ECOLOGY OF INARTICULATED BRACHIOPODS

CHRISTIAN C. EMIG [Centre d’Océanologie de Marseille]

INTRODUCTION need to be analyzed carefully at the popula- tion level before using them to interpret spe- Living inarticulated brachiopods are a cies and genera. highly diversified group. All are marine, with Assemblages with lingulides are routinely most species extending from the littoral wa- interpreted as indicating intertidal, brackish, ters to the bathyal zone. Only one species and warm conditions, but the evidence for reaches abyssal depths, and none is restricted such assumptions is mainly anecdotal. In to the intertidal zone. Among the living fact, formation of lingulide fossil beds gen- brachiopods, the lingulides, which have been erally occurred during drastic to catastrophic most extensively studied, are the only well- ecological changes. known group. Both living lingulide genera, Lingula and BEHAVIOR Glottidia, are the sole extant representatives INFAUNAL PATTERN: LINGULIDAE of a Paleozoic inarticulated group that have evolved an infaunal habit. They have a range Burrows of morphological, physiological, and behav- Lingulides live in a vertical burrow in a ioral features that have adapted them for an soft substrate. Their burrow has two parts endobiont mode of life that has remained (Fig. 407): the upper part, oval in section, remarkably constant at least since the early about two-thirds of the total length of the Paleozoic. The lingulide group shares many burrow, in which the shell moves along a features that are characteristic of this mode single plane, and the cylindrical lower part in of life including a shell shape that is oblong which only the pedicle moves (EMIG, 1981b, oval or rectangular in outline with straight, 1982). In a homogenous fine sand the length lateral, subparallel to parallel margins and an of the whole burrow is about ten times the anterior margin that is straight to slightly length of the shell (Fig. 407), but it can be concave for burrowing; a complex muscle reduced when the coarse fraction increases at system that operates the inarticulated valves; depth in the sediment or when a hard layer a mantle margin and its setae that serve sev- occurs (EMIG, 1982). In tropical areas, a layer eral basic purposes; and a pedicle that an- formed by pieces of coral and pebbles or by chors the animal at the bottom of the burrow shell fragments often limits the thickness of and shifts the position of the shell. Such the sandy sheet to about 15 to 20 cm. The characteristics can be considered as plesio- pedicle is anchored within this coarse layer, morphic among the Brachiopoda. and the detrital mass of the bulb is less than The ecological requirements of inarticu- that of individuals living in thick, sandy sedi- lated brachiopods indicate the need for a life- ment. The extension of the pedicle can reach history approach that emphasizes aspects of a length 20 times that of the shell to com- populations rather than individuals because pensate for sedimentation (EMIG, 1983a). many such factors as reproduction, survivor- Fossil burrows with lingulide shells in situ ship, dispersion, and evolution depend on show the same structure (Fig. 407). Thus populations. Accordingly there is no single when determining the relationship of the factor that determines the occupancy of a burrow to the length of the shell, the com- niche by a population and that is always di- paction of the sand layer can be estimated at rectly related to the biocoenosis in which the about one-third (EMIG & others, 1978; population is living. Those requirements EMIG, 1982). 474 Brachiopoda

opening

1 cm rubefied layer

1 cm lingulide in situ

5 mm clayey sandstone

faded layer

2

pedicle mass 1a 1b

FIG. 407.1a, Longitudinal section of a burrow of a living lingulide with the shell in normal position and retracted (and 1b, detailed pedicle mass); and 2, of a fossil lingulide (Triassic of Vosges Mountains; Emig & others, 1978).

The walls of the burrow are lined with der coelomic pressure, to reinforce the an- mucus secreted by the edges of the mantle choring in the substratum and is enhanced and the pedicle (EMIG, 1982). The mucous by crenulation of the pedicle bulb. layer binds the walls and lubricates the The lingulides often live in sediment that movements of the animal in its burrow. Only is in a reducing environment below the up- the distal bulb of the pedicle, surrounded by per 2 to 5 centimeters, but the peripheral a mass of sand and various detrital particles substrate, which is up to 1 to 2 mm thick agglutinated by the bulb’s sticky mucous se- along the burrow walls, is oxygenated by cretion, is firmly anchored into the substra- continuously renewed water in the burrow tum at the bottom of the burrow (Fig. 407). (Fig. 407). The size of this mass depends upon charac- Continuous filtering indicates that the teristics of the sediment. Functioning like normal position of the lingulide shell is at the ampulla in the Phoronida, the distal bulb the top of the burrow (EMIG, 1982). To of the lingulides is able, by turgescence un- maintain this position (Fig. 408), a weak Ecology of Inarticulated Brachiopods 475

FIG. 408. Longitudinal section of a lingulide in its burrow; 1, ventral side showing the mantle setae length; 2, nor- mal position (lateral view) by contraction of the lateral muscles; 3, quick valve closure by contraction of the ante- rior and posterior adductor muscles, first step of the escape reflex (new). contraction of the lateral body muscle layer fluid sediments the walls, even when bonded produces a hydrostatic pressure on the by mucous secretion, inadequately support body's coelomic cavity; the body volume is the shell in its normal filtering position shifted posteriorly and laterally; the valves (EMIG, 1983a). gape about 6° to rest against the lateral bur- At the surface of the sediment, three char- row walls, which act as supports; and the lo- acteristic pseudosiphons indicate the pres- phophore extends to become functional ence of a lingulide in normal, life position within an enlarged pallial cavity. The normal (Fig. 407–408, 410; Table 35). They are life position is static and can be maintained shaped by the highly specialized anterior se- without much effort. The pedicle plays no tae of the mantle. At the level of the shortest role in the maintenance of this position. setae, the anterior mantle margin of each Occasional scissorlike movements of the valve develops an epidermal crest. These valves assist in maintenance of the burrow come into contact with each other and in- (Fig. 409). duce tilting and interlacing of the setae Coarser or muddier substrates are less well borne by the crests. Simultaneously the long- suited for providing stable burrow walls. est setae, which can be as long as a third of Consequently, the animal is unable to live or the shell length, remain vertical (Fig. 408). to survive in sediment that is too coarse or The central aperture is exhalant, while the too muddy, contrary to general assumptions two lateral apertures are inhalant. The exhal- about habitats of lingulides. Thus living ant and inhalant water streams are com- lingulides have rarely been found in muddy pletely separated by the mantle crests and sediments with a fine fraction (< 63 µm) internally by tentacle tips without any mix- higher than 35 to 40 percent because in such ing of the flows. The diameter of setae varies 476 Brachiopoda

10°10°

rotational shear

opening quick adduction

slow adduction

slow reopening

3

1

2

FIG. 409. 1, U-shaped reburrowing by Glottidia (Emig & others, 1978); 2, sequence of the scissors burrowing move- ments of G. pyramidata (Thayer & Steele-Petrovic, 1975); 3, patterns in the burrowing sequence of Lingula anatina (Savazzi, 1991). from 15 to 60 µm, and they occur at inter- rent direction appears to be sufficient to vals of 15 to 30 µm so that they exclude large avoid recycling (Fig. 410). particles from the pallial cavity. Contrary to Shell Movements and Burrowing general assumptions (PAINE, 1970; THAYER & STEELE-PETROVIC, 1975), the absence of lin- Shell movements and burrowing behavior gulides from mud is not related to clogging are similar in both extant lingulide genera, of the lophophoral cavity by fine particles. In Lingula and Glottidia (YATSU, 1902b; THAYER a turbid water mass, fine particles may be & STEELE-PETROVIC, 1975; EMIG, 1981b, retained in large masses by the mucus on the 1982, 1983b; TRUEMAN & WONG, 1987; setae of the pseudosiphons and do not enter SAVAZZI, 1991), and have probably been the pallial cavity but are flushed out periodi- practiced by oblong or rectangular lingu- cally by scissorlike movements of the shell loides since early Paleozoic times (EMIG, (EMIG, 1983a). 1984b; SAVAZZI, 1991). No orientation related to current direction Opening and slow closing movements has been observed (WORCESTER, 1969; EMIG, (Fig. 408–409) of the valves are governed by 1981b) because the strong, jetlike, exhalant fluctuations in pressure within the meta- current precludes possible recycling by the coelomic body cavity and are generated by inhalant currents. A turn of the shell plane of contraction of the lateral muscle layers of the about 25 to 30° from the near-bottom cur- body, which are composed of circulo- Ecology of Inarticulated Brachiopods 477

TABLE 35. Summary of the two adult lophophore types in living inarticulated brachiopods in relation to the number of inhalant (in) and exhalant (ex) compartments and apertures in the shell and shell orientation in or on substratum (new).

Taxa Genera Species Shell orientation Schizolophe Spirolophe of Pelagodiscus atlanticus Lingulides 2 12 shell vertical - 2 in + 1 ex1 Craniids 4 19 dorsal valve above, ventral valve below - 2 in + 1 ex1 Discinids 1 1 dorsal valve above, ventral valve below 1 in + 2 ex 3 11 dorsal valve above, ventral valve below - 1 in + 2 ex

1in these groups, there are two small additional exhalant apertures behind the shell. longitudinal fibers. This body cavity func- pulses and, although the shell is moderately tions as a single, fluid-filled chamber, al- gaping, the setae, which prevent sediment though partially divided by a gastroparietal particles from entering the mantle cavity, aid band and, with the coelomic canal of the in the burrowing process. A complete rota- pedicle, acts as a fluid reservoir in the hy- tion takes five to eight seconds. Second, draulic system. This system that opens the there is a slow opening of the valve of one to valves performs the same function as the five seconds in duration, followed by a short elastic hinge ligament of the molluscs and pause (up to three seconds). Third, a slow the diductor muscles of articulated brachio- closure and then reopening of the valves are pods. Quick closure is obtained by the con- followed by a quick contraction of the ad- traction of the anterior and posterior adduc- ductor muscles that forces water jets into the tor muscles. Scissorlike movements of the surrounding sediment. Fourth, there is a valves occur by contraction of the well- pause of variable length. developed, oblique muscles. This complex Progression into the sand coincides with body musculature sustains the unique, infau- large pressure pulses and is facilitated by the nal mode of life of these brachiopods. secretion of a large amount of mucus. Con- When a lingulide is on a sandy substrate, trary to popular belief, the lingulide pedicle fluctuations in pressure within the coelomic is not used for burrowing; it is unable to dig body and pedicle cavities open and close the into the sediment. Instead it acts as a support valve. When the lingulide starts to burrow or prop while repeated scissorlike move- (Fig. 409), the pedicle stiffens with its distal ments, shell closure with water injection, and bulb pressing downward to prop up the shell openings accompanied by pressure valves, thereby bringing the anterior margins pulses result in successively deeper penetra- of the valves into contact with the sediment. tion. Burrowing follows a semicircular Penetration takes place by means of a com- course, the radius of which probably de- bination of scissorlike movements of the pends on shell size. The animal burrows ob- valves and ejection of water from them that liquely downward to a depth that has not yet loosen the sand prior to a downward move- been established in natural conditions, then ment of the shell and an upward transporta- curves upward and burrows vertically until it tion of mucous-bound sand by the lateral reaches the surface of the sediment. Pedicle setae of the mantle. anchoring following burial is achieved by The typical burrowing sequence consists mucoid adhesion of sand and various par- of the following phases (Fig. 409). First, ticles. Some fossil U-shaped burrows could scissorlike movements occur by oscillatory be related to reburrowing features (EMIG & rotation of the valves about an axis passing others, 1978). While reentering the sedi- dorsoventrally through the posterior shell; ment the animal is extremely susceptible to the movements coincide with small, pressure predation. 478 Brachiopoda

FIG. 410. Composite of two transverse sections of a lingulide in its burrow, one section at the level of the anterior mantle margin showing the epidermal crests and inhalant and exhalant opening (shown in heavy lines), the other at the level of the lophophore (shaded in gray) (adapted from Emig, 1982).

Burrowing is faster in small individuals to dig in such coarse sediments (THAYER & than in larger ones, and failure to reburrow STEELE-PETROVIC, 1975; CULTER, 1979). increases in Lingula with shell lengths ex- During rapid experimental sedimentation, ceeding 1.7 to 2 cm (MORSE, 1902; AWATI & autotomy of the pedicle occurs when accu- KSHIRSAGAR, 1957; WORCESTER, 1969; EMIG, mulation exceeds the pedicle extension. A 1981b, 1982, 1983a; HAMMOND, 1983; new pedicle is regenerated in four to eight SAVAZZI, 1991). Reburrowing could be inter- weeks in Lingula, but individuals without a preted as a size-related process, but such a pedicle maintain their filtering position with performance seems to vary between geo- difficulty and generally emerge onto the sedi- graphic populations, as contradicting ac- ment surface. Any damage to the pedicle al- counts have shown (EMIG, 1983a; SAVAZZI, ways impairs burrowing as it precludes the 1991). In experimental conditions the bur- use of the coelom as a hydraulic system. L. rowing speed is always five to ten times faster reevii is able to move pebbles of several cen- than in natural conditions (Table 36). Up- timeters in diameter that happen to lie on ward burrowing is essential for the survival top of its burrow (EMIG, 1981b). of the lingulides and can be accelerated to compensate for sedimentation above their Retraction into the Burrow burrows, perhaps a response to the increase Rapid retraction into the burrow is an es- of the sediment pressure (Table 36). A rapid cape reflex (FRANÇOIS, 1891; MORSE, 1902) influx of coarse sediment, which is not typi- that is well known in almost all animals that cal of the environments of lingulides, how- live in burrows or tubes. This protective re- ever, may occur during high-energy events action in response to unfavorable circum- (HAMMOND, 1983). The nature of the sedi- stances in the external environment is ac- ment has a direct influence on burrowing companied by the rapid closure of the shell. capability, which is about twice as fast in a This response by the lingulides is elicited by sandy substrate as in coarser sediment (par- tactile stimulations of the anterior marginal ticles > 2 mm). In experimental conditions setae (MORSE, 1902; TRUEMAN & WONG, Lingula anatina was able to burrow upward 1987), by an organism moving over the sedi- in coarse sediment but was unable to con- ment surface, or by a shadow falling on the struct a stable burrow and finally emerged brachiopod (EMIG, 1981b). Such stimuli re- onto the sediment surface, often after auto- sult in a quick closure of the shell with expul- tomy of its pedicle. The results are indecisive sion of water combined with contraction of under natural conditions (EMIG, 1983a), but the pedicle muscle, and the animal with- the temperature seems to have no influence draws quite quickly into the burrow. If the on the burrowing speed. Glottidia is unable disturbance continues the animal generally Ecology of Inarticulated Brachiopods 479

TABLE 36. Experimental and in situ (measurements in italics) burrowing conditions (data from a, Emig, 1981b; b, Emig, 1983a; c, Hammond, 1983; d, Paine, 1963; Thayer & Steele-Petrovic, 1975; e, Worcester, 1969). Mean burrowing speed is given in parentheses (new).

L. anatina (b) L. reevei (a, e) G. pyramidata (d)

Burrowing speed in normal conditions (cm/h) experimental 0.5–1.7 (0.9) 0.2–2.5 (0.75) 0.67–2.7 in situ 0.08–0.21 0.21–0.75 < 0.67–2.7

Mean upward speed (cm/h) during experimental sedimentation Thickness of sediment b c e d 10 cm (0.11) (0.45) - (1.3) 15 cm - - (0.14), (1.07) -

20 cm (0.13) (0.58) - (0.33) 30 cm (0.18) - (0.38) (0.40) retracts 1 to 3 cm from the surface into the larva during settlement. All discinids are at- lower section of the upper part of the bur- tached by a highly muscular pedicle to hard row. During retraction the upper part (0.5 to substrates except for Pelagodiscus, which is 1 cm) of the burrow collapses and is obtu- closely fixed to the hard substrate by means rated by sand grains, although in compact of its two main vertical body muscles. The sand the burrow remains open (EMIG, pedicle is very short, and the shell is held 1981b, 1982). near the substratum. Among living inar- At the end of a disturbance, the lingulide ticulated brachiopods only the pedicle of the is elevated by scissorlike and small opening discinids has a dual function. It acts as an movements of the shell combined with the anchor, and it supports the weight of the action of the setae and copious mucous se- shell and holds it in relative position to the cretion, all of which restore the upper part of substrate (Fig. 411.2). the burrow. During retraction and reex- The craniids, which are cemented by the tension, the coelomic pedicle canal functions entire surface of the ventral valve to a hard as a hydrostatic skeleton combined with the substrate, lack a pedicle; the larvae settle with contractions of the pedicle muscle and coelo- the posterior end expanded along the sub- mic pressures in the body. strate and secreting the ventral valve, which In intertidal environments, the lingulide is cemented to the substrate. This ventral retracts into the burrow during low tide. It valve is variably calcified in Neocrania species follows the water level down and then moves and has a calcified, alveolate structure in upward again with the advancing tide Neoancistrocrania norfolki. (CHUANG, 1956, 1961; EMIG & others, As in lingulides, the strong adductor 1978). muscles of discinids and craniids close the shell, which is opened mainly or exclusively EPIFAUNAL PATTERN: DISCINIDAE by hydrostatic mechanisms with longitudi- AND CRANIIDAE nal and outer body muscles working against The other extant inarticulated brachio- the pressure of the coelomic fluid. The setae pods are epifaunal, attached either by a fixa- of the mantle edges of the discinids are as tion organ (discinids) or by cementation to highly specialized as those of lingulides. some hard substrate (craniids). The ventral They have tactile sensitivity, resulting in a valve is always oriented toward the substra- protective closure of the shell, which is ac- tum, a feature that the discinids and craniids companied by the contraction of the pedicle share with the articulated thecideidines, drawing the shell near the substratum. The which is related to the orientation of the craniids have no setae. 480 Brachiopoda

0.5 cm

1

2

FIG. 411. 1, Live Discradisca strigata in pumping position, the anterior setae interlocked to form a functional siphon; arrows indicate in- and outcurrent directions (adapted from LaBarbera, 1985); 2, faunal distribution on a rocky substrate. All D. strigata are numbered; number 6 bears a D. strigata (number 7) and number 22 bears a barnacle; A, anemone; C, solitary coral; G, gastropod (LaBarbera, 1985).

The shells of discinids and craniids gape riorly and exits through two, posterolateral, quite widely at the anterior edge and more exhalant gapes (Table 35; PAINE, 1962b; narrowly at the posterior margin. A copious, LABARBERA, 1985; EMIG, 1992). Another dis- median, inhalant flow enters the shell ante- position for craniids is that two inhalant cur- Ecology of Inarticulated Brachiopods 481 rents flow in at the anterolateral margins, LIFE SPAN while one main exhalant current flows out at the anteromedian edge; additional exhalant The longevity of lingulides based on the currents occur at the posterolateral margins length of the shell is a matter of conjecture. (CHUANG, 1974). The life spans of Lingula anatina and L. In discinids the densely packed setae of reevii have been recently estimated theoreti- the anterior mantle margin function as an cally from five to eight years, while Glottidia incurrent siphon (Fig. 411.1). These anterior pyramidata is said to live from 14 months to setae can be nearly three times as long as the less than two years (Fig. 412; MORSE, 1902; diameter of the shell, while the length of the PAINE, 1963; CULTER, 1979). Shell growth in setae diminishes rapidly toward the posterior Lingula anatina and L. reevii decreases lin- end (MORSE, 1902). Discinids orient the lo- early with increasing size (WORCESTER, 1969; phophore relative to the current (LABARBERA, MAHAJAN & JOSHI, 1983). L. anatina attains 1985); but Pelagodiscus, because of the na- a length of 25.6, 36.8, and 47.6 mm at the ture of its attachment, probably undergoes a age of one, two, and three years respectively small degree of reorientation against the cur- (Fig. 412); consequently the theoretical life rent. In the cemented craniids the orienta- span appears to be six to seven years. Two tion may depend on the larval settlement previous shell growth curves have been estab- under the influence of the prevailing bottom lished for Lingula reevii in Hawaii (WORCES- current, with adjustments at that stage so TER, 1969) and for Lingula anatina in that the anterior region faces the local flow Australia (Fig. 412; KENCHINGTON & HAM- direction. MOND, 1978). Growth in a population in a Discradisca strigata has a characteristic restricted area, however, is directly related to behavior pattern (LABARBERA, 1985). At ir- such local environmental factors as water regular intervals or when disturbed, the characteristics, disturbances, nature of the valves nearly close, and the dorsal valve ro- substrate, and nutrients. These time-depen- tates clockwise and counterclockwise dent variations can affect the metabolism of through an arc of 60 to 120°. This move- the animal and consequently retard or favor ment rubs the lateral setae of the dorsal valve growth, although the shell grows continu- over and past the ventral setae. The setal si- ously throughout its life. Consequently, in- phon is disturbed by this movement but re- dividuals of equal shell length may differ in mains potent. When the dorsal valve returns age, sexual maturity, and longevity (CHUANG, to its normal alignment with the ventral 1961; PAINE, 1963; WORCESTER, 1969; C. valve, their margins clamp together tightly EMIG, personal unpublished data, 1983). and both valves rotate as a unit through an There are few data on the life spans of arc of 60 to 150°. On returning to a resting other inarticulated brachiopods. Populations position, the margins of the valves remain of Discradisca strigata (LABARBERA, 1985) clamped tightly to the substrate, but within take more than 10 years to become stabi- several minutes at most the shell returns to a lized. The three to six growth rings in the position slightly elevated above the substrate shell of Pelagodiscus atlanticus may be inter- and the valves slowly reopen. The subcentral preted as evidence of a life span of three to foramen of discinids affords greater protec- six years. However, shells from the continen- tion for the pedicle than the posterior open- tal slope have a greater length and a narrower ing of articulated brachiopods and ensures relative width and a smoother, less crenu- that the entire shell margin, including re- lated periostracum than those from the abys- gions adjacent to the pedicle, sweeps through sal plain (ZEZINA, 1981). These differences a sizeable arc when the animal rotates, thus seem to be the results of such environmental inhibiting growth of epifauna at a greater factors as temperature variations (2.65 to distance from the shell. 3.07°C on the slope, 2.2 to 2.35°C in the 482 Brachiopoda

