known of the behavior of zeiforms, as they live at depths where they are difficult to observe. Adults of the John dory (Z. faber) are mainly solitary. The buckler dory (Z. conchifer) often occurs in small aggregations.

Feeding ecology and diet

Zeiform fishes are carnivores; they feed mainly on a variety of fishes but also consume cephalopods and crustaceans. Juveniles of the larger species and adults of the dwarf dories (family Zeniontidae) and tinselfishes (family Grammicolepidae) feed on zooplankton (e.g., copepods, pteropods, fish and crustacean larvae). Adults of the larger zeiform fishes (John dory and the buckler dory) are eaten only by large piscivorous predators (e.g., some sharks, goosefishes [Lophus species], and lancetfishes [Alepisaurus species]). Juveniles and adults of small zeiforms (e.g., Zenion species) are subject to predation by a variety of piscivores.

Reproductive biology

The sexes are separate. Females grow larger than males. Spawning has not been observed, but zeiforms apparently are "broadcast spawners," with the eggs and sperm released into the water column and fertilization taking place in the sea. The eggs and larvae are pelagic and float near the surface; the eggs are spherical, 0.04–0.1 in (1–2.8 mm) in diameter, with a single oil globule. There are no reports of nests or egg guarding or parental care in zeiform fishes.

Conservation status

None of the zeiform fishes are listed by the IUCN. Some populations probably are overexploited in areas where intensive trawling takes place.

Significance to humans

The larger species of zeiforms are of some commercial importance as food fishes. Most zeiforms are caught by trawlers, but anglers also catch a few of the larger species.

Species accounts

List of Species

Red boarfish

Antigonia rubescens

family

Caproidae

taxonomy

Antigonia rubescens Günther, 1860, Japan.

other common names

English: Pink boarfish.

physical characteristics

Attains a total length (including the tail fin) of about 6 in (15 cm). The head and body are deep, greatly compressed, and shaped like a disk or diamond; the body depth is more than twice the head length. The dorsal head profile and the predorsal region are concave, and there is a bulge on the nape. The body, cheeks, and operculum are covered with small ctenoid scales. The mouth is small, and the upper jaw is shorter than the eye diameter. The dorsal fin has nine spines and 27–30 soft rays; the anal fin has three spines and 24–28 rays. The tail fin has 10 branched rays. The body is pale reddish silvery, with a dark red bar from the dorsal fin origin to the origin of the anal fin; another red bar lies above and below the eye, and there is a red band on the body at the rear of dorsal and anal fin bases. The abdomen and lower rear part of the head are silvery white.

distribution

Japan, Midway Island northwest of Hawaii, Taiwan, Philippines, China, and Australia.

habitat

Adults are usually caught with trawls near the bottom at depths of 333–3,000 ft (100–900 m).

behavior

The behavior of the red boarfish is poorly known, because it lives too deep to be observed easily. They occur in large aggregations, as many individuals may be caught in a single trawl haul.

feeding ecology and diet

No information has been published on the diet of the red boarfish. Probably feeds on plankton and small benthic invertebrates. Subject to predation by various piscivores that inhabit the outer shelf and slope region (e.g., sharks, lancetfishes, and gempylids).

reproductive biology

Poorly known. Probably a broadcast spawner, with pelagic eggs and larvae.

conservation status

Not listed by IUCN.

significance to humans

None known.

Thorny tinselfish

Grammicolepis brachiusculus

family

Grammicolepidae

taxonomy

Grammicolepis brachiusculus Poey, 1873, Cuba.

other common names

English: Diamond dory, deepscale dory; Spanish: Palometa oropel.

physical characteristics

Attains a total length (including the tail fin) of about 2 ft, 5 in (72 cm) and a weight of 9 lb (4 kg). The head and body are deep, very compressed, and shaped like an oblong disc, or dinner platter; juveniles are diamond-shaped. The body, cheeks, and operculum are covered with vertically elongated scales. The mouth is small, and the maxilla has two or three ridges, bound to the ascending processes of the premaxillae and loosely connected to the palatines. The jaws have one or two rows of minute, slender teeth, and the vomer may or may not have three or four minute, slender teeth. There are two dorsal fins, the first with six to seven spines and the second with 31–34 unbranched soft rays. The anal fin has two spines and 33–35 unbranched rays. The tail fin has 13 branched rays. The pelvic fins have one spine and six branched rays; there is a row of 34–36 small spines along each side of the dorsal and anal fin bases.

