lamp shells

▪ animal

Introduction

also called  brachiopod,  

      any member of the phylum Brachiopoda, a group of bottom-dwelling marine invertebrates. They are covered by two valves, or shells; one valve covers the dorsal, or top, side; the other covers the ventral, or bottom, side. The valves, of unequal size, are bilaterally symmetrical; i.e., the right and left sides are mirror images of one another. Brachiopods (from the Greek words meaning “arm” and “foot”) are commonly known as lamp shells because they resemble early Roman oil lamps.

      Brachiopods occur in all oceans. Although no longer numerous, they were once one of the most abundant forms of life.

      Members of this phylum first appeared rather early in zoological history. It is possible, by means of fossil representatives, to survey their evolution from the Cambrian Period (about 542,000,000 years ago) to the present. Although some of the evolutionary development is revealed, it is still imperfectly understood. Other than their usefulness in dating geological periods, members of this phylum have no economic value, except as curios and museum pieces.

General features

Size range and diversity of structure

      Most brachiopods are small, 2.5 centimetres (about one inch) or less in length or width; some are minute, measuring one millimetre (more than ¹/₃₀ of an inch) or slightly more; some fossil forms are relative giants—about 38 centimetres (15 inches) wide. The largest modern brachiopod is about 10 centimetres (four inches) in length.

      Great diversity existed among brachiopods in the past; modern brachiopods, however, exhibit little variety. They are commonly tongue-shaped and oval lengthwise and in cross section. The surface may be smooth, spiny, covered with platelike structures, or ridged. Most modern brachiopods are yellowish or white, but some have red stripes or spots; others are pink, brown, or dark gray. The tongue-shaped shells (Lingula) are brown with dark-green splotches; rarely, they are cream yellow and green.

Distribution and abundance

      Today, brachiopods, numbering about 300 species representing 80 genera, are abundant only locally. In parts of the Antarctic they outnumber all other large invertebrates. They are common in the waters around Japan, southern Australia, and New Zealand. Although rare in the Indian Ocean, some unusual types are common along the coast of South Africa. In Caribbean and West Indian waters, 12 species occur. The east and west coasts of the North Atlantic Ocean are sparsely occupied by brachiopods; the waters around the British Isles contain a few species, and a few genera live in the Mediterranean Sea. The West Coast of the United States and Hawaii have a number of brachiopod species, and the coasts of Chile and Argentina have a considerable variety, including the largest living species. Some live in the polar regions, and a few are abyssal; i.e., they inhabit deep parts of the ocean.

Natural history

Reproduction

      Not much is known about the reproduction of brachiopods. Except in three genera, the sexes are separate. Eggs and sperm are discharged into the mantle cavity through funnel-shaped nephridia, or excretory organs, on each side of the mouth. Fertilization takes place outside the shell. In a few genera the young develop inside the female in brood pouches formed by a fold of the mantle, a soft extension of the body wall. Some fossil forms had internal cavities that may have served as brood chambers. The egg develops into a free-swimming larva that settles to the bottom. The free-swimming stage of the articulate brachiopods (whose valves articulate by means of teeth and sockets) lasts only a few days, but that of the inarticulates may last a month or six weeks. In inarticulate larvae the pedicle, a stalklike organ, develops from a so-called mantle fold along the valve margin; in articulates it develops from the caudal, or hind, region.

Behaviour and ecology

      About 60 percent of brachiopods live in shallow water (less than 100 fathoms—about 180 metres [600 feet]) on the shelf areas around the continents. More than 35 percent occupy waters deeper than 100 fathoms, and a few live in the abyss down to more than 6,000 metres (about 20,000 feet). Lingula lives from the tidal zone to 23 fathoms (about 42 metres [138 feet]). Most modern branchiopods anchor by the pedicle to pebbles, to the undersides of stones, or to other hard objects. They prefer quiet water and protected surroundings. Lingula lives in mud or sand and is attached at the bottom of its burrow.

      Brachiopods feed by opening the shell and bringing in food-bearing currents by lashing of the cilia (hairlike structures) attached to the filaments of the lophophore, a horseshoe-shaped organ that filters (filter feeding) food particles from the seawater. Cilia in lophophore grooves bring food particles, often trapped in mucus, to the mouth. Brachiopods feed on minute organisms or organic particles. Articulate brachiopods, which have a blind intestine, may depend partly on dissolved nutrients.