50

45

40

35

30

25

20 shell length (mm)

15

10

5

0 0 123456789 years

FIG. 412. Growth curves of various lingulide species; ■, Lingula anatina (data from Mahajan & Joshi, 1983); ●, Lin- gula anatina (data from Kenchington & Hammond, 1978; , Lingula reevii (data from Worcester, 1969); , Glottidia pyramidata (data from Culter, 1979) (new). plain) and food supply. Presumably these or mangrove tree roots. The grain-size frac- factors control the growth rate more effec- tion that is transported by saltation (about tively on the slope than on the abyssal plain 90 to 220 µm) and generally associated with (ZEZINA, 1981). Neocrania anomala lived for the traction-load fraction (> 600 µm) deter- 14 months in aquaria at normal laboratory mines lingulide distribution. Where the trac- light and without changing the water tion fraction (about 220 to 600 µm) or the (JOUBIN, 1886). suspension fractions (< 90 µm) increase in the sediment relative to the saltation frac- ECOLOGY tion, lingulide density decreases rapidly. The ABIOTIC FACTORS distribution of lingulides in deeper waters sometimes depends on the presence of Qua- Substrates ternary littoral sands, as in New Caledonia. Soft substrates: Lingulidae.—Lingulides From the few available data, the organic con- live in compact and stable sediments under tent of substrates containing Lingula is rather the influence of moderate, near-bottom cur- low (one to four percent) (EMIG & LELOEUFF, rents (PAINE, 1970; EMIG, 1984a). The two 1978; BARON, CLAVIER, & THOMASSIN, preferred substrates are either well-sorted, 1993). Nevertheless, other ecological fea- fine- to very fine-grained sand and clayey tures affect the distribution and may be even sand (in which the 90 to 250 µm fraction more important. comprises more than 50 to 60 percent) and Hard substrates: Discinidae and Cra- coarse sand grains in a fine-grained or very niidae.—Discinids attached to various rocky fine-grained sandy matrix. The sediment can surfaces and to mollusc fragments occur sin- be further stabilized by marine phanerogams gly or in clusters of many individuals, for ex- Ecology of Inarticulated Brachiopods 483 ample, Discinisca lamellosa, D. laevis, and cause all are typically quite intolerant of Discradisca strigata. The last species forms lower salinity, none is adapted to brackish- clusters of more than 12 individuals sepa- water or freshwater conditions. Accordingly rated by less than 2 mm, while solitary indi- lingulides actually live in biotopes in normal- viduals are uncommon (LABARBERA, 1985). marine salinities but are capable of osmotic Pelagodiscus atlanticus is found attached to response to stresses of strong salinity varia- rocks ranging in size from pebbles to boul- tions, particularly at low tide in the intertidal ders (FOSTER, 1974) and is sometimes found zone when freshwater input occurs (HAM- on bivalve shells (Vesicomya, Bathyarca), MEN & LUM, 1977). The salinity range of the brachiopod shells (COOPER, 1975), scaph- populations of a species depends on the ge- opod shells, whale bones, and manganese ography of its habitat. Yet populations can nodules (ZEZINA, 1981). P. atlanticus occurs survive a greater range of salinity than that in deep-sea areas where fine-grained sub- occurring in its normal environmental con- strates accumulate slowly; both factors ap- ditions. The presence in a deltaic environ- pear to limit the distribution of this species ment does not, therefore, imply that the (ZEZINA, 1961). lingulides constantly live under reduced or Neocrania species show a wide depth tol- highly fluctuating salinities (EMIG, 1981a, erance and a preference for flat, hard surfaces 1986). Mean salinities during annual varia- on which they generally grow in clusters. In tions as low as 20‰ are exceptionally re- shallow water Neocrania occurs attached to ported in lingulide environments. Actually the undersides or sheltered sides of rocky lingulides are not tolerant of extremely low surfaces, including areas of bare rock, sub- salinity except for brief periods, generally less strates coated with coralline algae, and sub- than 24 hours. The lowest limit is about 16 marine caves. In deeper water, individuals to 18‰, which is not exceptional in com- occur on rocks, ranging from pebble to boul- parison to bivalve molluscs (HAMMEN & der size, shells, hard skeletons of other inver- LUM, 1977). tebrates, various hard fragments, and, more Temperature rarely, on other brachiopod shells (ROWELL, 1960; BERNARD, 1972; FOSTER, 1974; BRUN- Previously regarded as the limiting factor TON & CURRY, 1979; LOGAN, 1979; LEE, of the latitudinal extension of the lingulides, 1987). Neocrania larvae settle on hard sub- the range of temperature tolerance is highly strates where the sedimentation rate is very variable among populations; and a popula- low and often colonize substrates that are tion of a given area is generally unable to swept by strong currents reaching 3 to 5 km/ survive temperature variations, especially low h (ROWELL, 1960; FOSTER, 1974; LEE, 1987), temperatures, larger than those occurring in but they do not occur in more strongly cur- natural conditions. The salinity or tempera- rent-swept environments more frequently ture range under which an indigenous popu- than other brachiopods. The external shape lation normally lives can be lethal for an- and height of the craniid shell vary greatly in other population adapted to a different range response to the contours of the substrate to of conditions. Lingula anatina is a good ex- which they are attached (FOSTER, 1974; LO- ample as is illustrated by comparing the re- GAN, 1979; LEE, 1987). action of three populations that are widely Craniscus has been recorded from Japan dispersed (Table 37; EMIG, 1986, 1988). on various kinds of substrates from sandy Neither of the populations from northern Ja- mud to rocky bottoms. pan and New Caledonia could survive at sa- Salinity linities higher than 40 to 50‰. In northern Japan (EMIG, 1983a) and China (LEROY, At present, all inarticulated brachiopods 1936) the temperatures remain below 5°C live in seawater of normal salinity; and, be- for three months and below 11°C for more 484 Brachiopoda

TABLE 37. Annual variations of temperature, presence of hemerythrin within the coelom- salinity, and the bathymetric range of Lingula ocytes (YATSU, 1902b; HAMMEN, HANLON, & anatina in three locations (Emig, 1988). LUM, 1962; WORCESTER, 1969). Hemeryth- rin, however, seems to be used as a store un- Temperature Salinity Depth (°C) (g/l ) (m) der anoxic conditions or during cessation of respiration, such as may occur intertidally Persian Gulf 15–40 55–60 6–16 when the burrow is exposed and is part of New Caledonia* 18–30 15–25 intertidal (to 67) Northern Japan 1–22 28–30 5–18 the oxygen-transporting function in lingulides. Data on the rates of oxygen con- ° *lethal conditions at <15 C and salinity of >40 g/l. sumption are available only for lingulides but are difficult to compare because they are than six months, while populations from based on either total-animal wet weight New Caledonia that experience experimen- (HAMMEN, HANLON, & LUM, 1962) or on ° tal temperatures below 15 to 17 C undergo dry mass of tissue (SHUMWAY, 1982). Lingula a lethal, irreversible retraction of the mantle. reevii and Glottidia pyramidata have higher The onset of breeding and the length of rates of oxygen consumption than the articu- the spawning season of lingulides depend lated Terebratulina septentrionalis by a factor mainly on water temperature and latitudinal of two to nine, and the activity of metaboli- and seasonal effects. They vary from a 1.5- cally important enzymes, such as succinate month period in midsummer in temperate dehydrogenase, is up to 20 times higher waters (northern Japan, Virginia) and a five- (HAMMEN & LUM, 1977; HAMMOND, 1983). to nine-month period between late spring On the other hand, the oxygen consumption and late autumn in warm temperate waters rate of Lingula anatina is about 2.5 times to year-round breeding in tropical waters lower than in three articulated species (southern Florida, Singapore, Burma, and (SHUMWAY, 1982). India) if temperatures do not drop below 26 The redox layer, which often occurs some ° to 27 C. 2 to 5 cm below the sediment-water inter- There are no data on temperature require- face, does not signify a low oxygen concen- ments of the discinids except that Pelago- tration in the surrounding water mass, even discus is more abundant in the deep sea at in the burrow. Such anaerobic conditions as ° temperatures below 3.5 C. red tides can be responsible for a mass mor- Neocrania species tolerate a wide annual tality. Although Glottidia pyramidata was range of temperature related to their geo- one of the five species of 22 species surviving graphic and bathymetric distribution, from such events that temporarily lowered the ° ° -2 to 1.5 C for N. lecointei (FOSTER, 1974), mean density of the population from 42 to ° 14 to 21 C for N. huttoni (LEE, 1987), and 13 individuals per square meter, two years ° about 26 to 28 C for species living in equa- later this density had risen to 1,332 individu- torial waters. N. anomala, which is distrib- als per square meter (SIMON & DAUER, ° ° uted between 30 to 60 N in the Atlantic 1977). Individuals of Glottidia are probably Ocean and Mediterranean Sea, has a wide able to resist short-term anoxic events be- temperature tolerance. The almost complete cause they bear mantle papillae over the sec- absence of calcite in the pedicle valve of sev- ondary mantle canals in the pallial cavity. eral species of Neocrania does not represent The papillae allow an increase of the respira- an adaptation to very cold water (FOSTER, tory and nutritional exchanges. On the other 1974). The temperature range of the bio- hand, the volume of the lophophoral cavity ° topes of Craniscus is from 2 to 18 C. in Glottidia is less than that of Lingula. In the Oxygen same way Lingula anatina is more resistant to stress from loss of oxygen than bivalves col- Lingulides are able to survive temporarily lected from the same locality (ROBERTSON, in poorly oxygenated waters because of the 1989). Ecology of Inarticulated Brachiopods 485

02468101214161820 0246810 02468 02468 species 0

shelf break

20 lingulides craniids inarticulated brachiopods discinids

40

477

60

80

1,000

1,200

5,530 5,530 2,342 (7,600) (7,600)

FIG. 413. Bathymetric distribution of the living inarticulated species; numbers at bottom indicate deepest recorded living specimens; those in parentheses indicated deepest recorded empty shells (new).