Juveniles have a greatly elongated first anal fin spine and second dorsal fin spine. The pelagic juvenile stage (4–8 in, or 10–20 cm in standard length) looks quite different from the adult. The body is more compressed and angular, with 10–13 prominent, flattened, bladelike, spiny scutes projecting laterally from the surface of each side of the body. Each scute is an out-growth from a scale whose basal part is divided, overlapping both sides of the one behind. On the base of the larger scutes are retrorse spinules. The scutes become smaller as the fish grows, and they eventually shrink to nothing as the fish transforms to an adult at about 10–12 in (25–30 cm). Adults are silvery in color, with bronze reflections. Juveniles are silvery, with irregular black blotches on the body, black spots on the tail fin, and 5 black bars on anal fin.

distribution

Tropical and temperate waters of the eastern Atlantic from France to South Africa; also known from Japan, Hawaii, Australia, and the western Atlantic from Canada to Suriname.

habitat

Adults usually are caught with trawls near the bottom, at depths of 1,333–3,000 ft (400–900 m).

behavior

The behavior of the thorny tinselfish is poorly known, as this species has yet to be observed in shallow water.

feeding ecology and diet

No information has been published on the diet of the thorny tinselfish. It probably feeds on plankton and small benthic invertebrates. Juveniles are subject to predation by a variety of piscivores. Adults are eaten by some large sharks and lancetfishes.

reproductive biology

Poorly known. Probably a broadcast spawner with pelagic eggs and larvae.

conservation status

Not listed by the IUCN. Its apparent rarity may be due to the difficulty in sampling fishes from depths of 1,333–3,000 ft (400–900 m).

significance to humans

None known.

Tinselfish

Xenolepidichthys dalgleishi

family

Grammicolepidae

taxonomy

Xenolepidichthys dalgleishi Gilchrist, 1922, Natal, South Africa.

other common names

English: Diamond dory; spotted tinselfish.

physical characteristics

Attains a total length (including tail fin) of about 6 in (15 cm). The head and body are greatly compressed and vertically elongated, shaped like a flattened diamond. The body, cheeks, and operculum are covered with vertically elongated scales. The mouth is small and the maxilla, which has two or three ridges, is bound to the ascending processes of the premaxillae and loosely connected to the palatines. The jaws have one or two rows of minute, slender teeth; the vomer has a few minute, slender teeth. There are two dorsal fins, the first with five spines and the second with 27–30 unbranched soft rays. The anal fin has two spines and 27–29 unbranched rays. The tail fin has 13 branched rays, and the pelvic fins have one spine and six branched rays. There is a row of 29 small spines along each side of the dorsal fin and 26–27 small spines along the anal fin base. Juveniles have a greatly elongated (two or three times the length of the fish) first anal fin spine and second dorsal fin spine. The body is silvery with scattered, round black spots; the rear margin of tail fin is dusky.

distribution

Western Atlantic from Canada to southern Brazil and eastern Atlantic from Senegal to South Africa; also in Japan, Taiwan, Philippines, Australia, New Zealand, New Caledonia, Fiji, and Tonga.

habitat

Usually taken near the bottom in depths of 666–1,333 ft (200–400 m) but also taken in midwater and at the surface of the open ocean.

behavior

The behavior of the tinselfish is poorly known, as this species is rarely observed in shallow water.

feeding ecology and diet

There is no information. Probably feeds on zooplankton and small benthic invertebrates. The tinselfish is prey for some sharks, lancetfishes, scombrids, carangids, and gempylids.

reproductive biology

There is no information. Probably a broadcast spawner.

conservation status

Not listed by IUCN.

significance to humans

None known.

Buckler dory

Zenopsis conchifer

family

Zeidae

taxonomy

Zeus conchifer Lowe, 1852, Madeira.

other common names

English: Mirror dory (Australia), sailfin dory, silvery John Dory; Portuguese: Peixe galo, falo branco.

physical characteristics

Attains a total length (including the tail fin) of 2 ft, 8 in (80 cm) and a weight of 9 lb (4 kg). The head and body are very compressed and shaped like an oblong disc, or dinner platter. The dorsal head profile is slightly concave. Dorsal fin spines of the adult are elongated and filamentous. The body has no scales. There is a row of five to eight bony bucklers, each with a strong spine, along each side of the bases of the anal and both dorsal fins. There are two or three bucklers on the isthmus and ventral midline of the chest in front of the pelvic fins and seven or eight pairs of bucklers (ridged bony scutes) along the ventral midline of the abdomen, from the base of the pelvic fins to the first anal fin spine. The thoracic region is compressed. The pelvic fins are large and close together, inserted on the chest below the eyes and well in front of the pectoral fins. Each has six or seven soft rays. The first pelvic ray could be considered a spine, because it is unbranched and not segmented, but (unlike the pelvic fin spine of Zeus) it is a biserial (double) ray. There are five to eight bony bucklers (enlarged, platelike scales) along the base of the soft dorsal and anal fins and seven to 10 pairs of spiny scutes along the belly, from the base of the pelvic fins to the anus. The dorsal fin has nine to 10 spines and 24–27 rays; the anal fin has three spines (the first two movable and the third fixed) and 24–26 rays. The caudal fin has 11 branched rays; the pectoral fins are much smaller than the pelvic fins, with 12 or 13 rays. Dorsal, anal, and pectoral fin rays are unbranched. Adults are silvery with a faint, dusky mid-lateral spot above the pectoral fin and below the lateral line. Small juveniles, 1–4 in (2–10 cm) long, are covered with scattered small black spots.