      Shells of some articulate brachiopods have a fold, which forms a trilobed anterior that helps keep lateral, incoming food-bearing currents separated from outgoing, waste-bearing currents. When feeding, Lingula protrudes its anterior (front) end above the mud and arranges its setae (bristle-like structures) into three tubes. These channel the water into lateral incoming and medial, or central, outgoing currents. Some coralliform brachiopods of the Permian Period (299,000,000 to 251,000,000 years ago) are thought to have fed by rapid beating of the dorsal valve, causing a sucking in and expulsion of food-bearing water. Some ostreiform (oyster-shaped) types of the same period are believed to have fed by gentle pulsation of the dorsal valve.

Form and function

      Two major groups of brachiopods are recognized, based on the presence or absence of articulation of the valves by teeth and sockets. The valves of inarticulate brachiopods are held together by muscles. Lingula, with its elongated, tonguelike shell, is an example. Its convex valves bulge outward at the middle and taper posteriorly, or away from the hinge. A long, fleshy pedicle protrudes between the valves at the tapered end. The pedicle of Lingula differs from that of most other brachiopods in being flexible and capable of movement—an aid in burrowing and in attaching the animal in its burrow. The shell interior is divided into posterior coelomic (internal-body) and anterior mantle cavities. The internal organs are located in the coelom. The digestive system consists of mouth, gullet, stomach, intestine, and anus, all surrounded by a liver, or digestive gland. A complex set of muscles opens the valves and slides them laterally, or sideways, when feeding. The mantle cavity is occupied by the lophophore. Lingula lives in a burrow in mud or sand with the tip of its pedicle attached in mucus at the bottom of the burrow. The contractile pedicle permits extension of the shell when feeding or retraction if the animal is startled.

      The articulate-brachiopod shell is typified by Waltonia, which is small (about two centimetres [3/4 inch]) and red in colour, with a smooth or slightly ridged shell. This type of shell is more highly specialized than that of most inarticulate species and is composed of three layers. The outer layer, called periostracum, is made of organic substance and is seldom seen in fossils. A middle layer consists of calcium carbonate (calcite). The inner layer is composed of calcite fibres and may be punctate—i.e., perforated by minute pits—or it may be pseudopunctate, with rods (taleolae) of calcite vertical to the surface. Impunctate shells have neither pits nor taleolae.

      Many hinged brachiopods attach to the substrate, or surface, by a tough, fibrous pedicle; but some specialized forms are cemented to the substrate by the beak of the ventral valve. Cemented forms are commonly distorted, scalelike, or oyster shaped or resemble a cup coral. The pedicle of some brachiopods is atrophied; their shells lie loose on the sea floor.

      The shell of an articulate brachiopod tapers posteriorly to a beak. The ventral valve is usually the larger. The hinge may be narrow or wide. Many hinged genera have a flat or curved shelf, called the palintrope, between the beak and the hinge line. The ventral palintrope is divided at the middle by the delthyrium, a triangular opening for the pedicle. The delthyrium may remain open or be wholly or partly closed by small plates growing from its margins. In some families the delthyrium is closed completely or partly by one plate, the pseudodeltidium, anchored to the delthyrial margins. The articulating teeth occur at the angles of the delthyrium and may or may not be supported by vertical dental plates, which may be separate or united to form a so-called spondylium. Teeth are of two types, deltidiodont and cyrtomatodont. Deltidiodont teeth grow anteriorly with the palintrope and leave a growth path along the delthyrial edge; cyrtomatodont teeth are knoblike and occur in shells without a hinge line. They grow anteriorly but are kept knoblike by posterior resorption.

      The dorsal valve contains structures called crura that diverge from the beak. In some fossil forms the crural bases (brachiophores) bound a triangular cavity, the notothyrium, in which the diductor, or opening, muscles (muscle) are attached onto the floor or to a ridge, or boss, called the cardinal process at the apex. The notothyrium may be closed by a solid plate, the chilidium. In more highly developed genera a hinge plate bearing the pedicle or dorsal adjustor muscles occurs between the crural bases. The hinge plate is said to be divided when it is incomplete but undivided when it forms a flat or concave structure. The hinge plate is often supported by a median septum, or wall. The hinge sockets are located between the inside shell wall and a socket ridge to which the hinge plate is attached. In many specialized genera the crura support calcareous loops or spires (brachidia), the inner skeleton of the lophophore. Structures corresponding in function but of different origin and with different names occur in the pedicle region of some inarticulate brachiopods.

      The fleshy body of the articulate brachiopod is divided transversely by the body wall into a posterior visceral cavity filled with coelomic fluid and an anterior mantle cavity filled with seawater. The visceral cavity contains the U-shaped digestive canal, four reproductive glands, and a liver, or digestive gland, held in place by mesenteries (sheets of tissue). Extensions of the coelom into the mantle hold the eggs and sperm. The mouth leads into a saclike stomach that ends in an intestine; there is no anus. The liver surrounds the stomach. Waste is excreted through the mouth. The nervous system, which consists of two principal ganglia, or nerve centres, encircles the esophagus and sends branches to other parts of the body. One pair of excretory organs (nephridia) occurs in most brachiopods, but two pairs may be present.