Depth and density depth. More than 40 percent of inarticulated brachiopods, mainly lingulide and discinid Many living inarticulated species extend species, occur between 0 and 60 m depth; through a remarkable depth range from lit- and more than 40 percent of the craniids toral waters into the bathyal zone (ranging occur between 20 to 420 m. from the shelf break, generally about 100 m, The optimum environment for living Lin- to 3,000 m) down to about 500 m on the gula and Glottidia species is not intertidal, slope (Fig. 413). Only Pelagodiscus atlanticus although 11 of the 12 species of lingulides occurs at abyssal depths, i.e., in the zone have been recorded in the intertidal zone ranging from 3,000 to 6,000 m. Inartic- (PAINE, 1970; EMIG, 1984a) and in the ulated brachiopods seem not to have mi- infralittoral zone from 1 to 2 m to about 20 grated into deeper water in the course of m. The maximum recorded density of Lin- time and cannot be used as indicators of gula reevii is 500 individuals per square 486 Brachiopoda meter (WORCESTER, 1969); that of L. anatina Other Factors is 864 individuals per square meter (KEN- CHINGTON & HAMMOND, 1978). Glottidia As suspension feeders, brachiopods re- pyramidata reaches concentrations of more quire good circulation of the water. Seawater than 8,000 individuals per square meter in constituents also play a role in the ecological Florida (CULTER, 1979), and G. albida shows requirements. Some are used for formation a density peak of more than 500 individuals of the shell and their rate of assimilation may per square meter in depths of 22 to 47 m off have a direct influence on growth of the the coast of California (JONES & BARNARD, shell. Calcium ions, which are taken up from 1963). the seawater primarily by the lophophore, Pelagodiscus atlanticus, one of the deepest- move through the coelomic system into the water brachiopods, has been recorded mantle and are eventually deposited in the throughout the bathyal and abyssal zones inner layer of the shell. Yet the major source with one-third of the occurrences being at of inorganic phosphate for shell formation in depths of more than 4,000 m (ZEZINA, 1961) Glottidia pyramidata is likely to be food and and only a few of the records of its occur- not seawater (PAN & WATABE, 1988a). rence being from less than 1,000 m (ZEZINA, On the Florida coast, Glottidia pyramid- 1975). Its density may reach up to 480 indi- ata, together with the lancelet Branchiostoma viduals per square meter at the foot of sea- caribbaeum, are sensitive to deterioration of mounts and up to 76 individuals per square water quality and, thus, are used as indicator meter at 1,500 to 2,000 m in Antarctic wa- organisms of unspoiled areas and uncon- ters (ZEZINA, 1961). On the marginal ridge taminated waters in determining suitability of the Kurile-Kamchatka trench, however, a for fishing. eutrophic area with a rich food supply and Taphonomy rather active currents, the density of 12 indi- viduals per square meter is comparable to Recent ecological statements on tapho- that in the tropical oligotrophic parts of the nomic conditions of living lingulides (EMIG, ocean (ZEZINA, 1981). The other species of 1986, 1990) have been corroborated by re- discinids are mainly restricted to the conti- interpretations of fossil beds (Fig. 414). The nental shelves. Four of the 12 species of natural death of the lingulides leads to the discinids have been recorded in the intertidal extrusion of the animal from its burrow zone, although all of them are more abun- (WORCESTER, 1969; EMIG, 1986). The valves dant below the low-tide level or subtidally become separated, and the organic matrix (Fig. 413). degrades rapidly due to hydrolysis, microor- The craniids extend from shallow waters ganisms, and mechanical abrasion. The thin, to the bathyal zone and appear as a deeper- fragile, chitinophosphatic valves are reduced water group among the inarticulated brach- to unrecognizable fragments, the deteriora- iopods. Densities of N. anomala up to 500 tion occurring from the margins to the cen- individuals per square meter have been re- tral portion of the valve; and in general after corded on small, flat, hard surfaces at various two or three weeks the valves have com- depths between 10 and 200 m. N. lecointei pletely disappeared from the sediment has been found alive only on the seaward (EMIG, 1983a, 1990). This explains why only edge of the continental shelf in the Ross Sea, a catastrophic event, occurring over some which belongs presently to the bathyal zone, days, is the most significant source of mortal- between approximately 450 and 650 m, ity with respect to preservation of the shell where it is the dominant brachiopod with up and ultimate fossilization because there is to 46 individuals per square meter (FOSTER, little potential for fossilization in normal 1974). environments (EMIG, 1986). Consequently, Ecology of Inarticulated Brachiopods 487 fossil lingulides are not indicators of their biotopes but of drastic environmental temperature changes that led to their burial. salinity Fossilization can occur either in situ in life sea level position, for example, in conditions of rapid substratum level temperature decrease, salinity increase, des- iccation, emersion of the substratum or drop muddy sedimentation of sea level, or very fine sedimentation; or it 1 in situ (within the burrow) can occur as flat-lying disarticulated valves, for example, after prolonged reduction of salinity, coarse-grained sedimentation, and salinity storms (EMIG, 1986). Data obtained for liv- ing species obviously apply to the interpreta- tion of fossils (EMIG, 1986). Nevertheless survivorship under abnormal conditions can vary according to the geographical popula- tion and depends also on the synergy of the coarse sedimentation applicable environmental factors on a given population. storms When salinity increases to 40 to 50‰, the death of populations occurs in burrows in a 2 flat-lying valves few days. Osmotic pressure empties the ani- mal of its coelomic fluid, and the pedicle FIG. 414. Diagram summarizing the effects of abiotic factors that may induce lingulide fossilization (new). becomes detached from the shell. When the salinity decreases below 16 to 18‰, death occurs in one day to several weeks and quick- 1963; SOOTA & REDDY, 1976; EMIG, 1986, ens with lowering salinity, although the sa- and personal observations, 1983). Neverthe- linity of interstitial water remains high for less from experimental results the duration of several days. Individuals leave their burrows survivorship to low salinity is variable among as their bodies swell under osmotic pressure, species. At a salinity of 15‰, Lingula ana- and pedicles become limp or detached. The tina in Queensland (Australia) resists longer putrefaction of the soft body causes separa- than Lingula reevii in Hawaii, while Glottidia tion of the valves, which are then spread over pyramidata in Florida has a greater survivor- the sediment surface. Shells rarely float, but ship than populations of Lingula. it has been reported (EMIG, 1981b). At a sa- During an exceptional storm often associ- linity of 18‰, the initial body weight in- ated with heavy rains, the sediment is creases by 3.3 percent in three hours; at 5‰, churned up, and the lingulides are washed it increases by 3.8 percent in one hour. onto the shoreline and may form shell Weight then remains constant for about two masses up to 75 cm high (RAMAMOORTHI, hours followed by another weight increase VENKATARAMANUJAM, & SRIKRISHNADHAS, that is lethal (HAMMEN & LUM, 1977). Re- 1973; HAMMOND, 1983; EMIG, 1986). duced salinities in rapid transition are toler- When the sea level drops through tec- ated, for example, during tidal cycles in es- tonism, regression, or high sedimentation, tuarine or deltaic environments where the the animal retreats with the water level un- salinity can drop to less than 10‰. Several til it reaches the bottom of its burrow where observations have reported high mortality death occurs in about three days. after heavy rains of two to three days’ dura- Experiments on Lingula anatina in New tion causing nearby rivers to flood (PAINE, Caledonia with decreasing temperatures 488 Brachiopoda show that below 15 to 17°C (the lowest tem- welling system. Such deposits totally domi- perature in natural conditions is 18 to 19°C) nate the littoral sediment (HILLER, 1993). A individuals go down to and remain at the correspondence is suggested with the Esto- bottom of their burrows. At 6 to 10°C an ir- nian Lower Ordovician obolid conglomer- reversible retraction of the mantle occurs ates, which are likely to have formed under over several millimeters from the shell mar- similar conditions of upwelling. gins leading to death within one to three weeks because the lingulides are unable to BIOTIC FACTORS form their pseudosiphons, and, conse- quently, the pallial water streams are highly Nutritional sources perturbed (C. EMIG, personal unpublished Sources of nutrition are known for only a data, 1983). Mantle regression has been ob- few lingulide species. The type and abun- served in both inarticulated and articulated dance of ingested particles as well as the im- brachiopods, but a factor specifically portance of direct absorption of nutrients responsible for this regression is identified depend on such factors as season, depth, and for the first time herein. geographic area. Analyses of gut contents of When the temperature drops below 10°C Lingula reevii from Hawaii (EMIG, 1981b) in Florida, Glottidia pyramidata does not re- show the presence of two types of food: a spond to any stimuli, although slow warm- vegetal fraction, mainly phytoplanktonic and ing after three days at low temperature pro- consisting of diatoms, peridinians, and fila- duced signs of activity at 12°C (PAINE, mentous algae, and an animal fraction, 1963). mainly from the superficial meiobenthos and Muddy sediment with more than 35 to 40 macrobenthos, i.e., foraminifers, rotifers, percent of very fine fraction (< 50 µm) de- polychaetes, oligochaetes, and copepods. posited over original sandy bottoms leads to Both fractions are mixed with a constant the death of lingulides within their burrows amount of sedimentary particles of 2 to 3 µm in several weeks. A lingulide can maintain and various organic detritus (e.g., spicules only sporadically its normal position before and spines). Glottidia pyramidata ingests par- collapsing into the sandy layer, and this gen- ticles smaller than 125 µm in diameter, in- erally leads to death by debilitation. This cluding sand grains and various vegetal and observation is of paleoecological importance. animal matter, Coscinidiscus, gastropod ve- When lingulide valves occur at the bottom of ligers, nauplii, and even Glottidia eggs a shale overlying a sandstone, the sandstone (PAINE, 1963). Food particles from the sedi- unit is the normal substrate of the lingulides ment-water interface may be readily resus- that are sometimes fossilized within their pended by tidal or bottom currents or waves, burrows. The shale cannot be interpreted as by arm shaking of ophiurians, or by holothu- the normal substrate for lingulides but as a rians. deposit of muddy sedimentation that was Direct absorption of dissolved nutrients is responsible for the death of the lingulide known to occur in the lophophorates. The population. Conversely, coarse sedimenta- lophophore in lingulides (STORCH & tion (> 0.5 mm) leads to the emerging of the WELSCH, 1976) appears to be able to absorb lingulides at the sediment surface and finally directly dissolved organic matter from seawa- lying on the surface. ter. There is also evidence that digestion oc- The shallow-water species Discinisca curs in the lophophore, attested to by the tenuis occurs intertidally at a few localities. It presence within the tentacles of alkaline is known in the Walvis Bay area (Namibia) phosphatase and three esterases (STORCH & where large deposits formed by huge num- WELSCH, 1976). Like the phoronids, bers of shells are washed up onto the beach. lingulides are able to live in aquaria for some Its occurrence along the Namibian coast is weeks without having the water changed. linked to the existence of the Benguela up- Glottidia pyramidata can be maintained at Ecology of Inarticulated Brachiopods 489 least three months under starvation condi- tide (VARGAS, 1988); stomach contents of tions without apparent loss of vitality (PAINE, the willet Catoptrophorus semipalmatus but 1963). more frequently the short-billed dowitcher The body weight of Lingula anatina var- Limnodromus griseus revealed that G. aude- ies from 0.13 g for a shell length of 1.35 cm barti is an important food item. Catoptro- to 5.19 g for a shell length of 4.25 cm. (The phorus semipalmatus and the fish Symphurus mean value is 2.24 g for a length of 3.19 cm; plagiusa are known predators of Glottidia n=346; KAWAGUTI, 1943.) The body weight, pyramidata, which is their main source of like the shell height, increases more rapidly food (PAINE, 1963). People also eat Lingula than the shell length. The weight:length ra- anatina and L. rostrum on almost all the tio of Glottidia pyramidata changes at a western Pacific islands from Japan to New length of approximately 8 mm correspond- Caledonia. ing to the development of gonads (PAINE, 1963). In Lingula this development occurs at Parasites a shell length of 1.5 to 2 cm. Unencysted metacercariae of trematodes of the subfamily Gymnophallinae (usually Predation one to three in an individual) have been sea- Lingulides are eaten by such crustaceans as sonally recorded around the nephrostomes hermit, stone, and portunid crabs, crango- and in the gonads of Glottidia pyramidata, nids, stomatopods, shrimps, and amphipods mainly at the end of summer and in autumn. (PAINE, 1963; WORCESTER, 1969; CULTER, The infestations can reach 68 percent of the 1979; EMIG & VARGAS, 1990). The asteroid population. These parasites can reduce or Luidia clathrata is an important predator of destroy the gonads and have a secondary in- Glottidia pyramidata. Forty-three percent of fluence on the digestive glands and mantle the Luidia collected contained Glottidia canals (PAINE, 1962a, 1963). Adult parasites shells with little selectivity for size for shells are likely to occur in avian predators of G. less than 1 cm long, suggesting that larger pyramidata. The occurrence of two species of individuals may withdraw too deeply into poecilostomatoid copepods, Parostrincola the sediment to be preyed upon. Other echi- lingulae and Panjakus platygyrae, associated noderms are also reported as predators, such with Lingula anatina has been reported from as the ophiuroid Amphipholis germinata and Hong Kong (HULMES & BOXSHALL, 1988). the echinoid Encope stokessi (EMIG & VARGAS, Zooxanthellae are abundant within the di- 1990). Gastropods (mainly naticids and gestive gland of Lingula (KIRTISINGHE, 1949), muricids) are only occasional predators of and monocystid protozoa have been reported lingulides, but bored valves can represent up in Neocrania. to 14 percent of the valves recovered from FAUNAL RELATIONSHIPS the sediment (PAINE, 1963). Dead shells of craniids are sometimes drilled by gastropods Communities (LEE, 1987). Lingula parva has been recorded during Soft-substrate communities.—By their gen- the dry period (March to September 1953) eral characteristics, lingulides are nearly along the Sierra Leone and Nigerian coasts in stable in their evolutionary state. They the stomachs of several demersal fishes present all the features of a dominant group (LONGHURST, 1958; ONYIA, 1973). Several within a community (EMIG, 1989a): low tens of Glottidia pyramidata shells have been growth rate, uniformity of shape, larger size recorded in stomachs of sturgeons and vari- than the other members of the community, ous rays along the Florida coast. The mud long life span, low recruitment potential, flats inhabited by Glottidia audebarti are vis- generally just higher than the population ited seasonally by migratory birds, and at replacement (K-demography), and long geo- least 13 species were observed foraging at low logical range. Such characteristics allow high 490 Brachiopoda biomass to develop related to the available (DOLAH & others, 1984). Near Charleston energy and result in an excellent ability to Harbor (Florida, USA) it is present in coarse integrate and conserve energy. Such a domi- to fine sands from 8 to 17 m (DOLAH, nant group generally shows plesiomorphic CALDER, & KNOTT, 1983) with the highest characters compared to other taxa. density being at 17 m. In Tampa Bay Few lingulide communities have been (Florida, USA) the reestablishment of a studied. Data on the macrobenthic fauna are benthic community following natural given in Tables 38 and 39. Lingula anatina defaunation by red tide has been studied in has been investigated in the Mutsu Bay an intertidal sand flat (SIMON & DAUER, (northern Japan) in fine sands and muddy 1977). sands from 4 to 18 m depth (TSUCHIYA & The associated fauna of other locations is EMIG, 1983); on the west coast of Korea in a briefly listed here. On the western Korean tidal flat of silty sands from -2.5 to 2.3 m coast Lingula anatina occurs in sand to sandy (AN & KOH, 1992) where the number of mud flats with many other such endobiont species collected monthly varies from 28 to species as polychaetes, crabs, and molluscs, 41; in Taiwan in a tidal flat of fine, sandy which are dominant quantitatively (FREY & mud (DÖRJES, 1978); in Phuket Island others, 1987). In a New Hebridian man- (Thailand) in front of a mangrove in an in- grove community L. anatina occurs seaward tertidal, large, bay-shaped, fine-sand flat of the Rhizophora zone dominated by gastro- dominated by molluscs, mainly the gastro- pods and crabs (MARSHALL & MEDWAY, pod Cerithidea cingulata, where the other 1976). On the western African coast L. parva most abundant animals are the fiddler crab occurs in the Venus community, particularly Uca lactea and the sipunculid Phascolosoma at the Venus-Amphioplus transition (LONG- arcuatum (FRITH, TANTANASIRIWONG, & HURST, 1958). In the Ebrié Lagoon (Ivory BHATIA, 1976). In New Caledonia in associa- Coast) L. parva occurs in a shallow, sandy tion with the seaweed Halodule on coarse substrate in the Corbula trigona community sands the macrofauna is dominated respec- in which the main species are 12 polychaetes, tively by Lingula anatina, molluscs (mainly 9 gastropods, 14 bivalves (dominant), and the bivalve Gafrarium tumidum and a gastro- 10 crustaceans (ZABI, 1984). In Ambon, L. pod Cerithium sp.), and polychaetes (mainly rostrum occurs midlittorally seaward of a Caulleriella sp.) (BARON, CLAVIER, & THO- mangrove stand and on a sandy beach lo- MASSIN, 1993). Glottidia audebarti recorded cated between the ocypodid zone and the in Costa Rica (VARGAS, 1988; EMIG & VAR- clypeasterid zone (EMIG & CALS, 1979). In a GAS, 1990) in mud flats exposed only at a benthic survey on the eastern coast of India tide level below 0.1 m has an associated (BHAVANARAYANA, 1975) a Lingula-Solen zone macrofauna composed mainly of deposit was reported, almost exclusively populated feeders; the meiofauna comprises 88 percent by both taxa in considerable numbers. Off nematodes, 6 percent foraminifers, and 3 the Californian coast, Glottidia albida occurs percent ostracodes. Glottidia pyramidata oc- at high density in the Amphioplus commu- curs in Sapelo Island (Georgia, USA) in the nity inhabiting a compact, fine, sandy sub- Moira-atrops community located between 10 strate although it has also been recorded in and 13 m depth in coarse, relict sand domi- several other communities, including the nated by polychaetes followed in importance Listrolobus, Amphioda, Nothria, and Tellina by crustaceans, but the fauna shows a gener- communities (JONES & BARNARD, 1963). In ally low density (DÖRJES, 1977). In Winyah Mission Bay (San Diego, California), G. Bay (South Carolina, USA) it occurs in me- albida occurs with a macrofauna dominated dium- to fine-grained sands from 6 to 11 m by 65 percent polychaetes, 15 percent Ecology of Inarticulated Brachiopods 491 n ur. & IMON S 30–46 17–43 26 30 85 195 79–93 27–55 37–193 54–155 83 9 , 1977; AUER , 1958; & D IMON ONGHURST S L 8 8 , 1976; , 1976; HATIA HATIA L. anatina L. anatina L. anatina L. anatina L. anatina L. parva G. audebarti G. pyramidata G. pyramidata G. pyramidata G. pyramidata , & B , & B ANTANASIRIWONG ANTANASIRIWONG , T , T RITH RITH F F 7 7 , 1990; , 1990; ARGAS ARGAS & V & & V & MIG MIG E E 6 6 , 1978; , 1978; ÖRJES ÖRJES D D 5 5 , 1977; , 1977; ÖRJES ÖRJES D D 4 4 , 1983; , 1983; mean data from a different area of Florida. area mean data from a different NOTT NOTT 12 , & K , & K ALDER ALDER , C , C OLAH OLAH D D 3 3 , 1993; , 1993; , 1988. ARGAS total data from one area of Florida; of Florida; one area total data from V 11 HOMASSIN HOMASSIN 11 The results presented have been calculated from the data given by the cited authors (new). the data given by been calculated from have results presented The , & T , & , & T , & , 1988; , 1983; n% n%n%n%n% n n n % n%n% n%n% n MIG LAVIER LAVIER ARGAS 0273-520 15–298–178 39–59 3172762314312 9 30827723-- 39–47 5–12 3–14 12–27 18–34 3–9 4–7 8–21 16–29 1–6 0–3 3–15 0–8 0–4 0–1 0–7 0–3 383830–3815–31 45- 38–41 1931–88 38–56 15–1832 25 43–60 3–18 19 54 40–45 11–38 9–31 29 39 - 28 21–25 12–29 5–9 27 12 22 12–42 11–19 20–21 57 22–32 1–2 - 14 27 - 29 1–7 4 21–23 3 2–5 19 - 27 - 3–5 - 23 4 14 12–13 6–7 13–16 2 - - 7 19 - 10 8 - 12–14 10 12 195–9108–39 45–77580 1–49 35–150 46 3–12 12–7,000 14–99 5–230 340 2–20 4–27 1–6 5–185 27 1–26 0–14 285 0–80 0–12 23 0–10 0–9 55–150 20 0–6 455–1,195 2 1–66 79–7,100 15 1 15 1,255 1498-- 9308–2,378 22 27–64231–868 33–55(336) 7–37 15–32 70–424 103 130 - (5) 3–30 4–50 - 62 98–1,522 29 5–32 1–4 (2,478) 5–12 24–45 27 (38) 4–48 29 64–7,877 - 3–89 - 1–3 (2,181) 16 7 - (34) 29–47 5–19 158–207 - 67–53 - - - 39 - 2–48 - - 9 - 1,070–5,132 1–3 80–261 2–13 21 152 - - (1,501) 50–1,178 646–8,850 (23) 33 13 4–15 22–35 1,332 100 151 1 17 (6,496) 438–6,240 448 3,700–41,000 166 V , C , C 10 & E ARON ARON which lingulides occur. The results presented have been calculated from the data given by the cited authors (new). the data given been calculated from presented have The results which lingulides occur. 2 2 B B 8 2 2 , 1983; SUCHIYA T MIG 4 3 4 3 10 6,11 6,10 7 7 , 1992; , 1992; 4 & E 38. Richness and percentage of the species of the main taxonomic groups present in various communities which lingulides occ groups present 38. Richness and percentage of the species main taxonomic 39. Densities per square meter and percentage of the individuals of the main taxonomic groups present in various communities i present in various groups meter and percentage of the individuals main taxonomic per square 39. Densities 5 5 8,11 8,12 9 OH OH 1 1 9 10 , 1977; & K & K ABLE ABLE SUCHIYA N N AUER A A T D T 1 LocationJapan Korea Taiwan PolychaetesThailand Caledonia New Mollusks Crustaceans Echinoderms Others Lingulide species Total Western Africa Western Costa Rica Georgia S. Carolina S. Carolina Florida T 1 9 LocationJapan Korea Taiwan Thailand Polychaetes Mollusks Crustaceans Echinoderms Others Lingulides Total New Caledonia New Costa Rica S. Carolina S. Carolina Florida Florida 492 Brachiopoda molluscs, and 11 percent crustaceans, with into account when analyzing taphonomic mean density from 621 to 1,874 individuals factors to explain the poor, associated fauna per square meter (DEXTER, 1983). On the found in paleocommunities or when specu- coasts of Florida, Glottidia pyramidata is of- lating about diversity of the fossils. In fossil ten associated with the lancelet Bran- assemblages lingulides are often the only fos- chiostoma caribbaeum, polychaetes, cuma- sils found, indicating either that other kinds ceans, and amphipods; and its biomass, of organisms were not preserved or that the which has a mean value of 35 percent, can biocoenosis was oligotypical (EMIG, 1989a). reach up to 75 percent of the total biomass The occurrence of such a monospecific as- of the benthic invertebrates. G. pyramidata semblage of fossils requires an extensive occurs also in the biocoenosis of well-sorted, analysis of the environmental constraints and fine sands with the phoronid Phoronis of the characters of the occurring species to psammophila (PAINE, 1963; EMIG, 1983b). identify any patterns of the original commu- The associated fauna within a given com- nity. The oligotypical biocoenosis presents munity seems to play a minor role in lin- one or several of the following characteris- gulide distribution (EMIG, 1984a). Neverthe- tics: low-energy input resulting from the ef- less, when the density of Lingula increases fects of climatic factors, extreme harshness there is a small decrease in the total number due to edaphic factors reducing the physiol- of species with an increase of the total num- ogy of the individuals, or high daily or sea- ber of individuals; when the density of sonal variations of the edaphic and climatic Glottidia increases the total number of spe- factors. Thus the biocoenosis is characterized cies and individuals tends to increase. Com- by high dominance in faunal and environ- parisons of the distribution of the major mental features and develops conservatism groups (Table 38–39) with the increase of with highly reduced capacity for organisms density of lingulides show in western Korea to evolve. that polychaetes (number of species and in- Hard-substrate communities.—In the deep dividuals) and crustaceans (number of spe- parts of the slope in the Antarctic regions cies) tend to decrease, while echinoderms, (FOSTER, 1974), Pelagodiscus atlanticus is as- mainly suspension feeders, tend to increase sociated with a very meager fauna. In the or to appear; in the Mutsu Bay, polychaetes, lagoonal complex of Cananéia, Brazil the dominant group, molluscs, and crusta- (TOMMASI, 1970b), Discinisca sp. has been ceans tend to decrease; with the muddy frac- recorded at 6 and 8 m depth on a rocky-sand tion increasing with the depth there is a gen- bottom with the following macrofauna: eral decrease in the fauna. Near Charleston polychaetes respectively 600 and 60 indi- Harbor the number of individuals of mol- viduals per square meter (10 and 18 percent luscs and the number of individuals and spe- of the fauna), molluscs 1,170 and 30 (19 and cies of polychaetes tend to increase, while on 9 percent), decapods 380 and 150 (6 and 46 the southwestern coast of Florida opposite percent), amphipods 2,900 and 0 (47 per- variations of the densities have been ob- cent), others 1,080 and 70 (18 and 21 per- served over a year between polychaetes, crus- cent), and Discinisca 10 and 20 (0.2 and 6 taceans, and Glottidia. percent). In the Bahia Conceptión, Chile, Polychaetes are generally the dominant Discinisca lamellosa occurs in the intertidal group in numbers of individuals and species zone with a meager fauna of one cnidarian, followed by molluscs or crustaceans (see one nemertine, two molluscs, two polycha- Table 38–39). The presence of molluscs is etes, and one to three crustaceans (URIBE & not fundamentally related to the distribution LARRAIN, 1992). In Baja, California, Discra- of lingulides (BABIN & others, 1992). An- disca strigata lives under cobbles and small other important feature is the large number boulders, patchily distributed on an exten- of species and individuals of the associated sive sandy beach and extending down to the fauna (Table 38–39), which should be taken low-water mark (PAINE, 1962b) where it is Ecology of Inarticulated Brachiopods 493 associated with sponges, gastropods, and fluctuations in density are highly variable bivalves. The fractional area covered by epi- even within a restricted geographic area. fauna averaged 49 percent, and free space Episodic failure of recruitment observed in ranged from 32 to 74 percent on a rock area lingulide populations can be related to such from 21 to 116 cm2: D. strigata covered 2 to causes as protracted breeding season, bad 26 percent of the surface, bryozoans 5 to 38 environmental conditions for settlement, percent, serpulids 2.5 to 29 percent, spiror- food supply, and interactions with the sur- bids 0.01 to 7.3 percent, and sponges 0.5 to rounding fauna including predation. 8.2 percent. Some authors (PAINE, 1970; KENCHING- Neocrania anomala is recorded in shallow TON & HAMMOND, 1978) have raised the waters under rocky surfaces together with a question of unidentified factors affecting the sciaphilic fauna. It also occurs deeper in the absence of lingulides in apparently suitable sublittoral zone with a fauna dominated by sediments. Actually the distribution of sponges, cnidarians, spirorbid worms, and lingulides is restricted within the limits of bryozoans, and on the continental slope be- the biocoenosis in which a lingulide species low 100 m on smooth, fine-grained, hard is living, even if preferred substrates occur substrate to large rocks, particularly within beyond the limits of the community (EMIG, the community dominated by the brachio- 1984a, and personal unpublished data, pod Gryphus vitreus. N. anomala is also re- 1983). corded in Scottish lochs with hydroids, sponges, chitons, foraminifers, and molluscs Shell epibionts (CURRY, 1982) and in the Strait of Messina Epibionts preferentially settle on the hard between 80 and 200 m under conditions substrate provided by the brachiopod shell. where there are bottom currents where the According to the infaunal habit of lingulides, macrofauna is dominated by anthozoans almost all epibionts are restricted to the an- (eight species), bryozoans (31 species), anne- terior margins of the valves because only lids (14 species), molluscs (20 species), crus- these margins are accessible and are not dis- taceans (5 species), and echinoderms (2 spe- turbed during withdrawal into the burrow. cies) (DI GERONIMO & FREDJ, 1987). Cyanobacteria, however, frequently extend Neocrania huttoni forms part of a distinctive to the umbonal region along the margins. rocky substrate community with calcareous In one locality the following macro- algae, sponges, serpulids, ascidians, bivalves, organisms were recorded from 5,000 Lingula barnacles, and bryozoans including a variety shells (WORCESTER, 1969): 10 occurrences of of filter feeders (LEE, 1987). N. lecointei is algae, 14 anemone Aptasia, many bryozoans, associated with a varied fauna that includes 2 polychaetes, 6 barnacles, 1 amphipod and, corals, polychaetes, ophiuroids, bryozoans, on 16 percent of the shells, the limpet and ascidians (FOSTER, 1974). N. pourtalesi Cruciblum spinosum. From a large list of occurs not uncommonly throughout some epibionts (represented by two algal divisions communities of cryptic habitats of coral and six animal phyla) on the shells of Lingula reefs, where brachiopods and sponges are the anatina and L. reevii (HAMMOND, 1984), the dominant taxa (JACKSON, GOREAU, & HART- most commonly recorded taxa are cyano- MAN, 1971). Craniscus occurs in Japan with bacteria (frequency up to 30 percent), an associated fauna that comprises mainly polychaetes (frequency up to 45 percent), molluscs and two articulated brachiopod barnacles (frequency up to 20 percent), lim- species, Dallina and Terebratulina (HATAI, pets (frequency up to 16 percent), bryozoans 1940). (frequency up to 11 percent), and traces of the attachment of the egg cases of gastropods Population structure or the byssal threads of mussels (frequency Because the distribution of lingulides is up to 29 percent). In only ten percent of the controlled by environmental factors, annual infested Lingula were both valves affected. 494 Brachiopoda

Algae, specifically Enteromorpha sp., estab- Competitive interactions lished itself only on those valves that had regeneration scars (PAINE, 1963). The hy- Mechanisms of competitive interaction droid Campanulatiidae Clytia can occur on are likely to be characteristic of the discinids up to 20 percent of lingulide individuals and may have been important in ensuring with a shell length exceeding 1.4 mm. the success of the living genera since their An unidentified leptocean bivalve (per- earliest known occurrence in the Triassic haps Euciroa) is found byssally attached to (ROWELL, 1961). The only work that has the shell of Lingula anatina (SAVAZZI, 1991) addressed the competitive abilities of in densities of up to nine individuals per inarticulated brachiopods deals with shell. The posterior region of the bivalve Discradisca strigata, which invariably wins shell is oriented upward, located at the ante- competitive interactions for space with other rior margin near the exhalant currents of the sessile epifauna (LABARBERA, 1985). One brachiopod. The bivalve progressively mi- such competitive interaction is metamor- grates upward as is shown by a trail of byssal phosis on the surface of bryozoan colonies filaments left along their paths to compen- facilitated by a reversal of the flow patterns sate for growth of the Lingula shell. The bi- and the possession of a functional anterior valve feeds on feces of L. anatina. Lingulides siphon that allows juveniles to draw water were found to carry gastropods near the an- from above the bryozoan’s lophophores, so terior margin. Egg capsules of gastropods that mature individuals eventually usurp the occurred seasonally on the anterior margins space occupied by the colony. Another inter- of the valves of lingulides; up to 45 Nassarius action is maintenance of a pool of particle- and up to 5 Olivella egg capsules were found depleted water around most of the shell of on a single individual of Glottidia pyra- larger juveniles and adults, which probably midata. inhibits encroachment by bryozoans and Epibionts like the worm Polydora or the sponges. In addition, abrasion of underlying mollusc Brachiodontes may benefit from calcareous epifauna by the harder phosphatic lingulide inhalant currents, but their pres- shell occurs, which erodes these faunal ele- ence can have detrimental effects by causing ments to the level of the substrate. Numer- distortion of the shell of the host (PAINE, ous eroded epizoans occur under the ventral 1963; HAMMOND, 1984). The number of valves of Discradisca although no abrasion of worms on a valve varies from one to six, with the valves themselves was seen. The edge of a typical number of three or four while the these valves probably abrades neighboring number of small Brachiodontes may be as organisms during rotation of the shell even many as five. of juveniles, and it is probably made more Among the craniids, Neocrania shells fre- effective by the simultaneous sweep of lateral quently bear encrusting organisms including setae that mechanically damage the tissues of bryozoans, serpulids, barnacles, calcareous surrounding sponges and bryozoans. algae, and sponges. Most valves carry more Three of these mechanisms are not avail- than one epibiont, and the percentage of the able to articulated brachiopods, and the cover can reach 95 percent. fourth is apparently not exploited, which In some specimens of Discradisca laevis, may explain differences in competitive abili- great numbers of full-grown Pedicellinae ties between inarticulated and articulated adhered to the long, barbed setae (DAVIDSON, brachiopods. Numerous examples of appar- 1880). One-third of the Discradisca shells ent spatial competition between D. strigata (17 percent of the total valve area) bore and other epifauna, particularly sponges and epizoans, primarily bryozoans and spiror- bryozoans, have been recorded (LABARBERA, bids, and occasional other Discradisca, serpu- 1985); but this species was spatially domi- lids, and small sponges. nant on only 3 of the 11 rocks investigated, Ecology of Inarticulated Brachiopods 495 even though it dominates in competitive in- that the entire shell margin, including re- teractions; and no individual appeared to be gions adjacent to the pedicle, sweeps through in any danger of overgrowth. In discinids the a sizeable arc when the animal rotates. This foramen, which is more centrally located inhibits growth of epifauna at a greater dis- than in articulated brachiopods, affords tance from the shell than is possible for ar- greater protection for the pedicle and ensures ticulated species.