distribution

Mainly continental. Western Atlantic from Canada to Argentina, including the Gulf of Mexico and Caribbean Sea, and eastern Atlantic from France and British Isles to South Africa. Also in the Indian Ocean from South Africa to Kenya and India.

habitat

Adults are found over soft (sandy or muddy) or hard (rocky) substrata. They are demersal, usually caught near the bottom of the continental shelf or upper continental slope region at depths of 33–1,188 ft (10–360 m). Occasionally found in mid-water well above the bottom.

behavior

The behavior of the buckler dory is poorly known, as this species is rarely observed in shallow water. Adults usually are found in aggregations near the bottom. A slow, stalking, ambush-predator mode of hunting is assumed, and the greatly protrusible upper jaw compensates for the feeble swimming musculature.

feeding ecology and diet

Feeds on demersal fishes, crustaceans, and cephalopods and often makes excursions above the bottom to feed on midwater fishes. Prey species selection is influenced by availability and accessibility. The dominant species in the diet is likely to be the most abundant prey in the habitat. Adult buckler dory have few predators other than great white sharks and goosefish (Lophius spp.). Juveniles are likely prey for most piscivorous predators.

reproductive biology

Reproduction is similar to that of the John dory (see following account).

conservation status

Not listed by IUCN. The species is taken as bycatch in various trawl fisheries in the North Atlantic, off southern Brazil, Namibia, and South Africa. There are no fishing regulations or catch data that apply specifically to the buckler dory.

significance to humans

None known.

John dory

Zeus faber

family

Zeidae

taxonomy

Zeus faber Linnaeus, 1758, habitat in Pelago.

other common names

French: Saint-Pierre; German: Heringskönig; Spanish: Pez de San Pedro, gallo de San Pedro, barbero, gallo barbero; Portuguese: Peixe galo.

physical characteristics

Attains a total length (including the tail fin) of 3 ft (90 cm) and a weight of 18 lb (8.2 kg). The head and body are very compressed and shaped like an oblong disc, or dinner platter. The dorsal head profile is straight. The dorsal fin spines of the adult are greatly elongated and filamentous. Scales are minute, cycloid, and embedded; there are five to eight bony bucklers (enlarged, platelike scales) along the base of the soft dorsal and anal fins and seven to 10 pairs of spiny scutes along the belly from the base of the pelvic fins to the anus. The dorsal fin has nine to 11 spines and 22–24 rays, and the anal fin has four spines and 20–23 rays. Dorsal, anal, and pectoral fin rays are unbranched, and the caudal fin has 11 branched rays. The pelvic fins are enlarged, situated below and slightly in front of the pectoral fins, with one spine and six or seven branched rays. The pectoral fins are much smaller than the pelvic fins.

The head and body are silvery to olive-brown, with a conspicuous yellow or white-edged, blue-black ocellus on the middle of the body below the lateral line. The body often has indistinct dark, wavy stripes. The dorsal fin spine filaments are white. Juveniles are small, about 1 in (2–3 cm) long. Color is brownish, with small black spots along the base of the median fins, curved white and black stripes on the head and body, and the usual round black spot on the middle of the body below the lateral line.

distribution

Eastern Atlantic from the North Sea to South Africa, including the Mediterranean Sea, Black Sea, Azores, Madeira, and Canary

and Cape Verde Islands and along the west coast of Africa to Namibia. In the western Indian Ocean from South Africa to Somalia and India. In the western Pacific in Indonesia, the Philippines, China, Taiwan, Japan, Australia, and New Zealand. Not known from the eastern Pacific or western Atlantic.

habitat

Adults are found over soft (sandy or muddy) or hard (rocky) substrata; they are demersal, usually caught near the bottom of the continental shelf or upper continental slope region at depths of 33–1,188 ft (10–360 m). Adults also frequently occur in midwater or near the surface.