      The mantle cavity is lined by the thin, shell-secreting mantle that is fringed by setae at its edges. Within the mantle cavity is the lophophore, which may be a simple or complicated loop, often horseshoe-shaped. Ciliated filaments along the loop direct food-bearing currents to the mouth, which is located on the body wall between the branches of the lophophore and crura.

      The shell opens by contraction of diductor muscles that extend from near the centre of the ventral valve to the process under the dorsal beak. These muscles pull the dorsal beak forward, rotating it on a line joining the hinge teeth. Contraction of the adductor muscles closes the valves; in the ventral valve the adductors are located between the diductors. Pedicle muscles or adjustors extending from the pedicle to the hinge plate of the dorsal valve rotate the shell on the pedicle. In places where the muscles are attached to the shell there are scars, which are helpful in the identification of genera.

Paleontology

      Brachiopods were among the first animals to appear at the beginning of the Cambrian Period (542,000,000 years ago). Their evolution and distribution was wide and rapid. More than 35,000 species in more than 2,500 genera are known, and the number of described species increases yearly. Articulate and inarticulate brachiopods appeared at the same time in a relatively advanced state of development, indicating a long evolution from forms without shells, an evolution apparently lost or unrecorded in Precambrian times.

      The Inarticulata, the most abundant brachiopods of the Cambrian, soon gave way to the Articulata and declined greatly in number and variety toward the end of the Cambrian. They were represented in the Ordovician (488,000,000 to 444,000,000 years ago) but decreased thereafter. In the Cretaceous (146,000,000 to 66,000,000 years ago) the punctate calcareous Inarticulata proliferated, but this trend soon ended. The Inarticulata dwindled through the Cenozoic (66,000,000 years ago) to the Recent. Only nine genera are known during the Recent Epoch (last 10,000 years). Inarticulate genera represent about 6.5 percent of all brachiopod genera.

      The Articulata, diverse and most numerous from Ordovician times to the present, were, in the Cambrian, represented by several specialized forms. Articulate evolution tended toward shell elaboration for bottom dwelling and perfection of feeding mechanisms from the simple looped lophophore to the elaborate lobate and spiral forms. The Orthida, the most common articulate brachiopods of the Cambrian and Ordovician, decreased in numbers after the Ordovician, and the impunctate Orthida became extinct in the Early Devonian (416,000,000 to 398,000,000 years ago); the punctate Orthida lingered into the Permian Period (299,000,000 to 251,000,000 years ago). The Strophomenida appeared in the Early Ordovician and increased rapidly. They were abundant and varied in the Devonian, becoming even more so by Permian times. This large order became greatly reduced at the end of the Permian Period. The Pentamerida, never prolific, flourished in the Ordovician; an evolutional burst of huge forms occurred in the Silurian (444,000,000 to 416,000,000 years ago), but after that the pentamerids decreased into the Devonian (416,000,000 to 359,000,000 years ago) and became extinct early in the late part of that period. The Spiriferida are conspicuous for the great elaboration of the spiral brachidium. They appeared in the Ordovician, were widely distributed into the Permian, and survived into the Jurassic, which began about 200,000,000 years ago. The Rhynchonellida were abundant from mid-Ordovician throughout the Paleozoic. They survived into the Triassic (251,000,000 to 200,000,000 years ago) and had a rebirth in the Jurassic, after which they declined into the Cenozoic. They now number only 14 genera.

      The Terebratulida, now the dominant group, appeared in the early Devonian and rapidly expanded in the mid-Devonian to produce a number of gigantic forms; a few long-looped and short-looped genera persisted into the Permian. The Terebratulida survived the Permian and were widely distributed in the Triassic and evolved into a great variety of forms in the Jurassic, especially the short-looped types. Decline of the short-looped terebratulids began in the Late Cretaceous (100,000,000 to 66,000,000 years ago); they have continued to dwindle into the present and are now outnumbered by the long-looped terebratulids.

Classification

Distinguishing taxonomic features

      Brachiopods possess a lophophore (a feeding structure that filters food from seawater), excretory organs (nephridia), and simple circulatory, nervous, and reproductive systems. Brachiopods have usually been divided into two classes, Articulata and Inarticulata.

Annotated classification

      The classification below is based on that proposed by A. Williams and A.J. Rowell in 1965 in Treatise on Invertebrate Paleontology.