BIOGEOGRAPHY OF INARTICULATED BRACHIOPODS

CHRISTIAN C. EMIG [Centre d’Océanologie de Marseille]

INTRODUCTION persist in our knowledge of the distribution and ecology of inarticulated species. The distribution of the inarticulated Populations of inarticulated species un- brachiopods is largely controlled by environ- dergo seasonal to continuous recruitment mental factors (see chapter on ecology of depending on their latitudinal distribution. inarticulated brachiopods, p. 473–495). The early, shelled larvae of the lingulides are Most of the inarticulated genera have broad common members of the tropical plankton. geographic distributions on which the dis- A Lingula female can spawn 28,000 oocytes persal potential of the larvae has had only a over a six-month period, and a Glottidia fe- small influence; lingulides and discinids have male may produce 130,000 ova over a four- planktotrophic larvae, while craniids have month period. The duration of the plank- short-lived, lecithotrophic larvae. The differ- tonic stage of lingulide larvae varies from 3 ences between species in their ecological re- to 6 weeks (CHUANG, 1959a; PAINE, 1963). quirements are more related to their ability Discinid larvae, at least Discinisca itself, to settle, which is induced by biotic or abi- are also planktotrophic and acquire valves otic factors of the biocoenosis to which the only in late stages (CHUANG, 1977). Discinid species belongs. All adult inarticulated larvae have been reported from marine brachiopods are exclusively sedentary. plankton from the water surface to depths of PATTERNS IN DISTRIBUTION 3,000 m, sometimes at great distances from the shore (HELMCKE, 1940; ODHNER, 1960; Because the biogeographic analyses of the CHUANG, 1977). Larvae of Discinisca have inarticulated taxa, especially discinids and been recorded from littoral waters down to craniids, cannot presently be based on infra- 350 m, while discinid larvae recorded from generic and subtle ecological distinctions or deep waters belong probably to Pelagodiscus broad geographic records, this account will atlanticus. For example, Pelagodiscus larvae be limited to the distribution of the genera. have been collected with a calculated density Many published records are deficient in pre- of 2 to 3.5 larvae per 1,000 m3 (MILEIKOVSKY, cise information on the biogeography and 1970) between depths of 500 and 2,000 m ecology of species. Sampling inarticulated in the northwestern Pacific Ocean. Postlarval brachiopods at depths beyond the range of specimens dredged from great depths (2,700 scuba may also present a misleading picture to 3,200 m) indicate that Pelagodiscus larvae of brachiopod distribution and abundance. become sedentary at different valve sizes. The use of submersibles provides reliable The lecithotrophic larva of Neocrania information only on large species that can be anomala, the only craniid species in which observed directly or by video. Another factor development has been studied (NIELSEN, that introduces bias is the propensity of 1991), has a short swimming stage of about craniids and discinids to settle on more or four to six days before settlement. Hydrody- less extensive, hard substrates that are namic conditions that occur in the biotope difficult to investigate with traditional of N. anomala (EMIG, 1989b) can disperse oceanographic sampling gear. Furthermore, larvae over several hundred kilometers dur- the attention paid to brachiopods in benthic ing this short stage. Hence, the gregarious studies and during oceanographic cruises is pattern of Neocrania species must be related frequently perfunctory so that large gaps to an environmental factor that attracts and 498 Brachiopoda

FIG. 415. Latitudinal distribution of inarticulated brachiopods (new). induces larval settlement close to the adult tribution of inarticulated taxa can be globally forms, not to the short swimming stage of related to their bathymetric extension (see the larvae. Fig. 413) although more constraints are in- The upper and lower limits of tolerance to volved than the temperature, pressure, and such factors as temperature, salinity, and dynamics of seawater. depth have been used generally to explain Most inarticulated genera are cosmopoli- the range of geographic distribution of the tan (Fig. 416) and were common in past species. As stated in the section on the ecol- eras. Indeed among living brachiopod fami- ogy of inarticulated brachiopods (Table 37, lies only the Lingulidae, Discinidae, and p. 484), however, such tolerances can vary Craniidae can be traced back to the early Pa- subtly even among populations and have to leozoic. The radiations of the inarticulated be analyzed carefully before being used to ex- species and genera represented in recent ma- plain the biogeographic distribution of rine faunas (Table 40) are related to geologi- higher taxa. cal events. Most genera began their develop- ment in the Cenozoic with the global DISTRIBUTION OF FAMILIES biotope changes marking the end of the Cre- AND GENERA taceous crisis, at the end of the Paleogene threshold, and during the Neogene as a re- The three extant inarticulated families sult of changes in the circulation of the ocean have a worldwide distribution. The Lingu- waters that allowed the development of lidae are dominant in tropical and subtropi- deep-sea species. cal areas; the Discinidae occur mainly in in- tertropical areas; the Craniidae are widely LINGULIDAE distributed from northern to southern high Living Lingulidae belong to two genera: latitudes, into which the discinid Pelagodiscus Lingula (seven species), which is worldwide also extends (Fig. 415). The latitudinal dis- in distribution, except along the coasts of Biogeography of Inarticulated Brachiopods 499

FIG. 416. Geographic distribution of inarticulated brachiopod genera (new). 500 Brachiopoda

TABLE 40. First geological record of the inartic- politan distribution and extend mainly over ulated genera represented in present the continental shelf. Discina striata has a re- marine faunas (new). stricted distribution in the intertropical zone of the western coast of Africa. Lingulidae Discinidae Craniidae Triassic Discinisca CRANIIDAE Upper Jurassic Craniscus Paleocene Lingula? Discradisca Neocrania (13 species) has a worldwide Glottidia? distribution (Fig. 416). Its latitudinal range Eocene Neocrania Miocene Pelagodiscus is as wide as that of Pelagodiscus, but its Holocene Discina Valdiviathyris? bathymetric distribution is from shallow Neoancistrocrania waters of the continental shelf to about 1,000 m depth on the bathyal slope (Fig. 417). Only one species, Neocrania lecointei, America, where Glottidia (five species) occurs is recorded in the deeper parts of the bathyal exclusively (Fig. 416). Large variations in zone (to 2,342 m). The two other genera edaphic factors during the late Mesozoic have restricted distributions. Craniscus japon- (EMIG, 1984b; BIERNAT & EMIG, 1993) are icus occurs in the western Pacific from 23 to probably responsible for the radiation of 885 m, while Valdiviathyris quenstedti is both genera. Glottidia may have originated known from a single location at 672 m. Neo- on the western coast of North and Central ancistrocrania norfolki has been collected in America and Lingula possibly in the islands two locations of the South Pacific Ocean at of the western Pacific. Their latitudinal 233 and 250 m depth. distribution occurs within the 40° belt from All three inarticulated brachiopod families temperate to equatorial areas (Fig. 417), and are of ancient stocks and are fairly cosmo- their bathymetric distribution is restricted to politan in distribution, extending from the the continental shelf except for Glottidia al- shoreline to the bathyal depths. Most species bida, which extends onto the upper part of have a distribution restricted to the 45° lati- the bathyal slope. Such a geographic distri- tudinal belt and occur on the continental bution appears to be a consequence of the shelf from intertidal to a depth of about 100 opening of the Atlantic Ocean and of the m. Species extending to latitudes higher than Paleocene-Eocene extension of the tropical- 45° occur also in the bathyal zone (between subtropical belt to about 45° latitude, with about 100 and 3,000 m). Their bathymetric optimal conditions for the development of extent, however, is limited mainly to the new temperate marine biotopes with good upper bathyal part (to 1,000 m). Only Pel- prospects for speciation. Yet the distribution agodiscus atlanticus, one of the most recent of the lingulides appears rather similar at species, is widespread in the abyssal zone least since the early Paleozoic when taking (3,000 to 6,000 m). Species of the two into account the paleolatitudinal positions in monospecific genera Discina striata and Val- correlation with temperatures of water diviathyris quenstedti and also Neoancistro- masses. crania norfolki, which is said to be recent as well, have a restricted geographic and bathy- DISCINIDAE metric distribution (Fig. 416–417). Pelagodiscus atlanticus occurs worldwide in In contrast to the lingulides, speculating deep water in the bathyal and abyssal zones on the origins and paths of dispersal of the and is undoubtedly the most widespread discinids and craniids is difficult. The brachiopod species geographically and present distribution of inarticulated taxa bathymetrically (Fig. 416–417). Discinisca cannot be explained as the consequence of (four species) and Discradisca (six species) their age or their dispersal rate as suggested have a warm-temperate to tropical, cosmo- for the taxa of articulated brachiopods. The Biogeography of Inarticulated Brachiopods 501

FIG. 417. Latitudinal and bathymetric extension of inarticulated brachiopod genera (new). 502 Brachiopoda diversification of the genera and the long inarticulated brachiopods. As ZEZINA (1970) geological history of species are relevant to previously stated the reasons for the our understanding of the extent of the biogeography of supraspecific brachiopod geographic and bathymetric distribution of taxa are elusive. REFERENCES Ackerly, Spafford. 1991. Hydrodynamics of rapid shell University of Glasgow. 116 p. closure in articulate brachiopods. Journal of Experi- Alvarez, Fernando. 1990. Devonian athyrid brachio- mental Biology 156:287–314. pods from the Cantabrian Zone (N.W. Spain). ———. 1992. Rapid shell closure in the brachiopods Biostratigraphie du Paléozoïque 11:311 p., 30 pl. Terebratulina retusa and Terebratalia transversa. Jour- Alvarez, Fernando, Covadonga Brime, & G. B. Curry. nal of the Marine Biological Association of the 1987. Growth and function of the micro-frills United Kingdom 72:579–598. present on the Devonian brachiopod Athyris cam- Adoutte, André, & H. Philippe. 1993. The major lines pomanesi (Verneuil & Archiac). Transactions of the of metazoan evolution: summary of traditional evi- Royal Society of Edinburgh 78:65–72. dence and lessons from ribosomal RNA sequence Alvarez, Fernando, & C. H. C. Brunton. 1990. The analysis. In Y. Pichon, ed., Comparative Molecular shell-structure, growth and functional morphology Neurobiology. Birkhauser. Basel. p. 1–30. of some Lower Devonian athyrids from northwest Afzelius, B. A. 1979. Sperm structure in relation to Spain. Lethaia 23(2):117–131, 12 fig. phylogeny in the lower Metazoa. In D. W. Fawcett Amsden, T. W. 1953. Some notes on the Pen- & J. M. Bedford, eds., The Spermatozoon. Matura- tameracea, including a description of one new ge- tion, Motility, Surface Properties and Comparative nus and one new subfamily. Washington Academy Aspects. Urban and Schwarzenberg. Baltimore. p. of Sciences, Journal 43(5):137–147, 7 fig. 243–251. An, S., & C. H. Koh. 1992. Environments and distri- Afzelius, B. A., & M. Ferraguti. 1978. Fine structure of bution of benthic animals on the Mangyung- brachiopod spermatozoa. Journal of Ultrastructural Dongjin tidal flat, west coast of Korea. Journal of Research 63:308–315. the Oceanological Society of Korea 27(1):78–90. Afzelius, B. A., & H. Mohri. 1966. Mitochondria re- spiring without exogenous substrate. A study of In Korean. aged sea urchin spermatozoa. Experimental Cell Re- Arber, M. A. 1942. The pseudodeltidium of the search 42:11–17. strophomenid brachiopods. Geological Magazine Aldridge, A. E. 1981. Intraspecific variation of shape 79:170–187. and size in subtidal populations of two recent New Armstrong, J. 1969. The cross-bladed fabrics of the Zealand articulate brachiopods. New Zealand Jour- shells of Terrakea solida (Etheridge and Dun) and nal of Zoology 8:169–174. Streptorhynchus pelicanensis Fletcher. Palaeontology ———. 1991. Shape variation of Neothyris (Brach- 12:310–320. iopoda, Terebratellinae). In D. I. MacKinnon, D. E. Asgaard, Ulla, & R. G. Bromley. 1991. Colonization Lee, & J. D. Campbell, eds., Brachiopods Through by micromorph brachiopods in the shallow subtidal Time, Proceedings of the 2nd International Brach- of the eastern Mediterranean Sea. In D. I. Mac- iopod Congress, University of Otago, Dunedin, Kinnon, D. E. Lee, & J. D. Campbell, eds., Brach- New Zealand, 1990. Balkema. Rotterdam. p. 115– iopods Through Time, Proceedings of the 2nd In- 122. ternational Brachiopod Congress, University of Alexander, R. R. 1986. Frequency of sublethal shell- Otago, Dunedin, New Zealand, 1990. Balkema. breakage in articulate brachiopod assemblages Rotterdam. p. 261–264. through geologic time. In P. R. Rachebœuf & C. C. Asgaard, Ulla, & N. Stentoft. 1984. Recent micro- Emig, eds., Les Brachiopodes Fossiles et Actuels, morph brachiopods from Barbados; palaeo- Actes du 1er Congrès International sur les ecological and evolutionary implications. Géobios, Brachiopodes, Brest 1985. Biostratigraphie du Mémoire spécial 8:29–37, pl. 1–2. Paléozoïque 4. p. 159–166, pl. 1. Ashworth, J. H. 1915. On larvae of Lingula and Allan, R. S. 1937. On a neglected factor in brachiopod Pelagodiscus (Discinisca). Transactions of the Royal migration. Records of the Canterbury Museum Society of Edinburgh 51:45–69, pl. 4–5. 4:157–165. Atkins, Dorothy. 1956. Ciliary feeding mechanisms of ———. 1949. Notes on a comparison of the Tertiary brachiopods. Nature 177:706. and recent Brachiopoda of New Zealand and South ———. 1958. A new species and genus of America. Transactions of the Royal Society of New Kraussinidae (Brachiopoda) with a note on feeding. Zealand 77:288–289. Proceedings of the Zoological Society of London ———. 1960. The succession of Tertiary brachiopod 131:559–581. faunas in New Zealand. Records of the Canterbury ———. 1959. The growth stages of the lophophore of Museum 7:233–268. the brachiopods Platidia davidsoni (Eudes Des- ———. 1963. On the evidence of Austral Tertiary and Longchamps) and P. anomioides (Phillipi), with recent brachiopods on Antarctic biogeography. In notes on the feeding mechanism. Journal of the Proceedings of the 10th Science Congress of the Marine Biological Association of the United King- Pacific Science Congress, University of Hawaii, dom 38:103–132. Honolulu. Bishop Museum Press. Honolulu. p. ———. 1960a. A new brachiopod from the Western 451–454. Approaches, and the growth stages of the lopho- Al-Rikabi, Ikbal. 1992. A molecular approach to phore. Journal of the Marine Biological Association palaeontology: Biochemical method applications of of the United Kingdom 39:71–89, fig. 1–14, pl. 1. brachiopod proteins. Master of Science thesis. ———. 1960b. The ciliary feeding mechanism of the 504 Brachiopoda

Megathyridae (Brachiopoda), and the growth stages Baker, P. G., & K. Laurie. 1978. Revision of the of the lophophore. Journal of the Marine Biologi- Aptian thecideidine brachiopods of the Faringdon cal Association of the United Kingdom 39:459– sponge gravels. Palaeontology 21:555–570. 479. Balakirev, E. S., & G. P. Manchenko. 1985. High lev- ———. 1961a. A note on the growth stages and struc- els of allozymic variation in brachiopod Coptothyris ture of the adult lophophore of the brachiopod grayii and Ascidia Halocynthia aurantium. Genetika Terebratella (Waltonia) inconspicua (G. B. Sowerby). 21:239–244. Proceedings of the Zoological Society of London Balinski, Andrzej. 1975. Secondary changes in micro- 136:255–271. ornamentation of some Devonian ambocoeliid ———. 1961b. The growth stage of the adult struc- brachiopods. Palaeontology 18:179–189. ture of the lophophore of the brachiopod Megerlia Bancroft, B. B. 1945. The brachiopod zonal indices of truncata (L.) and M. echinata (Fischer & Oelhert). the stages Costonian-Onnian in Britain. Journal of Journal of the Marine Biological Association of the Paleontology 19:181–252, pl. 22–38. United Kingdom 41:95–111. Baron, J., J. Clavier, & B. A. Thomassin. 1993. Struc- ———. 1963. Notes on the lophophore and gut of the ture and temporal fluctuations of two intertidal brachiopod Tegulorhynchia nigricans (G. B. seagrass-bed communities in New Caledonia (SW Sowerby). Proceedings of the Zoological Society of Pacific Ocean). Marine Biology 117:139–144. London 140:15–24. Bassett, M. G., L. E. Holmer, L. E. Popov, & John Atkins, Dorothy, & M. J. S. Rudwick. 1962. The lo- Laurie. 1993. Phylogenetic analysis and classifica- phophore and ciliary feeding mechanism of the tion of the Brachiopoda—reply and comments. brachiopod Crania anomala (Müller). Journal of the Lethaia 26:385–386. Marine Biological Association of the United King- Bayne, B. L., & R. C. Newell. 1983. Physiological dom 42:469–480. energetics of marine molluscs. In A. S. M. Avise, J. C. 1994. Molecular markers, natural history Saleuddin & K. M. Wilbur, eds., The Mollusca, and evolution. Chapman & Hall. New York & Vol. 4. Academic Press. New York. p. 407–515. London. 511 p. Bayne, B. L., J. Widdows, & R. J. Thompson. 1976. Awati, P. R., & G. R. Kshirsagar. 1957. Lingula from Physiological integrations. In B. L. Bayne, ed., western coast of India. Zoological Memoirs of the Marine Mussels: Their Ecology and Physiology. University of Bombay 4:1–87. Cambridge University Press. Cambridge. p. 261– Ayala, F. J., J. W. Valentine, T. E. DeLaca, & G. S. 291. Zumwalt. 1975. Genetic variability of the Antarc- Beecher, C. E. 1891. Development of the Brach- tic brachiopod Liothyrella notorcadensis and its bear- iopoda. Part I. Introduction. American Journal of ing on mass extinction hypotheses. Journal of Pale- Science (series 3) 41:343–357. ontology 49:1–9. ———. 1892. Development of the Brachiopoda. Part Babin, C., J. H. Delance, C. C. Emig, & P. R. II. Classification of the stages of growth and de- Racheboeuf. 1992. Brachiopodes et Mollusques cline. American Journal of Science (series 3) Bivalves: concurrence ou indifférence? Géobios, 44:133–155. Mémoire spécial 14:35–44. ———. 1897. Morphology of the brachia. Bulletin of Backeljau, T., B. Winnepenninckx, & L. De Bruyn. the United States Geological Survey 87:105–112. 1993. Cladistic analysis of metazoan relationships: Beecher, C. E., & J. M. Clarke. 1889. The develop- a reappraisal. Cladistics 9:167–181. ment of some Silurian Brachiopoda. New York Bada, J. L., M. Y. Shou, E. H. Man, & R. A. State Museum Memoir 1(1):1–95, pl. 1–8. Schroeder. 1978. Decomposition of hydroxy amino Bemmelen, J. F. van. 1883. Untersuchungen über den acids in foraminifera tests; kinetics, mechanism and anatomichen und histologichen Bau der Brachio- geochronological implications. Earth and Planetary poda Testicardina. Jena. Zeitschrift für Natur- Science Letters 41:67–76. wissenschaften 16:88–161. Baker, P. G. 1970a. Significance of the punctation Benigni, Chiara, & Carla Ferliga. 1988. Carniana mosaic of the Jurassic thecidellinid brachiopod Thecospiridae (Brachiopoda) from San Cassiano Moorellina. Geological Magazine 107:105–113. Formation (Cortina d’Ampezzo, Italy). Rivista ———. 1970b. The growth and shell microstructure Italiana di Palaeontologia 94:515–560. of the thecideacean brachiopod Moorellina granu- Benson, D. A., M. Boguski, D. L. Lipman, & J. Ostell. losa (Moore) from the Middle Jurassic of England. 1994. GenBank. Nucleic Acids Research 22:3441– Palaeontology 13:76–99. 3444. ———. 1990. The classification, origin and phylogeny Benton, M. J., ed. 1993. The Fossil Record 2. of thecideidine brachiopods. Palaeontology Chapman & Hall. London. 845 p. 33(1):175–191, 3 fig. Bernard, F. R. 1972. The living Brachiopoda of British ———. 1991. Morphology and shell microstructure Columbia. Syesis 5:73–82. of Cretaceous thecideidine brachiopods and their Beyer, H. G. 1886. A study of the structure of Lingula bearing on thecideidine phylogeny. Palaeontology (Glottidia) pyramidata Stim. (Dall). Studies from 34:815–836. the Biology Laboratory, Johns Hopkins University Baker, P. G., & D. G. Elston. 1984. A new polyseptate 3:227–265. thecideacean brachiopod from the middle Jurassic Bhavanarayana, P. V. 1975. Some observations on the of the Cotswolds, England. Palaeontology 27:777– benthic faunal distribution in the Kakinada Bay. In 791. R. Natarajan, ed., Recent Researches in Estuarine References 505