behavior

Adults usually are solitary, although they probably congregate for spawning. They typically occur near the bottom, but they often make excursions above the bottom to feed on midwater fishes. With its greatly compressed head and body, large mouth, and extremely protrusible upper jaw, the John Dory is a successful ambush predator. It slowly approaches an unsuspecting small fish by means of undulating waves of the transparent soft dorsal and anal fins. In one quick motion it drops the "trapdoor" lower jaw, shoots out the upper jaw, and expands the gill cavity, sucking in the hapless prey along with a considerable volume of water.

feeding ecology and diet

The John dory is a carnivore, feeding primarily on fishes, crustaceans, and cephalopods. Prey species selection is influenced by the availability and accessibility of prey. The dominant species in the diet is likely to be the most abundant prey in the habitat. In the North Atlantic, juveniles, 3–10 in (8–25 cm), feed mainly on benthic fishes, for example, dragonets (family Callionymidae) and Norway pout (family Gadidae). Adults, 10–22 in (25–56 cm), switch to a diet of larger schooling fishes, primarily blue whiting, Micromesistius poutassou (Risso, 1826), and snipefish, Macroramphosus scolopax (Linnaeus, 1758), of the family Macroramphosidae. Adults have few predators, except for great white sharks and goosefishes (Lophius spp.). Juveniles are probably eaten by piscivorous predators.

reproductive biology

The John dory is a broadcast spawner. The eggs and sperm are shed simultaneously into the water, where fertilization takes place. In the northeastern Atlantic, spawning occurs in April or May. The eggs and larvae are pelagic; the eggs are small (0.1 in or 2 mm diameter) and spherical, with a large oil globule. No nest building or parental care has been reported for the species. In the North Atlantic, John dory males mature in their second or third year at a length of about 10 in (26 cm), and they may live to an age of 13 years. Females are three or four years old when they become mature at a length of 14 in (35 cm). They grow larger than males do, and they may live 15 years. In New Zealand waters males attain maturity at 11.8 in (30 cm) in two years; their growth then slows considerably, with a maximum size of about 15.7 in (40 cm) and an age of nine years. Females grow faster and attain a maximum size of about 18.1 in (46 cm) and an age of nine years.

conservation status

Not listed by IUCN. As of 1998 there were no regulations regarding John dory fishery in the English Channel or in Australian waters. The species is taken as bycatch in the trawl fisheries of the eastern Atlantic, South Africa, Australia, New Zealand, Indonesia, Philippines, Taiwan, and Japan.

significance to humans

The John dory is prized as a food fish and is of commercial importance because of its high price in fish markets.

Resources

Books

Karrer, C., and P. C. Heemstra. "Family No. 140: Grammicolepididae." In Smiths' Sea Fishes, edited by M. M. Smith and P. C. Heemstra. Johannesburg: Macmillan South Africa, 1986.

Nelson, J. S. Fishes of the World. 3rd edition. New York: John Wiley & Sons. 1994.

Smith, M. M., and P. C. Heemstra, eds. Smiths' Sea Fishes. Johannesburg: Macmillan South Africa. 1986.

Periodicals

Barnard, H. K. "A Monograph of the Marine Fishes of South Africa. Part 1." Annals of the South African Museum 21(1925): 1–105.

Bigelow, H. B., and W. C. Schroeder. "Fishes of the Gulf of Maine." U.S. Fish and Wildlife Service Fishery Bulletin 53(1953): 1–577.

Dunn, M. R. "The Biology and Exploitation of John Dory, Zeus faber (Linnaeus, 1758), in Waters of England and Wales." ICES Journal of Marine Science 58, no. 1 (2001): 96–105.

Fowler, H. W. "The Buckler Dory and Descriptions of Three New Fishes from off New Jersey and Florida." Proceedings of the Academy of Natural Sciences of Philadelphia 86 (1934): 353–361.

Greenwood, P. H., D. E. Rosen, S. H. Weitzman, and G. S. Myers. "Phyletic Studies of Teleostean Fishes, with a Provisional Classification of Living Forms." Bulletin of the American Museum of Natural History 131, no. 4 (1966): 339–455.

Heemstra, P. C. "A Revision of the Zeid Fishes (Zeiformes: Zeidae) of South Africa." Ichthyology Bulletin of the J. L. B. Smith Institute of Ichthyology 41 (1980): 1–18.

Parin, N., and O. D. Borodulina. "Preliminary Review of the Bathypelagic Fish Genus Antigonia Lowe (Zeiformes, Caproidae)." Transactions of the P. P. Shirsov Institute of Oceanology 121 (1986): 1–105, 141–172.

Yoneda, M., S. Yamasaki, K. Yamamoto, H. Horikawa, and M. Matsuyama. "Age and Growth of John Dory, Zeus faber (Linnaeus, 1758), in the East China Sea." ICES Journal of Marine Science 59, no. 4 (2002): 749–756.

Phillip C. Heemstra, PhD