Phylum Brachiopoda (lamp shells)

 Marine invertebrates with two valves, or shells; lophophore horseshoe-shaped; about 300 living species known; more than 30,000 extinct species described; occur in all oceans.

      Class Inarticulata

 Shell does not articulate, is usually composed of chitinophosphatic material; shell muscles complex; pedicle (stalk) develops from ventral mantle, a soft extension of the body wall; intestine with anal opening.

      Order Lingulida (lingulid)

 Shell usually contains phosphate, rarely calcareous, biconvex (i.e., both valves convex), beak for attachment to surface apical, or located at the tip, in both valves; fleshy pedicle emerging between the valves at the tapered end; about 51 genera; Cambrian to Recent.

      Order Acrotretida

 Usually circular in outline; shell either contains phosphate or is punctate calcareous; pedicle opening confined to the ventral valve; 62 genera; Early Cambrian to Recent.

      Order Obolellida

 Mostly calcareous, biconvex, shape nearly circular to elongated; position of pedicle opening variable; dorsal valve with marginal beak; 5 genera; Early to mid-Cambrian.

      Order Paterinida

 Shell with phosphate, rounded or elliptical; pedicle opening partly closed by cover called homeodeltidium; dorsal valve similar to the ventral but with a convex homeochilidium; 7 genera; Early Cambrian to mid-Ordovician.

      Class Articulata

 Shells articulate by means of teeth and sockets; shells always calcareous; musculature less complicated than in Inarticulata; larval pedicle develops from rear region; no outside opening from intestine.

      Order Kutorginida

 Calcareous, biconvex interarea (smooth surface in area between beak and hinge line) present; delthyrium (opening in the pedicle) closed by a plate, the pseudodeltidium; dorsal valve with interarea; muscle area narrow and elongated in both valves; 3 genera; Early to mid-Cambrian.

      Order Orthida

 Usually biconvex, wide-hinged, with interareas in both valves; teeth deltidiodont (leave a growth path along margin of pedicle opening); hinge structures consist of brachiophores (supporting structures), shell substance punctate or impunctate—i.e., with or without pits; more than 200 genera; Early Cambrian through Permian.

      Order Strophomenida

 Teeth deltidiodont when present; ventral muscles large; shell substance pseudopunctate (with rods of calcite), rarely impunctate; more than 400 genera; mid-Ordovician to Early Jurassic.

      Order Pentamerida

 Biconvex, ventral valve usually with a spondylium (united dental plates); delthyrium usually open; dorsal-valve brachiophores supported by bracing plates; impunctate; nearly 100 genera; mid-Cambrian to Late Devonian.

      Order Rhynchonellida

 Narrow-hinged with functional pedicle; dorsal valve with or without a median septum; lophophore (of Recent genera) dorsally spiral and attached to crura (supporting structures); spondylia rare; nearly 300 genera; Ordovician to Recent.

      Order Spiriferida

 Lophophore supported by a calcareous spiral structure (brachidium); punctate or impunctate, usually biconvex; delthyrium open or closed; more than 300 genera; mid-Ordovician to Jurassic.

      Order Terebratulida

 Pedicle functional, cyrtomatodont teeth; lophophore supported wholly or in part by a calcareous loop, short or long and free or attached to a median septum; more than 300 genera; Early Devonian to Recent.

Critical appraisal

      The classes Articulata and Inarticulata were first proposed by T.H. Huxley in 1869. Before 1932 they were further subdivided into four orders based on the imperfectly known larval development and formation of the shell around the pedicle opening. In 1927 a fifth order was proposed, and it was suggested that a classification be based on the pedicle development of the larvae.

      Most brachiopods are extinct, and larval development can only be conjectured. Because of this, the early classification schemes have been abandoned. Eleven orders distributed in Huxley's classes have been retained in the present classification, which is still being modified. On the basis of hinge and tooth types some systematists have divided the Articulata into two subclasses, Protremata and Telotremata. The Protremata are wide-hinged forms with deltidiodont teeth. The Telotremata are narrow-hinged brachiopods with cyrtomatodont teeth.

Additional Reading

There are few works specifically on lamp shells. M.J.S. Rudwick, Living and Fossil Brachiopods (1970), is a modern, readable, and comprehensive account. Raymond C. Moore (ed.), Treatise on Invertebrate Paleontology, pt. H, Brachiopoda, 2 vol. (1965), is a technical, thorough account of brachiopods and their classification, with extensive topical bibliographies. Rex A. Doescher, Living and Fossil Brachiopod Genera, 1775–1979: Lists and Bibliography (1981), is also of interest.G. Arthur Cooper Ed.

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