Biology. Hindustan Publishing Co. Dehli. p. 146– Evolution 42:795–803. 150. Britten, R. J., & E. H. Davidson. 1971. Repetitive and Biernat, Gertrude, & C. C. Emig. 1993. Anatomical non-repetitive DNA sequences and a speculation on distinctions of Mesozoic lingulide brachiopods. the origins of evolutionary novelty. Journal of Mo- Acta Palaeontologica Polonica 38(1/2):1–20, 8 fig. lecular Evolution 46:111–133. Biernat, Gertrude, & Alwyn Williams. 1970. Ultra- Britten, R. J., & D. E. Kohne. 1968. Repeated se- structure of the protegulum of some acrotretide quences in DNA. Science 161:529–540. brachiopods. Palaeontology 13:491–502. Bromley, R. G., & Finn Surlyk. 1973. Borings pro- ———. 1971. Shell structure of the siphonotretacean duced by brachiopod pedicles, fossil and recent. Brachiopoda. Palaeontology 14:423–430. Lethaia 6:349–365. Bitter, P. H. von, & R. Ludvigsen. 1979. Formation Brooks, W. K. 1879. The development of Lingula and and function of protegular pitting in some North the systematic position of the Brachiopoda. Johns American acrotretid brachiopods. Palaeontology Hopkins University, Chesapeake Zoology Labora- 22:705–720. tory, Scientific Results of the Session of 1878:34– Blochmann, F. 1892. Untersuchungen über den Bau 112. der Brachiopoden. I. Die Anatomie von Crania Brown, I. A. 1953. Martinopsis Waagen from the Salt anomala (Müller). Jena. Gustav Fischer. p. 1–65. Range, India. Journal and Proceedings of the Royal ———. 1898. Die larve von Discinisca (Die Society of New South Wales 86:100–107, 3 fig., pl. Muellersche Brachiopodenlarve). Zoologische 9. Jahrbücher Abteilungen Anatomie und Ontogenie Brunton, C. H. C. 1966. Silicified productoids from der Tiere 11:417–426. the Visean of County Fermanagh. British Museum ———. 1900. Untersuchungen über den Bau der (Natural History) Bulletin (Geology) 12(5):175– Brachiopoden. I, Die Anatomie von Crania 243, pl. 1–19. anomala O.F.M. (1892). II. Die Anatomie von ———. 1969. Electron microscopic studies of growth Discinisca lamellosa (Broderip) und Lingula anatina margins of articulate brachiopods. Zeitschrift für (Bruguière). Gustav Fischer. Jena. p. 1–124. Sellforsch 100:189–200. ———. 1906. Neue Brachiopoden der Valdivia-und ———. 1971. An endopunctate rhynchonellid brach- Gaussexpeditionen. Zoologischer Anzeiger 30:690– iopod from the Viséan of Belgium and Britain. 702. Palaeontology 14:95–106. ———. 1908. Zur Systematik und geographischen ———. 1972. The shell structure of chonetacean Verbreirung der Brachiopoden. Zeitschrift für brachiopods and their ancestors. Bulletin of the Bri- wissenschaftliche Zoologie 90:596–644, pl. 36–40. tish Museum of Natural History (Geology) 21:1– Boore, J. L., & W. M. Brown. 1994. Complete DNA 26. sequence of the mitochondrial genome of the black ———. 1988. Some brachiopods from the eastern chiton, Katharina tunicata. Genetics 138:423–443. Mediterranean Sea. Israel Journal of Zoology Borman, A. H., E. W. de Jong, R. Thierry, P. West- 35:151–169. broek, & L. Bosch. 1987. Coccolith-associated Brunton, C. H. C., & G. B. Curry. 1979. British polysaccharides from cells of Emiliania huxleyi brachiopods. Synopses of the British fauna (new se- (Haptophycae). Journal of Phycology 23:118–123. ries) 17:64 p., 30 fig. Bosi Vanni, M. R., & A. M. Simmonetta. 1967. Con- Brunton, C. H. C., & Norton Hiller. 1990. Late tributo alla conoscenza dell’anatomia ed isologia di Cainozoic brachiopods from the coast of Nama- Muehlfedtia disculus (Pallas) 1766 (Brachiopoda, qualand, South Africa. Palaeontology 33(2):313– Testicardines). Gli apparati lofoforale, digerente, 342, 11 fig. nefridial e reproduttore. Societa Toscana di Scienze Brunton, C. H. C., & D. J. C. Mundy. 1988. Naturali, Memorie (series B) 74B:21–34. Strophalosiacean and aulostegacean productoids Boucot, A. J. 1959. Brachiopods of the lower Devo- (Brachiopoda) from the Craven Reef Belt (late nian rocks at Highland Mills, New York. Journal of Viséan) of North Yorkshire. Proceedings of the Paleontology 33(5):727–769, 5 fig., pl. 90–103. Yorkshire Geological Society 47:55–88. Bozzo, M. G., R. Bargalló, M. Durfort, R. Fontarnau, Brusca, R. C., & G. J. Brusca. 1990. Invertebrates. & J. López-Camps. 1983. Ultrastructura de la Sinauer Associates, Inc. Sunderland, Massachusetts. coberta des oòcits de Terebratula vitrea (Brach- 922 p. iopoda: Testicardina). Butllettí de la Institució Buchan, P., L. S. Peck, & N. Tublitz. 1988. A light, Catalana d’Història Natural 49 (Secció de Zoologia, portable apparatus for the assessment of inverte- 5):13–18, fig. 1–11. brate heart beat rate. Journal of Experimental Biol- Brafield, A. E., & D. J. Solomon. 1972. Oxycalorific ogy 136:495–498. coefficients for animals respiring nitrogenous sub- Bullivant, J. S. 1968. The method of feeding of strates. Comparative Biochemistry and Physiology lophophorates (Bryozoa, Phoronida, Brachiopods). 43A:837–841. New Zealand Journal of Marine and Freshwater Branch, G. M. 1981. The biology of limpets: physical Research 2:135–146. factors, energy flow and ecological interactions. Bulman, O. M. B. 1939. Muscle systems of some in- Oceanography and Marine Biology: An Annual articulate brachiopods. Geological Magazine Review 19:235–380. 76:434–444. Bremer, K. 1988. The limits of amino acid sequence Campbell, K. S. W., & B. D. E. Chatterton. 1979. data in angiosperm phylogenetic reconstruction. Coelospira: do its double spires imply a double 506 Brachiopoda

lophophore? Alcheringa 3:209–223, fig. 1–7. II. John Wiley and Sons Ltd. p. 517–530. Carlson, S. J. 1989. The articulate brachiopod hinge ———. 1990. Brachiopoda. In K. G. Adiyodi & R. G. mechanism: morphological and functional varia- Adiyodi, eds., Reproductive Biology of Inverte- tion. Paleobiology 15(4):364–386, 13 fig. brates. Fertilisation, Development and Parental ———. 1993a. Investigating brachiopod phylogeny Care, Vol. VI. John Wiley and Sons Ltd. p. 211– and classification—response to Popov et al. 1993. 254. Lethaia 26:383–384. ———. 1991. Common and evolutionary features of ———. 1993b. Phylogeny and evolution of recent brachiopods and their bearing on the rela- “pentameride” brachiopods. Palaeontology tionship between, and the monophyletic origin of, 36(4):807–837, 10 fig. the inarticulates and the articulates. In D. I. Mac- Carpenter, W. B. 1851. On the intimate structure of Kinnon, D. E. Lee, & J. D. Campbell, eds., Brach- shells of Brachiopoda. In T. Davidson, British Fos- iopods Through Time, Proceedings of the 2nd In- sil Brachiopoda. Monograph of the Palaeon- ternational Brachiopod Congress, University of tographical Society 1:22–40. Otago, Dunedin, New Zealand, 1990. Balkema. Carter, J. L., J. G. Johnson, R. Gourvennec, & H.-F. Rotterdam. p. 11–14. Hou. 1994. A revised classification of the spiriferid Clegg, H. 1993. Biomolecules in recent and fossil ar- brachiopods. Annals of the Carnegie Museum ticulate brachiopods. Ph.D. thesis. University of 63:327–374. Newcastle Upon Tyne. 246 p. Caryl, Anthony. 1992. DNA fingerprinting of brachio- Cloud, P. E. Jr. 1942. Terebratuloid Brachiopoda of the pod DNA. Unpublished Honours Bachelor of Sci- Silurian and Devonian. Geological Society of ence thesis. University of Glasgow. 37 p. America Special Paper 38:1–182, pl. 1–26. Chuang, S. H. 1956. The ciliary feeding mechanisms ———. 1948. Notes on recent brachiopods. American of Lingula unguis (L.) (Brachiopoda). Proceedings Journal of Science 246:241–250. of the Zoological Society of London 127(2):167– Cock, A. G. 1966. Genetical aspects of metrical 189. growth and form in animals. Quarterly Review of ———. 1959a. The breeding season of the brachiopod Biology 41:131–190. Lingula unguis (L.). Biological Bulletin, Marine Bi- Cohen, B. L. 1992. Utility of molecular phylogenetic ology Laboratory, Woods Hole, Massachusetts methods: a critique of immuno-taxonomy. Lethaia 117(2):202–207. 24:441–442. ———. 1959b. The structure and function of the ali- ———. 1994. Immuno-taxonomy and the reconstruc- mentary canal in Lingula unguis (L.) Brachiopoda. tion of brachiopod phylogeny. Palaeontology Proceedings of the Zoological Society of London 37:907–911. 132:283–311. Cohen, B. L., Peter Balfe, Moyra Cohen, & G. B. ———. 1960. An anatomical, histological and his- Curry. 1991. Molecular evolution and morphologi- tochemical study of the gut of the brachiopod Cra- cal speciation in North Atlantic brachiopods nia anomala. Quarterly Journal of Microscopical (Terebratulina spp.). Canadian Journal of Zoology Science 101:9–18. 69:2903–2911. ———. 1961. Growth of the postlarval shell in Lin- ———. 1993. Molecular and morphometric variation gula unguis (L.) (Brachiopoda). Proceedings of the in European populations of the articulate brachio- Zoological Society of London 137(2):299–310. pod Terebratulina retusa. Marine Biology 115:105– ———. 1968. The larvae of a discinid (Inarticulata, 111. Brachiopoda). Biological Bulletin 135:263–272. Cohen, B. L., Peter Balfe, & G. B. Curry. 1986. Ge- ———. 1971. New interpretation of the morphology netics of the brachiopod Terebratulina. In P. R . of Schizambon australis Ulrich and Cooper (Ordovi- Rachebœuf & C. C. Emig, eds., Les Brachiopodes cian siphonotretid inarticulate brachiopod). Journal Fossiles et Actuels, Actes du 1er Congrès Interna- of Paleontology 45:824–832. tional sur les Brachiopodes, Brest 1985. Biostra- ———. 1973. The inarticulate brachiopod larvae of tigraphie du Paléozoïque 4. p. 55–63. the International Indian Ocean Expedition. Journal Cohen, B. L., A. B. Gawthrop, & T. Cavalier-Smith. of the Marine Biological Association of India In preparation. Phylogeny of lophophorates, espe- 15:538–544. cially brachiopods, based on nuclear-encoded SSU ———. 1974. Observations on the ciliary feeding rRNA gene sequences. mechanisms of the brachiopod Crania anomala. Collins, M. J. 1991. Growth rate and substrate-related Journal of the Zoological Society of London mortality of a benthic brachiopod population. 173:441–449. Lethaia 24:1–11. ———. 1977. Larval development in Discinisca (inar- Collins, M. J., G. B. Curry, G. Muyzer, R. Quinn, S. ticulate brachiopod). American Zoologist 17:39– Xu, P. Westbroek, & S. Ewing. 1991. Immunologi- 53. cal investigations of relationships within the tere- ———. 1983a. Brachiopoda. In K. G. Adiyodi & R. bratulid brachiopods. Palaeontology 34:785–796. G. Adiyodi, eds., Reproductive Biology of Inverte- Collins, M. J., G. B. Curry, R. Quinn, G. Muyzer, T. brates. Oogenesis, Oviposition and Oosorption, Zomerdijk, & P. Westbroek. 1988. Sero-taxonomy Vol. I. John Wiley and Sons Ltd. p. 571–584. of skeletal macromolecules in living terebratulid ———. 1983b. Brachiopoda. In K. G. Adiyodi & R. brachiopods. Historical Biology 1:207–224. G. Adiyodi, eds., Reproductive Biology of Inverte- Collins, M. J., G. Muyzer, G. B. Curry, P. Sandberg, & brates. Spermatogenesis and Sperm Function, Vol. P. Westbroek. 1991. Macromolecules in brachiopod References 507

shells: characterisation and diagenesis. Lethaia Smithsonian Contributions to Paleobiology 41:1– 24:387–397. 43, fig. 1–4, pl. 1–7. Collins, M. J., G. Muyzer, P. Westbroek, G. B. Curry, ———. 1983. The Terebratulacea (Brachiopoda), Tri- P. A. Sandberg, S. J. Xu, R. Quinn, & D. I. assic to recent: A study of the brachidia (loops). MacKinnon. 1991. Preservation of fossil biopoly- Smithsonian Contributions to Paleobiology 50:1– meric structures: conclusive immunological evi- 445, 17 fig., 77 pl., 86 tables. dence. Geochimica et Cosmochimica Acta ———. 1988. Some Tertiary brachiopods of the east 55:2253–2257. coast of the United States. Smithsonian Contribu- Conklin, E. G. 1902. The embryology of a brachio- tions to Paleobiology 64:1–45. pod, Terebratulina septentrionalis Couthouy. Pro- Cooper, G. A., & P. J. Doherty. 1993. Calloria ceedings of the American Philosophical Society variegata, a new recent species of brachiopod 41:41–76. (Articulata: Terebratulida) from northern New Conover, R. J., & E. D. S. Corner. 1968. Respiration Zealand. Journal of the Royal Society of New and nitrogen excretion by some marine zooplank- Zealand 23:271–281. ton in relation to their life cycles. Journal of the Cooper, G. A., & R. E. Grant. 1974. Permian brachio- Marine Biological Association of the United King- pods of West Texas, II. Smithsonian Contributions dom 48:49–75. to Paleobiology 15:233–793. Conway Morris, Simon. 1994. Why molecular biology ———. 1975. Permian brachiopods of West Texas, III. needs palaeontology. In M. Akam, P. Holland, P. Smithsonian Contributions to Paleobiology Ingham, & G. Wray, eds., The Evolution of Devel- 19:795–1298, pl. 192–502. opmental Mechanisms. Development, 1994 ———. 1976. Permian brachiopods of West Texas, IV. Supplement. Company of Biologists. Cambridge. p. Smithsonian Contributions to Paleobiology 1–13. 21:1923–2285, pl. 503–662. ———. 1995. Nailing the lophophorates. Nature Cooper, G. A., & F. G. Stehli. 1955. New genera of 375:365–366. Permian brachiopods from West Texas. Journal of Conway, Morris, Simon, B. L. Cohen, A. B. Gaw- Paleontology 29:469–474, pl. 52–54. throp, T. Cavalier-Smith, & B. Winnepenninckx. Copper, Paul. 1965. A new Middle Devonian atrypid 1996. Lophophorate phylogeny. Science 272:282. brachiopod from the Eifel, Germany. Sencken- Conway Morris, Simon, & J. S. Peel. 1995. Articulated bergiana Lethaea 46(4/6):309–325, 13 fig., pl. 27. halkieriids from the Lower Cambrian of north ———. 1986. Evolution of the earliest smooth spire- Greenland and their role in early protostome evolu- bearing Atrypoids (Brachiopoda: Lissatrypidae, Or- tion. Philosophical Transactions of the Royal Soci- dovician–Silurian). Palaeontology 29(4):827–866, ety (series B) 347:305–358. pl. 73–75. Cooper, G. A. 1942. New genera of North American Cornish-Bowden, A. 1983. Relating proteins by amino brachiopods. Washington Academy of Sciences acid composition. Methods in Enzymology 91:60– Journal 32(8):228–235. 75. ———. 1952. Unusual specimens of the brachiopod Cowen, R. 1966. The distribution of punctae on the family Isogrammidae. Journal of Paleontology brachiopod shell. Geological Magazine 103:269– 26:113–119, pl. 21–23. 275. ———. 1954. Unusual Devonian brachiopods. Jour- ———. 1968. A new type of delthyrial cover in the nal of Paleontology 28:325–332, 5 fig., pl. 36–37. Devonian brachiopod Mucrospirifer. Palaeontology ———. 1955. New genera of middle Paleozoic brach- 11:317–327, pl. 63–64. iopods. Journal of Paleontology 29:45–63, pl. 11– Cowen, R., & M. J. S. Rudwick. 1970. Deltidial 14. spines in the Triassic brachiopod Bittnerula. ———. 1956. New Pennsylvanian brachiopods. Jour- Palaeontologische Zhurnal 44:82–85, pl. 9. nal of Paleontology 30:521–530, pl. 61. Crenshaw, M. A. 1972. The soluble matrix from ———. 1957. Loop development of the Pennsylvanian Mercenaria mercenaria shell. Biomineralization 6:6– terebratuloid Cryptacanthia. Smithsonian Miscella- 11. neous Collection 134(3):1–18, pl. 1–2. Culter, J. M. 1979. A population study of the inarticu- ———. 1973. Vema’s Brachiopoda (recent). Smithso- late brachiopod Glottidia pyramidata (Stimpson). nian Contributions to Paleobiology 17:1–51. Unpublished master of science thesis. University of ———. 1975. Brachiopods from west African waters South Florida. Tampa. 53 p. with examples of collateral evolution. Journal of Pa- ———. 1980. The occurrence of hermaphrodites of leontology 49:911–927, fig. 1–7, pl. 1–4. the inarticulate brachiopod Glottidia pyramidata. ———. 1977. Brachiopods from the Caribbean Sea Florida Scientist 43(1):20. and adjacent waters. Studies in Tropical Oceanog- Culter, J. M., & J. L. Simon. 1987. Sex ratios and the raphy Miami 14. Rosenstiel School of Marine and occurrence of hermaphrodites in the inarticulate Atmospheric Science. University of Miami Press. brachiopod, Glottidia pyramidata (Stimpson) in 212 p., 6 fig., 35 pl. Tampa Bay, Florida. Bulletin of Marine Science of ———. 1981. Brachiopods from the southern Indian the Gulf and Caribbean 40:193–197. Ocean. Smithsonian Contributions to Paleobiology Curry, G. B. 1981. Variable pedicle morphology in a 43:1–93, fig. 1–30, pl. 1–14. population of the recent brachiopod Terebratulina ———. 1982. New Brachiopoda from the southern septentrionalis. Lethaia 14:9–20. hemisphere and Cryptopora from Oregon (recent). ———. 1982. Ecology and population structure of the 508 Brachiopoda

recent brachiopod Terebratulina retusa from Scot- ———. 1974. Triasovye brakhiopody (morfologiya, land. Palaeontology 25(2):227–246. sistema, filogeniya, stratigraficheskoye znacheniye i ———. 1983. Microborings in recent brachiopods and biogeografiya) [Triassic Brachiopoda (morphology, the functions of caeca. Lethaia 16:119–127. systematics, phylogeny, stratigraphic distribution, Curry, G. B., & A. D. Ansell. 1986. Tissue mass in liv- and biogeography)]. Akademia Nauk SSSR ing brachiopods. In P. R. Rachebœuf & C. C. Sibiroskoe Otdelenie Institut Geologii I Geofiziki Emig, eds., Les Brachiopodes Fossiles et Actuels, (IGIG) Trudy [Institute of Geology and Geophys- Actes du 1er Congrès International sur les Brach- ics, Academy of Sciences of the USSR, Siberian iopodes, Brest 1985. Biostratigraphie du Paléo- Branch, Transactions] 214:386 p., 171 fig., 49 pl. zoïque 4. p. 231–241. Darwin, Charles. 1859a. On the Origin of Species. 1st Curry, G. B., A. D. Ansell, Mark James, & L. S. Peck. edition. John Murray. London. 490 p. 1989. Physiological constraints on living and fossil ———. 1859b. On the origin of the species by means brachiopods. Transactions of the Royal Society of of natural selection or the preservation of favoured Edinburgh, Earth Sciences 80:255–262. races in the struggle for life. The Folio Society edi- Curry, G. B., Maggie Cusack, Kazuyoshi Endo, Derek tion (1990). 298 p. Walton, & R. Quinn. 1991. Intracrystalline mol- Davidson, Thomas. 1850. Sur quelques Brachiopodes ecules from brachiopod shells. In S. Suga & H. nouveaux ou peu connus. Bulletin de la Société Nakahara, eds., Mechanisms and Phylogeny of Géologique de France (série 2) 8:62–74, 1 pl. Mineralization in Biological Systems. Springer- ———. 1853. A Monograph of the British fossil Verlag. Tokyo. p. 35–40. Brachiopoda (Volume 1: General introduction). Curry, G. B., Maggie Cusack, Derek Walton, Palaeontographical Society (London) Monograph Kazuyoshi Endo, H. Clegg, G. Abbott, & H. 7:1–136, pl. 1–9. Armstrong. 1991. Biogeochemistry of brachiopod ———. 1880. Report on the Brachiopoda dredged by intracrystalline molecules. Philosophical Transac- the HMS Challenger during the years 1873–1876. tions of the Royal Society of London 333B:359– Report of the Scientific Results of the Voyage of 366. HMS Challenger 1(1):1–67, pl. 1–4. Curry, G. B., R. Quinn, M. J. Collins, Kazuyoshi Davis, C. C. 1949. Observations of plankton taken in Endo, S. Ewing, G. Muyzer, & P. Westbroek. 1991. marine waters of Florida in 1947 and 1948. Quar- Immunological responses from brachiopod skeletal terly Journal of the Florida Academy of Science macromolecules; a new technique for assessing 11:67–103. taxonomic relationships using shells. Lethaia Degens, E. T., D. W. Spencer, & R. H. Parker. 1967. 24:399–407. Paleobiochemistry of molluscan shell proteins. ———. 1993. Molecules and morphology—the prac- Comparative Biochemistry and Physiology 20:553– tical approach. Lethaia 26:5–6. 579. Cusack, Maggie, G. B. Curry, H. Clegg, & G. Abbott. Devereaux, P., P. Haeberli, & O. Smithies. 1984. A 1992. An intracrystalline chromoprotein from red comprehensive set of sequence analysis programs brachiopod shells: implications for the process of for the VAX. Nucleic Acids Research 12:387–395. biomineralisation. Comparative Biochemistry and Dexter, D. M. 1983. Soft bottom infaunal communi- Physiology 102B:93–95. ties in Mission Bay. California Fish Game 69(1):5– Cusack, Maggie, & Alwyn Williams. 1996. Chemico- 17. structural degradation of Carboniferous lingulid Di Geronimo, I., & G. Fredj. 1987. Les fonds à Errina shells. Philosophical Transactions of the Royal So- aspera et Pachylasma giganteum. Documents et ciety of London B351:33–49. Travaux de l’Institut de Géologie "Albert de Dagys, A. S. 1968. Jurskiye i rannemelovye Lapparent" (Paris) 11:243–247. brakhiopody Severa Sibiri [Jurassic and Early Cre- Doherty, P. J. 1976. Aspects of the feeding ecology of taceous brachiopods from northern Siberia]. the subtidal brachiopod, Terebratella inconspicua Akademia Nauk SSSR Sibiroskoe Otdelenie Institut (Sowerby, 1846). Master of Science Thesis. Depart- Geologii I Geofiziki (IGIG) Trudy [Institute of ment of Zoology, University of Auckland. 183 p. Geology and Geophysics, Academy of Sciences of ———. 1979. A demographic study of a subtidal the USSR, Siberian Branch, Transactions] 41:167 population of the New Zealand articulate brachio- p., 81 fig., 26 pl. pod Terebratella inconspicua. Marine Biology ———. 1972. Postembrional’noye razvitiye brakhidiya 52:331–342. pozdnepaleozoyskikh i rannemezozoyskikh Tere- ———. 1981. The contribution of dissolved amino bratulida [Postembryonic development of the acids to the nutrition of brachiopods. New Zealand brachidium of late Paleozoic and early Mesozoic Journal of Zoology 8:183–188. terebratulids]. In A. S. Dagys & A. B. Ivanovskii, Dolah, R. F. van., D. R. Calder, & D. M. Knott. 1983. eds., Morphologicheskiye i filogeneticheskiye Assessment of the benthic macrofauna in an ocean voprosy paleontologii [Morphological and Phyloge- disposal area near Charleston, South Carolina. netic Questions of Paleontology]. Akademia Nauk Technical Report, South Carolina Marine Resource SSSR Sibiroskoe Otdelenie Institut Geologii I Center 56:1–97. Geofiziki (IGIG) Trudy [Institute of Geology and Dolah, R. F. van., D. M. Knott, E. L. Wenner, T. D. Geophysics, Academy of Sciences of the USSR, Si- Mathews, & M. P. Katuna. 1984. Benthic studies berian Branch, Transactions] 112:22–58, fig. 1–28. and sedimentological studies of the Georgetown References 509

ocean dredged material disposal site. Technical Re- ———. 1981a. Implications de données récentes sur port, South Carolina Marine Resource Center les lingules actuelles dans les interprétations 59:1–97. paléoécologiques. Lethaia 14(2):151–156. Donoghue, M. J., & P. D. Cantino. 1984. The logic ———. 1981b. Observations sur l’écologie de Lingula and limitations of the outgroup substitution ap- reevii Davidson (Brachiopoda: Inarticulata). Journal proach to cladistic analysis. Systematic Botany of Experimental Marine Biology and Ecology 9:192–202. 52(1):47–61. Dörjes, J. 1977. Marine macrobenthic communities of ———. 1982. Terrier et position des lingules the Sapelo island, Georgia region. In B. C. Coull, (brachiopodes, inarticulés). Bulletin de la Société ed., Ecology of Marine Benthos. University of Zoologique de France 107(2):185–194. South Carolina Press, Columbia, Belle W. Baruch ———. 1983a. Comportement expérimental de Lin- Library in Marine Science 6:399–421. gula anatina (brachiopode, inarticulé) dans divers ———. 1978. Sedimentology and faunistics of tidal substrats meubles (Baie de Mutsu, Japon). Marine flats in Taiwan. II. Faunistic and ichnocoenotic Biology 75(2/3):207–213. studies. Senckenbergiana Maritima 10(1/3):85– ———. 1983b. Taxonomie du genre Glottidia 115. (brachiopodes inarticulés). Bulletin du Museum Doumen, C., & W. R. Ellington. 1987. Isolation and National d’Histoire Naturelle de Paris (série 4) characterization of a taurine-specific opine dehydro- 5(section 4; no. 2):469–489. genase from the pedicles of the brachiopod, Glot- ———. 1984a. Importance du sédiment dans la distri- tidia pyramidata. Journal of Experimental Zoology bution des lingules (brachiopodes, inarticulés). 243:25–31. Lethaia 17:115–123. Du Bois, H. M. 1916. Variation induced in brachio- ———. 1984b. Pourquoi les lingules (brachiopodes, pods by environmental conditions. Publications of inarticulés) ont survécu à la transition Secondaire- the Puget Sound Marine Station 1:177–183. Tertiaire. Bulletin de la Commission des Travaux Dunning, H. N. 1963. Geochemistry of organic pig- historiques et scientifiques, Section Sciences 6:87– ments: carotenoids. In I. A. Berger, ed., Organic 94. Geochemistry. Macmillan. New York. p. 367–430. ———. 1986. Conditions de fossilisation du genre Dweltz, N. E. 1961. The structure of β-chitin. Lingula (Brachiopoda) et implications paléoéco- Biochimica et Biophysica Acta 51:283–294. logiques. Palaeogeography, Palaeoclimatology, Eernisse, D. J., J. S. Albert, & F. E. Anderson. 1992. Palaeoecology 53:245–253. Annelida and arthropoda are not sister taxa: a phy- ———. 1987. Offshore brachiopods investigated by logenetic analysis of spiralian metazoan morphol- submersible. Journal of Experimental Marine Biol- ogy. Systematic Biology 41:305–330. ogy and Ecology 108:261–273. Eernisse, D. J., & A. G. Kluge. 1993. Taxonomic con- ———. 1988. Les brachiopodes actuels sont-ils des gruence versus total evidence and amniote phylog- indicateurs (paléo) bathymétriques? Géologie eny inferred from fossils, molecules and morphol- Méditerranéenne 15(1):65–71. ogy. Molecular Biology and Evolution ———. 1989a. Les lingules fossiles, représentants 10:1170–1195. d’écosystèmes oligotypiques? Atti 3° Simposio di Eglinton, G., & G. A. Logan. 1991. Molecular preser- Ecologia e Paleoecologia delle Comunita bento- vation. Philosophical Transactions of the Royal So- niche, Catania 1985:117–121. ciety of London 333B:315–328. ———. 1989b. Distribution bathymétrique et spatiale Egorov, A. N., & L. Ye. Popov. 1990. Novyi rod des populations de Gryphus vitreus (brachiopode) Lingulid iz niznepermskikh otlozhenii Sibroskoi sur la marge continentale (Nord-Ouest Méditer- Platformy [A new lingulid genus from the Early ranée). Oceanologica Acta 12(2):205–209. Permian deposits of the Siberian Platform]. ———. 1989c. Distributional patterns along the Paleontologicheskii Zhurnal 1990(4):111–115, fig. Mediterranean continental margin (upper bathyal) 1–3. using Gryphus vitreus (Brachiopoda) densities. Eichler, P. 1911. Die Brachiopoden der deutschen Palaeogeography, Palaeoclimatology, Palaeoecology Südpolar-Expedition 1901 bis 1903. Deutsche 71:253–256. Südpolar-Expedition 1901–1903 (series 12) ———. 1990. Examples of post-mortality alteration in Zoologie 4:381–401. recent brachipod shells and (paleo) ecological con- Ekman, T. 1896. Beitrage zur Kenntnis des Steiles der sequences. Marine Biology 104:233–238. Brachiopoden. Zeitschrift für Wissenschaftliche ———. 1992. Functional disposition of the lopho- Zoologie 62:169–249. phore in living Brachiopoda. Lethaia 25:291–302. Elliott, G. F. 1951. On the geographical distribution of Emig, C. C., & P. Cals. 1979. Lingules d’Amboine, terebratelloid brachiopods. Annals and Magazine of Lingula reevii Davidson et Lingula rostrum (Shaw), Natural History (series 12) 5:305–333. données écologiques et taxonomiques concernant ———. 1953. Brachial development and evolution in les problèmes de spéciation et de répartition. terebratelloid brachiopods. Cambridge Philosophi- Cahiers de l’Indo-pacifique 2:153–164. cal Society Biological Reviews 28:261–279, 9 fig. Emig, C. C., J. C. Gall, D. Pajaud, & J. C. Plaziat. Emig, C. C. 1976. Le lophophore-structure significa- 1978. Réflexions critiques sur l’écologie et la tive des Lophophorates (Brachiopoda, Bryozoa, systématique des Lingules actuelles et fossiles. Phoronida). Zoologica Scripta 5:133–137. Géobios 11(5):573–609. 510 Brachiopoda

Emig, C. C., & P. LeLoeuff. 1978. Description de Lin- from diverse metazoan invertebrates. Molecular gula parva Smith (Brachiopoda, Inarticulata), Marine Biology and Biotechnology 3:294–299. récoltée en Côte d’Ivoire avec quelques remarques Foster, M. W. 1974. Recent Antarctic and subantarc- sur l’écologie de l’espèce. Téthys 8(3):271–274. tic brachiopods. Antarctic Research Series 21:1– Emig, C. C., & J. A. Vargas. 1990. Notes on Glottidia 189. audebarti (Broderip) (Brachiopoda, Lingulidae) ———. 1989. Brachiopods from the extreme South from the Gulf of Nicoya, Costa Rica. Revista de Pacific and adjacent waters. Journal of Paleontology Biologia Tropical 38(2A):251–258. 63:268–301. Endo, Kazuyoshi. 1987. Life habit and relative growth Fouke, B.W. 1986. The functional significance of spi- of some laqueid brachiopods from Japan. Transac- cule-reinforced connective tissues in Terebratulina tions and Proceedings of the Palaeontological Soci- unguicula (Carpenter). In P. R. Rachebœuf & C. C. ety of Japan (new series) 147:180–194. Emig, eds., Les Brachiopodes Fossiles et Actuels, ———. 1992. Molecular systematics of recent and Actes du 1er Congrès International sur les Brach- Pleistocene brachiopods. Ph.D. thesis. University of iopodes, Brest 1985. Biostratigraphie du Paléo- Glasgow. 208 p. zoïque 4. p. 271–279, fig. 1–4. Endo, Kazuyoshi, & G. B. Curry. 1991. Molecular and François, P. 1891. Choses de Nouméa. II. Observations morphological taxonomy of a recent brachiopod biologiques sur la lingule. Archives de Zoologie genus Terebratulina. In D. I. MacKinnon, D. E. expérimentale et générale (2)9:229–245. Lee, & J. D. Campbell, eds., Brachiopods Through Franzen, Åke. 1956. On spermiogenesis, morphology Time, Proceedings of the 2nd International Bra- of the spermatozoon, and biology of fertilization chiopod Congress, University of Otago, Dunedin, among invertebrates. Zoologiska Bidrag fran New Zealand, 1990. Balkema. Rotterdam. p. 101– Uppsala 31:355–482. 108. ———. 1969. On the larval development and meta- Endo, Kazuyoshi, G. B. Curry, R. Quinn, M. J. morphosis in Terebratulina, Brachiopoda. Zoolo- Collins, G. Muyzer, & P. Westbroek. 1994. Re-in- giska Bidrag fran Uppsala 38:155–174. terpretation of terebratulide phylogeny based on ———. 1982. Ultrastructure of spermatids and sper- immunological data. Palaeontology 37:349–373, matozoa in three polychaetes with modified biology fig. 1–10. of reproduction: Autolytus sp., Chitinopoma serrula Erdmann, V. A., J. Wolters, E. Huysmans, & R. De and Capitella capitata. International Journal of In- Wachter. 1985. Collection of published 5S, 5.8S vertebrate Reproduction 5:185–200. and 4.5S ribosomal RNA sequences. Nucleic Acids ———. 1987. Spermatogenesis. In A. C. Giese, J. S. Research 13:r105–r153. Pearse, & V. B. Pearse, eds., Reproduction of Ma- Eshleman, W. P., & J. L. Wilkens. 1979a. Actinomysin rine Invertebrates. Volume 9. General aspects: seek- ATPase activities in the brachiopod Terebratalia ing unity in diversity. Blackwell Scientific Publica- transversa. Canadian Journal of Zoology 57:1944– tions. California. p. 1–47. 1949, fig. 1–3. Freeman, Gary. 1994. The endocrine pathway respon- ———. 1979b. Brachiopod orientation to current di- sible for oocyte maturation in the inarticulate bra- rection and substrate position (Terebratalia chiopod Glottidia. Biological Bulletin 186:263– transversa). Canadian Journal of Zoology 57:2079– 270. 2082, fig. 1. Frey, R. W., J. S. Hong, J. D. Howard, B. K. Park, & Eshleman, W. P., J. L. Wilkens, & M. J. Cavey. 1982. S. J. Han. 1987. Zonation of benthos on a macro- Electrophoretic and electron microscopic examina- tidal flat, Inchon, Korea. Senckenbergiana Mari- tion of the adductor and diductor muscles of an ar- tima 19(5/6):295–329. ticulate brachiopod, Terebratalia transversa. Cana- Frith, D. W., R. Tantanasiriwong, & O. Bhatia. 1976. dian Journal of Zoology 60:550–559, fig. 1–5. Zonation of macrofauna on a mangrove shore, Farris, J. S. 1972. Outgroups and parsimony. System- Phuket island. Phuket Marine Biological Center atic Zoology 31:328–334. Research Bulletin 10:1–37. Felsenstein, Joseph. 1993. PHYLIP (Phylogeny Infer- Gaspard, Danièle. 1978. Biominéralisation chez les ence Package). Computer program distributed by brachiopodes articulés—microstructure et forma- the author. Department of Genetics, University of tion de la coquille. Annales de Paléontologie 64:1– Washington. Seattle, Washington. 25. Field, Katherine, G. Olsen, D. J. Lane, S. J. ———. 1982. Microstructure de térébratules biplisées Giovannoni, M. T. Ghiselin, E. C. Raff, N. R. Pace, (Brachiopodes) du Cénomanien de la Sarthe & R. A. Raff. 1988. Molecular phylogeny of the (France). Affinités d’une des formes avec le genre animal kingdom. Science 239:748–753. Sellithyris Midd. Annales de Paléontologie Fischer, P., & D.-P. Oehlert. 1892. Brachiopodes de (Vertébrés-Invertébrés) 68(1):1–14, 3 pl. l’Atlantique Nord. Resultats des Campagnes ———. 1986. Aspects figurés de la biominéralisation Scientifiques du Prince de Monaco 3:1–30. unités de base de la sécrétion carbonatée chez les Flower, N. E., & C. R. Green. 1982. A new type of Terebratulida actuels. In P. R. Rachebœuf & C. C. gap junction in the phylum Brachiopoda. Cell and Emig, eds., Les Brachiopodes Fossiles et Actuels, Tissue Research 227:231–234. Actes du 1er Congrès International sur les Folmer, O., M. Black, W. Hoeh, R. Lutz, & R. Brachiopodes, Brest 1985. Biostratigraphie du Vrijenhoek. 1994. DNA primers for amplification Paléozoïque 4. p. 77–83. of mitochondrial cytochrome c oxidase subunit I ———. 1989. Quelques aspects de la biodégradation References 511

des coquilles de brachiopodes; conséquences sur les Orthida (Brachiopoda). Compte-Rendus de leur fossilisation. Bulletin de la Société Géologique l’Académie des Sciences de Paris 311(II):1273– de France 6:1207–1216. 1277. ———. 1990. Diagenetic modifications of shell micro- Grant, R. E. 1963. Unusual attachment of a Permian structure in the Terebratulida (Brachiopoda, linoproductid brachiopod. Journal of Paleontology Articulata) II. In J. G. Carter, ed., Skeletal Bio- 37:134–140, 1 fig., pl. 19. mineralization: Patterns, Processes and Evolution- ———. 1968. Structural adaptation in two Permian ary Trends, Volume 1. Van Nostrand Reinhold. brachiopod genera, Salt Range, West Pakistan. Jour- New York. p. 53–56. nal of Paleontology 42:1–32. ———. 1991. Growth stages in articulate brachiopod ———. 1972. The lophophore and feeding mecha- shells and their relation to biomineralization. In D. nism of the Productidina (Brachiopoda). Journal of I. MacKinnon, D. E. Lee, & J. D. Campbell, eds., Paleontology 46:213–248, pl. 1–9. Brachiopods Through Time, Proceedings of the 2nd ———. 1980. Koskinoid perforations in brachiopod International Brachiopod Congress, University of shells: function and mode of formation. Lethaia Otago, Dunedin, New Zealand, 1990. Balkema. 13:313–319. Rotterdam. p. 167–174. ———. 1987. Brachiopods of Enewetak Atoll. In D. Gee, Henry. 1995. Lophophorates prove likewise vari- M. Devaney & others, eds., The Natural History of able. Nature 374:493. Enewetak Atoll, vol. 2. Office of Scientific and George, T. N. 1932a. Ambocoelia Hall and certain Technical Information, U. S. Department of En- similar British Spiriferidae. Geological Society of ergy. Oak Ridge, Tennessee. p. 77–84. London, Quarterly Journal 87:30–61, pl. 3–5. Gratiolet, M. P. 1860. Études anatomiques sur la ———. 1932b. The British Carboniferous reticulate Lingule anatine (L. anatina Lam.). Journal de Con- Spiriferidae. Geological Society of London, Quar- chyliologie 4:49–172. terly Journal 88:516–575, pl. 31–35. Gustus, R. M., & R. A. Cloney. 1972. Ultrastructural Ghiselin, M. T. 1988. The origin of molluscs in the similarities between setae of brachiopods and poly- light of molecular evidence. In P. H. Harvey & L. chaetes. Acta Zoologica 53:229–233. Partridge, eds., Oxford Surveys in Evolutionary Bi- Halanych, K. M. 1991. 5S ribosomal RNA sequences ology. Oxford University Press. Oxford. p. 66–95. inappropriate for phylogenetic reconstruction. Mo- Giese, A. C., J. S. Pearse, & V. B. Pearse. 1987. Repro- lecular Biology and Evolution 8:249–253. duction of Marine Invertebrates. Volume 9. General Halanych, K. M., J. D. Bacheller, A. M. A. Aguinaldo, aspects: seeking unity in diversity. Blackwell Scien- S. M. Liva, D. M. Hillis, & J. A. Lake. 1995. Evi- tific Publications. California. 712 p. dence from 18S ribosomal DNA that the lopho- Gilmour, T. H. J. 1978. Ciliation and function of the phorates are protostome animals. Science food-collecting and waste-rejecting organs of the 267:1641–1643. lophophorates. Canadian Journal of Zoology Hall, J., & J. M. Clarke. 1892–1894. An introduction 56:2142–2155. to the study of the genera of Palaeozoic ———. 1981. Food-collecting and waste-rejecting Brachiopoda. New York Geological Survey 8. Pt. mechanisms in Glottidia pyramidata and the persis- 1(1892):i–xvi, 1–367, pl. 1–20; Pt. 2: i–xvi, 1–394, tence of lingulacean brachiopods in the fossil pl. 21–84. record. Canadian Journal of Zoology 59:1539– ———. 1894–1895. An introduction to the study of 1547. the Brachiopoda intended as a Handbook for the Golding, G. B., & R. S. Gupta. 1994. Protein-based use of Students. New York State Geologist Annual phylogenies support a chimeric origin for the eu- Report for 1891 (1894):134–300, pl. 1–22; con- karyotic genome. Molecular Biology and Evolution tinuation (pt. 2) New York State Geologist Annual 12:1–6. Report for 1893 (1894) [1895]:751–943, pl. 23– Gorjansky, V. Iu. [see also Gorjansky, W. Ju.], & L. E. 54. Popov. 1985. Morphologiia, systematicheskoe Hammen, C. S. 1968. Aminotransferase activities and polozhenie I proiskhozhdenie bezzamkovykh amino acid excretion of bivalve molluscs and brach- brakhiopod s karbonatnoi rakovinoi [Morphology, iopods. Comparative Biochemistry and Physiology systematic position and origin of inarticulate brach- 26:697–705. iopods with a carbonate shell]. Paleontologicheskii ———. 1969. Lactate and succinate oxidoreductases Zhurnal 1985(3):3–14, 5 fig., pl. 1. in marine invertebrates. Marine Biology 4:233– Gorjansky, W. Ju. [see also Gorjansky, V. Iu.], & L. Ye. 238. Popov. 1986. On the origin and systematic position ———. 1971. Metabolism of brachiopods and bivalve of the calcareous-shelled inarticulate brachiopods. molluscs. In R. Alvardo, E. Gadea, & A. de Haro, Lethaia 19:233–240, fig. 1–3. eds., Actas del I Symposio International de Gourvennec, Rémy. 1987. Morphologie des épines Zoofilogenia, Salamanca, 1969, Acta Salman- chez les brachiopodes Delthyrididae. Lethaia ticensia, Ciencias 36:471–478. 20:21–31. ———. 1977. Brachiopod metabolism and enzymes. ———. 1989. Radial microornament in spiriferid American Zoologist 17:141–147. brachiopods and paleogeographical implications. Hammen, C. S., & R. C. Bullock. 1991. Opine oxi- Lethaia 22:405–411. doreductases in brachiopods, bryozoans, phoronids Gourvennec, Rémy, & Michel Mélou. 1990. and molluscs. Biochemical Systematics and Ecology Découverte d’un cas d’ornamentation épineuse chez 19:263–269. 512 Brachiopoda

Hammen, C. S., D. P. Hanlon, & S. C. Lum. 1962. Ordovician sequence of Bohemia. Sbornik Geo- Oxidative metabolism of Lingula. Comparative logickych ved (Paleontologie) 25:9–82, 16 fig., pl. Biochemistry and Physiology 5:185–192. 1–16. Hammen, C. S., & S. C. Lum. 1966. Fumarate reduc- Hay-Schmidt, A. 1992. Ultrastructure and immunocy- tase and succinate dehydrogenase activities in bi- tochemistry of the nervous system of the larvae of valve mollusks and brachiopods. Comparative Bio- Lingula anatina and Glottidia sp. (Brachiopoda). chemistry and Physiology 19:775–781. Zoomorphology 112:189–205. ———. 1977. Salinity tolerance and pedicle regenera- Heinrikson, R. L., & S. C. Meredith. 1984. Amino tion of Lingula. Journal of Paleontology 51:548– acid analysis by reverse-phase high-performance liq- 551. uid chromatography: precolumn derivatization with Hammen, C. S., & P. J. Osborne. 1959. Carbon diox- phenylisothiocyanate. Analytical Biochemistry ide fixation in marine invertebrates: a survey of 136:65–74. major phyla. Science 130:1409–1410. Heller, M. 1931. Exkretorische Tätigkeitder Brach- Hammond, L. S. 1980. The larvae of a discinid iopoden. Zeitschrift für Morphologie und Oeko- (Brachiopoda, Inarticulata) from inshore waters logie der Tiere 24:238–258. near Townsville, Australia, with revised identifica- Helmcke, J. G. 1940. Die Brachiopoden der deutschen tions of previous records. Journal of Natural His- Tiefsee-Expedition. Wissenschaftliche Ergebnisse tory 14:647–661. der deutschen Tiefsee-Expedition auf dem Dampfer ———. 1982. Breeding season, larval development “Valdivia” 1898–1899. G. Fischer Verlag, 1932– and dispersal of Lingula anatina (Brachiopoda, 1940. Jena. Wissenschaftliche Ergebnissen der inarticulata) from Townsville, Australia. Journal of deutschen Tiefsee-Expedition 24(3):217–316. Zoology, London 198:183–196. Hemmingsen, A. M. 1960. Energy metabolism as re- ———. 1983. Experimental studies of salinity toler- lated to body size and respiratory surfaces and its ance, burrowing behavior and pedicle regeneration evolution. Report to the Steno Memorial Hospital in Lingula anatina (Brachiopoda, Inarticulata). 9:7–110. Journal of Paleontology 57:1311–1316. Hennig, Willi. 1966. Phylogenetic Systematics. Uni- ———. 1984. Epibiota from the valves of recent Lin- versity of Illinois Press. Urbana. 263 p. gula (Brachiopoda). Journal of Paleontology Hertlein, L. G., & U. S. Grant IV. 1944. The Ceno- 58:1528–1531. zoic Brachiopoda of western North America. Pub- Hammond, L. S., & I. R. Poiner. 1984. Genetic struc- lications of the University of California at Los An- ture of three populations of the “living fossil” geles in Mathematical and Physical Sciences 3:236 brachiopod Lingula from Queensland, Australia. p., pl. 1–21. Lethaia 17:139–143. Hiller, Norton. 1988. The development of growth Hancock, A. 1859. On the organisation of the lines on articulate brachiopods. Lethaia 21:177– Brachiopoda. Philosophical Transactions of the 188. Royal Society 148:791–869. ———. 1991. The southern African recent brachiopod Hare, P. E. 1962. The amino acid composition of the fauna. In D. I. MacKinnon, D. E. Lee, & J. D. organic matrix of some recent and fossil shells of Campbell, eds., Brachiopods Through Time, Pro- some west coast species of Mytilus. Ph.D. thesis. ceedings of the 2nd International Brachiopod Con- California Institute of Technology. gress, University of Otago, Dunedin, New Zealand, Hare, P. E., & R. M. Mitterer. 1969. Laboratory simu- 1990. Balkema. Rotterdam. p. 439–445. lation of amino acid diagenesis in fossils. Carnegie ———. 1993. A modern analogue of the Lower Or- Institute of Washington Yearbook 67:205–208. dovician Obolus conglomerate of Estonia. Geologi- Haro, A. de. 1963. Estructura y anatomia comparadas cal Magazine 130(2):265–267. des las gonadas y pedunculo des los Braquiópodos Hillis, D. M., & M. T. Dixon. 1991. Ribosomal DNA: testicardinos. Publicaciones del Instituto de molecular evolution and phylogenetic inference. Biología Applicada, Barcelona 35:97–117. Quarterly Review of Biology 66:411–453. Harvey, E. N. 1928. The oxygen consumption of lumi- Hillis, D. M., J. P. Huelsenbeck, & C. W. Cunning- nous bacteria. Journal of General Physiology ham. 1994. Application and accuracy of molecular 11:469–475. phylogenies. Science 264:671–677. Hatai, K. M. 1940. The Cenozoic Brachiopoda of Ja- Hillis, D. M., & D. Moritz. 1990. Molecular System- pan. Science Reports of the Tohoku Imperial Uni- atics. Sinauer Associates, Inc. Sunderland, Massa- versity, Sendai, Japan (second series, Geology) chusetts. 588 p. 20:1–413, pl. 1–12. Hirai, E., & T. Fukushi. 1960. The development of Haugen, J.-E., H.-P. Sejrup, & N. B. Vogt. 1989. two species of lamp shells, Terebratalia coreanica and Chemotaxonomy of Quaternary benthic foramin- Coptothyris grayi. Bulletin of the Ashamushi Marine ifera using amino acids. Journal of Foraminiferal Biological Station 10:77–80. Research 19:38–51. Hochachka, P. W., & G. H. Somero. 1973. Strategies Havlicék,ˇ Vladimir. 1961. Rhynchonelloidea des of Biochemical Adaptation. W. B. Saunders Com- Boehmischen aelteren palaeozoikums (Brach- pany. Philadelphia. 358 p. iopoda). Ustredni Ustav Geologicky Rozpravy ———. 1984. Biochemical Adaptation. Princeton 27:211 p., 27 pl. University Press. Princeton. 480 p. ———. 1982. Lingulacea, Paterinacea and Holland, P. W. H. 1992. Homeobox genes in verte- Siphonotretacea (Brachiopoda) in the Lower brate evolution. BioEssays 14:267–273. References 513

Holland, P. W. H., & B. L. M. Hogan. 1986. Phylo- Mathematics Department, University of Bielefeld, genetic distribution of Antennapedia-like Germany. Bielefeld. homeoboxes. Nature 321:251–253. Huxley, J. S. 1932. Problems of Relative Growth. The Holland, P. W. H., N. A. Williams, & J. Lanfear. 1991. Dial Press. New York. 244 p. Cloning of segment polarity gene homologues from Hyman, L. H. 1959a. The Invertebrates: Smaller Co- the unsegmented brachiopod Terebratulina retusa elomate Groups. Chaetognatha, Hemichordata, (Linnaeus). Federation of European Biochemical Pogonophora, Phoronida, Ectoprocta, Brachiopoda, Societies Letters 291:211–213. Sipunculida, the Coelomate Bilateria, Vol. 5. Holmer, L. E. 1989. Middle Ordovician phosphatic McGraw-Hill. New York, London, Toronto. viii + inarticulate brachiopods from Västergötland and 783 p., fig. 1–240. Dalarna, Sweden. Fossils and Strata 26:1–172, 118 ———. 1959b. The Lophophorate Coelomates—Phy- fig. lum Brachiopoda. In L. H. Hyman, The Inverte- Holmer, L. E., & L. Ye. Popov. 1990. The Acrotre- brates: Smaller Coelomate Groups. McGraw-Hill. tacean brachiopod Ceratreta tanneri (Metzger) from New York, London, Toronto. p. 228–609. the Upper Cambrian of Baltoscandia. Geologiska Iijima, M., T. Hiroko, M. Yutaka, & K. Yoshinori. Foreningens i Stockholm Forhandlingar 1991. Difference of the organic component be- 112(3):249–263. tween the mineralized and the non-mineralized lay- d’Hondt, Jean-Loup. 1986. Étude de l’intestin et de la ers of Lingula shell. Comparative Biochemistry and glande digestive de Terebratulina retusa (L.) Physiology 98A:379–382. (Brachiopode). IV. Comparaison avec les activités Iijima, M., H. Kamemizu, N. Wakamatsu, T. Goto, & enzymatiques d’autres brachiopodes du même Y. Moriwaki. 1991. Thermal decomposition of Lin- biotope. In P. R. Rachebœuf & C. C. Emig, eds., gula shell apatite. Calcified Tissue International Les Brachiopodes Fossiles et Actuels, Actes du 1er 49:128–133. Congrès International sur les Brachiopodes, Brest Iijima, M., & Y. Moriwaki. 1990. Orientation of apa- 1985. Biostratigraphie du Paléozoïque 4. p. 301– tite and organic matrix in Lingula unguis shell. Cal- 305. cified Tissue International 47:237–242. d’Hondt, Jean-Loup, & E. Boucaud-Camou. 1982. Iijima, M., Y. Moriwaki, Y. Doi, & Y. Kuboki. 1988. Étude l’intestin et de la glande digestive de la The orientation of apatite crystals in Lingula unguis Terebratulina retusa (L.) (Brachiopode). Ultrastruc- shell. Japanese Journal of Oral Biology 30:20–30. ture et recherche d’activités amylasiques et Iijima, M., Y. Moriwaki, & Y. Kuboki. 1991. Orienta- protéasiques. Bulletin de la Société Zoologique de tion of apatite and the organic matrix in Lingula France 107(2):207–216. shells, p. 433–437. In S. Suga & H. Nakahara, eds., Hori, H., B. L. Lim, T. Ohama, T. Kumazaki, & S. Mechanisms and Phylogeny of Mineralization of Osawa. 1985. Evolution of organisms deduced Biological Systems. Springer-Verlag. Tokyo. from 5S rRNA sequences, p. 369–384. In T. Ohta Ikeda, T. 1974. Nutritional ecology of marine zoo- & A. Aoki, eds., Population Genetics and Molecu- plankton. Memoires of the Faculty of Fisheries, lar Evolution. Springer. Berlin. 503 p. Hokkaido University 21:19–112. Hori, H., & S. Osawa. 1986. Evolutionary change in Irwin, D. H. 1991. Metazoan phylogeny and the Cam- 5S rRNA secondary structure and a phylogenetic brian radiation. Trends in Ecology and Evolution tree of 352 5S rRNA species. BioSystems 19:163– 6:131–134. 172. Ishikawa, Hajime. 1977. Comparative studies on the Hoverd, W. A. 1985. Histological and ultrastructural thermal stability of animal ribosomal RNAs: V. observations of the lophophore and larvae of the Tentaculata (Phoronids, moss-animals and lamp- brachiopod, Notosaria nigricans (Sowerby, 1846). shells). Comparative Biochemistry and Physiology Journal of Natural History 19:831–850. 57B:9–14. ———. 1986. The adult lophophore of Notosaria Ivanov, C. P., & R. Z. Stoyanova. 1972. Aliphatic hy- nigricans (Brachiopoda). In P. R. Rachebœuf & C. drocarbons in fossils of Mesozoic belemnites. C. Emig, eds., Les Brachiopodes Fossiles et Actuels, Comptes-Rendus de l’Académie Bulgare des Sci- Actes du 1er Congrès International sur les ences 25(5):637–639. Brachiopodes, Brest 1985. Biostratigraphie du Ivanov, C. P., R. Z. Stoyanova, & C. P. Daskalov. 1975. Paléozoïque 4. p. 307–312. Content and composition of higher fatty acids in Huelsenbeck, J. P. 1995. Performance of phylogenetic fossils of Mesozoic belemnites. Comptes-Rendus de methods in simulation. Systematic Biology 44:17– l’Académie Bulgare des Sciences 28(7):935–938. 48. Iwata, K. 1981. Ultrastructure and mineralization of Hughes, W. W., G. D. Rosenberg, & R. D. Tkachuck. the shell of Lingula unguis Linné (Inarticulate Bra- 1988. Growth increments in the shell of the living chiopod). Journal of the Faculty of Science of brachiopod Terebratalia transversa. Marine Biology Hokkaido University (series IV) 20(1):35–65. 98:511–518. ———. 1982. Ultrastructure and calcification of the Hulmes, A. G., & G. A. Boxshall. 1988. Peocilostome shells in inarticulate brachiopods Part 2. Ultrastruc- copepods associated with bivalve molluscs and a ture of the shells of Glottidia and Discinisca. Jour- brachiopod at Hong-Kong. Journal of Natural His- nal of the Geological Society of Japan 88:957–966, tory 22(2):537–544. 5 pl. Huson, D. H., & R. Wetzel. 1994. SplitsTree 1.0: Jaanusson, Valdar. 1971. Evolution of the brachiopod Computer program distributed by the authors. hinge. In J. T. Dutro, Jr., ed., Paleozoic Perspectives: 514 Brachiopoda

A Paleontological Tribute to G. Arthur Cooper. acid composition and disc electrophoresis of fossil Smithsonian Contributions to Paleobiology 3:33– articulate shell protein. Comparative Biochemistry 46, 6 fig., pl. 1–2. and Physiology 30:225–232. Jackson, J. B., T. F. Goreau, & W. D. Hartman. 1971. ———. 1971. Constituents of brachiopod shells. In Recent brachiopod-coralline sponge communities M. Florkin & E. H. Stotz, eds., Comprehensive and their paleoecological significance. Science Biochemistry. Elsevier. Amsterdam. p. 749–784. 173:623–625. ———. 1973. The protein of brachiopod shell. V. N- Jacobs, H. T., P. Balfe, B. L. Cohen, A. Farquharson, & terminal end groups. Comparative Biochemistry L. Comito. 1988. Phylogenetic implications of ge- and Physiology 45B:17–24. nome rearrangement and sequence evolution in ———. 1977. Brachiopod shell proteins: their func- echinoderm mitochondrial DNA. In C. R. C. Paul tion and taxonomic significance. American Zoolo- & A. B. Smith, eds., Echinoderm Phylogeny and gist 17:133–140. Evolutionary Biology. Clarendon Press. Oxford. p. ———. 1979. The protein of brachiopod shell. VI. C- 121–137. terminal end groups and sodium dodecylsulphate- Jaeger, J. A., D. H. Turner, & M. Zuker. 1989a. Im- polyacrylamide gel electrophoresis: molecular con- proved predictions of secondary structure for RNA. stitution and structure of the protein. Comparative Proceedings of the National Academy of Sciences, Biochemistry and Physiology 63B:163–173. USA 86:7706–7710. ———. 1980. Phylogenetic information from fossil ———. 1989b. Predicting optimal and suboptimal brachiopods. In P. E. Hare, T. C. Hoering, & K. J. secondary structure for RNA. Methods in Enzy- King, eds., Biogeochemistry of Amino Acids. John mology 183:281–306. Wiley & Sons. New York. p. 83–94. James, M. A., A. D. Ansell, & G. B. Curry. 1991a. Re- ———. 1986. Evolution of the Brachiopoda: the mo- productive cycle of the brachiopod Terebratulina lecular approach. In P. R. Rachebœuf & C. C. retusa on the west coast of Scotland. Marine Biol- Emig, eds., Les Brachiopodes Fossiles et Actuels, ogy 109:441–451. Actes du 1er Congrès International sur les Brach- ———. 1991b. Functional morphology of the gonads iopodes, Brest 1985. Biostratigraphie du Paléo- of the articulate brachiopod Terebratulina retusa. zoïque 4. p. 103–111. Marine Biology 111:401–410. Jørgensen, C. B. 1981. A hydromechanical principle ———. 1991c. Oogenesis in the articulate brachiopod for particle retention in Mytilus edulis and other Terebratulina retusa. Marine Biology 111:411–423. ciliary suspension feeders. Marine Biology 61:277– James, M. A., A. D. Ansell, M. J. Collins, G. B. Curry, 282. L. S. Peck, & M. C. Rhodes. 1992. Biology of liv- Jørgensen, C. B., T. Kiørboe, F. Mohlenberg, & H. U. ing brachiopods. Advances in Marine Biology Riisgard. 1984. Ciliary and mucus-net filter feed- 28:175–387. ing, with special reference to fluid mechanical char- Jin, Yu-gan, & Hua-yu Wang. 1992. Revision of the acteristics. Marine Ecology Progress Series 15:283– Lower Cambrian brachiopod Heliomedusa Sun & 292. Hou, 1987. Lethaia 25(1):35–49, fig. 1–14. Joshi, J. G., & B. Sullivan. 1973. Isolation and pre- Johnson, J. G., & P. Westbroek. 1971. Cardinalia ter- liminary characterisation of haemerythrin for Lin- minology in rhynchonellid brachiopods. Geological gula unguis. Comparative Biochemistry and Physi- Society of America Bulletin 82:1699–1702. ology 44B:857–867. Jones, G. J., & J. L. Barnard. 1963. The distribution Joubin, L. 1886. Recherches sur l’anatomie des and abundance of the inarticulate brachiopod brachiopodes inarticulés. Archives de Zoologie Glottidia albida (Hinds) on the mainland of south- Expérimentale et Générale (série 2) 4:161–303. ern California. Pacific Naturalist 4(2):27–52. Jux, U., & F. Strauch. 1966. Die mitteldevonische Jope, H. M. 1965. Composition of brachiopod shell. Brachiopoden-Gattung Uncites de France 1825. In R. C. Moore, ed., Treatise on Invertebrate Pale- Palaeontographica 56:176–222. ontology. Part H, Brachiopoda. Geological Society Källersjö, M., J. S. Farris, A. G. Kluge, & C. Bult. of America & University of Kansas Press. New York 1992. Skewness and permutation. Cladistics 8:275– & Lawrence. p. 156–164. 287. ———. 1967a. The protein of brachiopod shell. I. Karlin, S., & A. M. Campbell. 1994. Which bacterium Amino acid composition and implied protein tax- is the ancestor of the animal mitochondrial ge- onomy. Comparative Biochemistry and Physiology nome? Proceedings of the National Academy of 20:593–600. Sciences, USA 91:12842–12846. ———. 1967b. The protein of brachiopod shell. II. Kawaguti, S. 1941. Hemerythrin found in the blood of Shell protein from fossil articulates: amino acid Lingula. Memoirs of the Faculty of Science and composition. Comparative Biochemistry and Physi- Agriculture, Taihoku Imperial University, Formosa ology 20:601–605. (Zoology, Number 12) 23:95–98. ———. 1969a. The protein of brachiopod shell. III. ———. 1943. A biometrical study on Lingula unguis Comparison with structural protein of soft tissue. (Linné). Venus 12(3/4):171–182. Comparative Biochemistry and Physiology 30:209– 224. In Japanese. ———. 1969b. The protein of the brachiopod shell. Kelly, P. G., P. T. P. Oliver, & F. G. E. Pautard. 1965. IV. Shell protein from fossil inarticulates: amino The shell of Lingula unguis. In Proceedings of the References 515

Second European Symposium on Calcified Tissue. quelques spirifères Dévoniens de la Chaine Liège, Belgium. p. 337–345. Cantabrique (Espagne). Leidse Geologische Kemezys, K. J. 1965. Significance of punctae and Mededelingen 33:73–148, 71 fig., pl. 1–16. pustules in brachiopods. Geological Magazine Kuga, Hiroto, & Akira Matsuno. 1988. Ultrastructural 102:315–324. investigations on the anterior adductor muscle of a ———. 1968. Arrangements of costellae, setae and brachiopod, Lingula unguis. Cell Structure and vascula in enteletacean brachiopods. Journal of Pa- Function 13:271–279. leontology 42:88–93, 1 fig. Kume, M. 1956. The spawning of Lingula. Natural Kenchington, R. A., & L. S. Hammond. 1978. Popu- Science Report of Ochanomizu University, Tokyo lation structure, growth and distribution of Lingula 6:215–223. anatina (Brachiopoda) in Queensland, Australia. Kume, M., & K. Dan. 1968. Invertebrate Embryology. Journal of Zoology 184:63–81. Nolit Publishing House. Belgrade. 605 p. Kimble, J. 1994. An ancient molecular mechanism for LaBarbera, Michael. 1977. Brachiopod orientation to establishing embryonic polarity? Science 266:577– water movement. I. Theory, laboratory behavior 578. and field observation. Paleobiology 3:270–287. King, K., Jr., & P. E. Hare. 1972. Amino acid compo- ———. 1978. Brachiopod orientation to water move- sition of the test as a taxonomic character for living ment: functional morphology. Lethaia 11:67–79. and fossil planktonic foraminifera. Micropaleontol- ———. 1981. Water flow patterns in and around three ogy 18:285–293. species of articulate brachiopods. Journal of Experi- Kirtisinghe, P. 1949. Unicellular algae in association mental Marine Biology and Ecology 55:185–206. with invertebrates. Nature 164:970. ———. 1984. Feeding currents and particle capture Kleiber, M. 1947. Body size and metabolic rate. Physi- mechanisms in suspension feeding animals. Ameri- ological Reviews 27:511–541. can Zoologist 24:71–84. ———. 1965. Metabolic body size. In K. L. Blaxter, ———. 1985. Mechanisms of spatial competition of ed., Energy Metabolism. Proceedings of the 3rd Discinisca strigata (Inarticulata: Brachiopoda) in the Symposium, Troon, Scotland. Academic Press. Lon- intertidal of Panama. Biological Bulletin don. p. 427–435. 168(1):91–105. Klippenstein, G. L. 1980. Structural aspects of ———. 1986a. The evolution and ecology of body haemerythrin and myohaemerythrin. American size. In D. M. Raup & D. Jablonski, eds., Patterns Zoologist 20:39–51. and Processes of Life, Dahlem Conferenzen. Kniprath, E. 1975. Das Wachstum des Mantels von Springer-Verlag. Berlin, Heidelberg. p. 69–98. Lymnaea stagnalis (Gastropoda). Cytobiologie ———. 1986b. Brachiopod lophophores: functional 10:260–267. diversity and scaling. In P. R. Rachebœuf & C. C. Knowlton, N. 1993. Sibling species in the sea. Annual Emig, eds., Les Brachiopodes Fossiles et Actuels, Review of Ecology and Systematics 24:189–216. Actes du 1er Congrès International sur les Brach- Kolesnikov, C. M., & E. L. Prosorovskaya. 1986. Bio- iopodes, Brest 1985. Biostratigraphie du Paléo- chemical investigation of Jurassic and recent brach- zoïque 4. p. 313–321. iopod shells. In P. R. Rachebœuf & C. C. Emig, ———. 1989. Analyzing body size as a factor in ecol- eds., Les Brachiopodes Fossiles et Actuels, Actes du ogy and evolution. Annual Review of Ecology and 1er Congrès International sur les Brachiopodes, Systematics 20:97–117. Brest 1985. Biostratigraphie du Paléozoïque 4. p. ———. 1990. Principles of design of fluid transport 113–119. systems in zoology. Science 249:992–1000. Komiya, H., N. Shimizu, M. Kawakami, & S. Lacaze-Duthiers, F. J. H. de. 1861. Histoire naturelle Takemura. 1980. Nucleotide sequence of 5S riboso- des Brachiopodes vivantes de la Méditerranée. Iie mal RNA from Lingula anatina. Journal of Bio- Monographie. Histoire naturelle de la Thécide chemistry 88:1449–1456. (Thecidium mediterraneum). Annales des sciences Kovalevskiy, A. O. 1874. Nablyudeniya Nad Razvitiem naturelles Zoologie (série 4) 15:259–330. Brachiopoda [On the development of the Brach- Lankester, E. R. 1873. Summary of zoological observa- iopoda]. Izvestiya Obschestvo Liubiteley Est- tions made at Naples in the winter of 1871–1872. estvoznaniya, Anthropologii i Etnografii [Proceed- Annals and Magazine of Natural History (series 4) ings of the Imperial Society for Natural Science, 11:81. Anthropology, and Ethnology, Moscow] 14:1–40, Laurie, John. 1987. The musculature and vascular sys- pl. 1–5. tems of two species of Cambrian Paterinide (Brachiopoda). Bureau of Mineral Resources Jour- In Russian. nal of Australian Geology and Geophysics 10:261– ———. 1883. Observations sur le développement des 265. brachiopodes, Analyse par Oehlert et Deniker. Ar- Law, R. H., & C. W. Thayer. 1991. Articulate fecun- chives de Zoologie Expérimentale et Générale (se- dity in the Phanerozoic: Steady state or what? In D. ries 2) 1:57–76. I. MacKinnon, D. E. Lee, & J. D. Campbell, eds., Kozlowski, Roman. 1929. Les brachiopodes goth- Brachiopods Through Time, Proceedings of the 2nd landiens de la Podolie polonaise. Palaeontologia International Brachiopod Congress, University of Polonica 1:1–254, pl. 1–12. Otago, Dunedin, New Zealand, 1990. Balkema. Krans, Th. F. 1965. Études morphologiques de Rotterdam. p. 183–190. 516 Brachiopoda

Lee, D. E. 1978. Aspects of the ecology and paleoecol- ultrastructure. Neues Jahrbuch für Geologie und ogy of the brachiopod Notosaria nigricans Palaeontologie, Monatshefte 3:133–148. (Sowerby). Journal of the Royal Society of New Lum, S. C., & C. S. Hammen. 1964. Ammonia excre- Zealand 8:395–417. tion in Lingula. Comparative Biochemistry and ———. 1987. Cenozoic and recent inarticulate bra- Physiology 12:185–190. chiopods of New Zealand: Discina, Pelagodiscus and Lutz, R. A., & D. C. Rhoads. 1977. Anaerobiosis and Neocrania. Journal of the Royal Society of New a theory of growth line formation. Science Zealand 17(1):49–72. 198:1222–1227. LeGeros, R. Z., C.-M. Pan, S. Suga, & Norimitsu Mackay, Sarah, & R. A. Hewitt. 1978. Ultrastructural Watabe. 1985. Crystallo-chemical properties of studies on the brachiopod pedicle. Lethaia 11: 331– apatite in atremate brachiopod shells. Calcified Tis- 339. sue International 37:98–100. Mackay, Sarah, D. I. MacKinnon, & Alwyn Williams. Leroy, P. 1936. “Lingula anatina” Lamarck (1809) 1994. Ultrastructure of the loop of terebratulide dans les mers froides de Chine. Bulletin de la brachiopods. Lethaia 26:367–378. Société des Sciences de Nancy 5:121–124. MacKinnon, D. I. 1971. Perforate canopies to canals Livingstone, A. 1983. Invertebrate and vertebrate in the shells of fossil Brachiopoda. Lethaia 4:321– pathways of anaerobic metabolism: evolutionary 325. considerations. Journal of the Geological Society of ———. 1974. The shell structure of spiriferide Brach- London 140:27–37. iopoda. British Museum (Natural History) Bulletin Livingstone, D. R., A. De Zwann, M. Leopold, & E. (Geology) 25(3):189–261, 27 fig., pl. 1–32. Marteljn. 1983. Studies on the phylogenetic distri- ———. 1977. The formation of muscle scars in articu- bution of pyruvate oxidoreductases. Biochemical late brachiopods. Philosophical Transactions of the Systematics and Ecology 11:415–425. Royal Society of London (series B, Biological Sci- Lockhart, P. J., M. A. Steel, M. D. Hendy, & D. ences) 280(970):1–27, 86 fig., pl. 1–11. Penney. 1994. Recovering evolutionary trees under ———. 1991. A fresh look at growth of spiral a more realistic model of sequence evolution. Mo- brachidia. In D. I. MacKinnon, D. E. Lee, & J. D. lecular Biology and Evolution 11:605–612. Campbell, eds., Brachiopods Through Time, Pro- Logan, Alan. 1979. The recent Brachiopoda of the ceedings of the 2nd International Brachiopod Con- Mediterranean Sea. Bulletin de l’Institut Océano- gress, University of Otago, Dunedin, New Zealand, graphique de Monaco 72(1434):1–112, 10 pl. 1990. Balkema. Rotterdam. p. 147–153, 6 fig. ———. 1983. Brachiopoda collected by CANCAP 1- MacKinnon, D. I., & Alwyn Williams. 1974. Shell 111 expeditions to the south-east North Atlantic. structure of terebratulid brachiopods. Palaeontology 1976–1978. Zoologische Mededelingen 57:165– 17(1):179–202, 3 fig., pl. 21–27. 189, 1 pl. Macomber, R. W., & Lenore Macomber. 1983. Rib- ———. 1988. A new thecideid genus and species bing patterns in the brachiopod Diceromyonia. (Brachiopoda, recent) from the southeast North Lethaia 16:25–37. Atlantic. Journal of Paleontology 62:546–551. Maddison, W. P., M. J. Donoghue, & D. R. Mad- ———. 1990. Recent Brachiopoda from the Snellius dison. 1984. Outgroup analysis and parsimony. and Luymes expeditions to the Surinam-Guyana Systematic Zoology 33:83–103. Shelf, Bonaire-Curacao, and Saba Bank, Caribbean Mahajan, S. N., & M. C. Joshi. 1983. Age and shell Sea, 1966 and 1969–1972. Zoologische Medede- growth in Lingula anatina (Lam.). Indian Journal of lingen 63:123–136. Marine Sciences 12:120–121. ———. 1993. Recent brachiopods from the Canarian- Maidak, B. L., N. Larsen, M. J. McCaughey, R. Over- Cape Verdean region: diversity, biogeographic beek, G. L. Olsen, K. Fogel, J. Blandy, & C. R. affinities, bathymetric range and life habits. Courier Woese. 1994. The ribosomal database project. Forschungs-Institut Senckenberg, Frankfurt-am- Nucleic Acids Research 22:3485–3487. Main 159:229–233. Malakhov, V. V. 1976. Certain stages of embryogenesis Logan, Alan, J. P. A. Noble, & G. R. Webb. 1975. An in Cnismatocentrum sakhaliensis parvum (Brach- unusual attachment of a recent brachiopod, Bay of iopoda, Testicardines) and the problem of evolution Fundy, Canada. Journal of Paleontology 49:557– of the way of origin of coelomic mesoderm. Zoo- 558. logicheski Zhurnal 55:66–75. Long, J. A. 1964. The embryology of three species rep- Manankov, I. N. 1979. Pseudopunctae in stropho- resenting three superfamilies of articulate Brach- menids. Paleontological Journal 3:332–338. iopoda. Unpublished Ph.D. thesis. University of Washington. Seattle. 184 p. Translation of O psevdoporakh strofomenid. Long, J. A., & S. A. Stricker. 1991. Brachiopoda. In A. Paleontologicheskii Zhurnal 3(13):72–78. Geise, J. S. Pearse, & V. B. Pearse, eds., Reproduc- Mancenido, M. O., & Miguel Griffin. 1988. Distribu- tion of Marine Invertebrates, Vol. 6. Blackwell Sci- tion and palaeoenvironmental significance of the entific. California. p. 47–84. genus Bouchardia (Brachiopoda, Terebratellidina): Longhurst, A. R. 1958. An ecological survey of the its bearing on the Cenozoic evolution of the South west African marine benthos. Colonial Office Fish- Atlantic. Revista Brasileira de Geociencias ery Publications 11:1–102. 18(2):201–211, pl. 1. Ludvigsen, R. 1974. A new Devonian acrotretid Mano, R. 1960. On the metamorphosis of the brachio- (Brachiopoda, Inarticulata) with unique protegular pod Frenulina sanguinolenta (Gmelin). Bulletin of References 517

the Marine Biological Station of Asamushi 10:171– Mileikovsky, S. A. 1970. Distribution of pelagic larvae 175. of bottom invertebrates in Kurile-Kamtchatka area. Marshall, A. G., & L. Medway. 1976. A mangrove In V. G. Bogorov, ed., Fauna of the Kurile- community in the New Hebrides, southwest Pa- Kamtchatka Trench and Its Environment. Nauka. cific. Biological Journal of the Linnean Society Moscow. p. 124–143. 8:319–336. Martinez-Chacon, M. L., & J. L. Garcia-Alcalde. In Russian, translated in English. 1978. La genesis del koskinoide en braquiopodos Mineur, R. J., & J. R. Richardson. 1984. Free and articulados. Revista de la Facultad de Ciencias, mobile brachiopods from New Zealand Oligocene Universidad de Oviedo 17–19:261–279. deposits and Australian waters. Alcheringa 8(3– Mattox, M. T. 1955. Observations on the brachiopod 4):327–334, fig. 1–5. communities near Santa Catalina. In Essays in the Mohlenberg, F., & H. U. Riisgard. 1978. Efficiency of Natural Sciences in Honor of Captain Allan Han- particle retention in 13 species of suspension feed- cock. University of Southern California Press. Los ing bivalves. Ophelia 14:239–246. Angeles. p. 73–86. Moore, R. C., ed. 1965. Treatise on Invertebrate Pale- Mauzey, K. P., C. Birkeland, & P. K. Dayton. 1968. ontology. Part H, Brachiopoda. The Geological Feeding behaviour of asteroids and escape responses Society of America & The University of Kansas of their prey in the Puget Sound region. Ecology Press. New York & Lawrence. xxxii + 927 p., 746 49:603–619. fig. Mayzaud, P. 1973. Respiration and nitrogen excretion Morse, E. S. 1873. Embryology of Terebratulina. of zooplankton II. Studies of the metabolic charac- Memoirs of the Boston Society of Natural History teristics of starved animals. Marine Biology 21:19– 2:249–264, pl. 8–9. 28. ———. 1902. Observations on living Brachiopoda. McCammon, H. M. 1965. Filtering currents in brach- Memoirs of the Boston Society of Natural History iopods measured with a thermistor flowmeter. 5(8):313–386. Ocean Science Engineering 2:772–779. Morton, J. E. 1960. The functions of the gut in ciliary ———. 1969. The food of articulate brachiopods. feeders. Biological Reviews 35:92–140. Journal of Paleontology 43:976–985. Muir-Wood, H. M. 1934. On the internal structure of ———. 1970. Variation in recent brachiopod popula- some Mesozoic Brachiopoda. Royal Society of Lon- tions. Bulletin of the Geological Institutions of the don Philosophical Transactions (series B) 223:511– University of Uppsala (new series) 2:41–48. 567, pl. 62–63. ———. 1971. Behaviour in the brachiopod Terebra- ———. 1962. On the morphology and classification tulina septentrionalis (Couthouy). Journal of Experi- of the brachiopod suborder Chonetoidea. British mental Marine Biology and Ecology 6:35–45. Museum (Natural History), monograph. viii + 132 ———. 1973. The ecology of Magellania venosa, an p., 24 fig., 16 pl. articulate brachiopod. Journal of Paleontology ———. 1965. Mesozoic and Cenozoic Terebratulidina. 47:266–278. In R. C. Moore, ed., Treatise of Invertebrate Paleon- ———. 1981. Physiology of the brachiopod digestive tology. Part H, Brachiopoda. Geological Society of system. In T. W. Broadhead, ed., Lophophorates, America & University of Kansas Press. Boulder & Notes for a Short Course. University of Tennessee Lawrence. p. 762–816, fig. 622–695. Department of Geological Sciences, Studies in Ge- Muir-Wood, H. M., & G. A. Cooper. 1960. Morphol- ology 5:170–204. ogy, classification and life habits of the McCammon, H., & R. Buchsbaum. 1968. Size and Productoidea (Brachiopoda). Geological Society of shape variation of three recent brachiopods from America Memoir 81:477 p., 135 pl. the Strait of Magellan. Antarctic Research Series Muir-Wood, H. M., G. F. Elliott, & Kotora Hatai. 2:215–225. 1965. Mesozoic and Cenozoic Terebratellidina. In McCammon, H. M., & W. A. Reynolds. 1976. Experi- R. C. Moore, ed., Treatise of Invertebrate Paleontol- mental evidence for direct nutrient assimilation by ogy. Part H, Brachiopoda. Geological Society of the lophophore of articulate brachiopods. Marine America & University of Kansas Press. Boulder & Biology 34:41–51. Lawrence. p. 816–857, fig. 696–741. McConnell, D. 1963. Inorganic constituents in the Müller, F. 1860. Beschreibung einer Brachiopoden shell of the living brachiopod Lingula. Geological Larva. Archives für Anatomie und Physiologie:72– Society of America Bulletin 74:363–364. 79. McCrady, J. 1860. On the Lingula pyramidata de- Neall, V. E. 1972. Systematics of the endemic New scribed by Mr. W. Stimpson. American Journal of Zealand brachiopod Neothyris. Journal of the Royal Science Arts (series 2) 30:157–158. Society of New Zealand 2:229–247. McDaniel, N. G. 1973. A survey of the benthic Negri, A., G. Tedeschi, F. Bonomi, J-H. Zhang, & D. macroinvertebrate fauna and solid pollutants in M. Kirtz. 1994. Amino-acid sequences of the alpha- Howe Sound. Fisheries Board of Canada, Technical and beta-subunits of hemerythrin from Lingula Report 385:1–64. reevii. Biochimica et Biophysica Acta 1208:277– Mickwitz, August. 1896. Über die Brachiopoden- 285. gathung Obolus Eichwald. Académie Impériale des Nekvasilova, O., & D. Pajaud. 1969. Le mode de fixa- Sciences de St. Petersbourg, Mémoires (série 8) tion chez Bifolium lacazelliforme Elliot (Thecideidae 4(2):215 p., 7 fig., 3 pl. Gray, Brachiopodes) au substrat. Casopis pro 518 Brachiopoda

Mineralogii a geologii 14:323–330. Owen, G., & Alwyn Williams. 1969. The caecum of Nielsen, Claus. 1987. The structure and function of articulate Brachiopoda. Proceedings of the Royal metazoan ciliary bands and their phylogenetic Society of London (series B) 172:187–201. significance. Acta Zoologica 68(4):205–262. Paine, R. T. 1962a. Ecological notes on a ———. 1991. The development of the brachiopod gymnophalline Metacercaria from the brachiopod Crania (Neocrania) anomala (O. F. Müller) and its Glottidia pyramidata. Journal of Parasitology phylogenetic significance. Acta Zoologica 72:7–28. 48(3):509. ———. 1994. Larval and adult characters in animal ———. 1962b. Filter-feeding pattern and local distri- phylogeny. American Zoologist 34:492–501. bution of the brachiopod Discinisca strigata. Bio- ———. 1995. Animal Evolution: Interrelationships of logical Bulletin 123:597–604. the Living Phyla. Oxford University Press. Oxford. ———. 1963. Ecology of the brachiopod Glottidia 467 p. pyramidata. Ecological Monographs 33:187–213. Nixon, K. C., & J. M. Carpenter. 1993. On out- ———. 1969. Growth and size distribution of the groups. Cladistics 9:413–426. brachiopod Terebratalia transversa Sowerby. Pacific Noble, J. P. A., & A. Logan. 1981. Size-frequency dis- Science 23:337–343. tributions and taphonomy of brachiopods: a recent ———. 1970. The sediment occupied by recent model. Palaeogeography, Palaeoclimatology, Palaeo- lingulid brachiopods and some paleoecological im- ecology 36:87–105. plications. Palaeogeography, Palaeoclimatology, Noble, J. P. A., A. Logan, & G. R. Webb. 1976. The Palaeoecology 7:21–31. recent Terebratulina community in the rocky sub- Palmer, A. R. 1981. Do carbonate skeletons limit the tidal zone of the Bay of Fundy, Canada. Lethaia rate of body growth? Nature 292:150–152. 9:1–17. ———. 1983. Relative cost of producing skeletal or- Nomura, S., & K. M. Hatai. 1936. A note on ganic matrix versus calcification: evidence from Coptothyris grayi (Davidson). Saito Ho-on Kai Mu- marine gastropods. Marine Biology 75:287–292. seum Research Bulletin 10:209–217. Pan, C.-M., & Norimitsu Watabe. 1988a. Uptake and Norford, B. S., & H. M. Steele. 1969. The Ordovician transport of shell minerals in Glottidia pyramidata trimerellid brachiopod Eodinobolus from southeast Stimpson (Brachiopoda: Inarticulata). Journal of Ontario. Palaeontology 12(1):161–171, pl. 32–33. Experimental Marine Biology and Ecology Norman, A. M. 1893. A month on the Trondhjem. 118:257–268, fig. 1–14. Annals and Magazine of Natural History (series 6) ———. 1988b. Shell growth of Glottidia pyramidata 12:340–367. Stimpson (Brachiopoda: Inarticulata). Journal of Odhner, N. H. 1960. Brachiopoda. Report of the Experimental Marine Biology and Ecology 119:43– Swedish Deep-Sea Expedition 2(4):23. 53, fig. 1–21. Ohuye, T. 1937. On the coelomic corpuscles in the ———. 1989. Periostracum formation and shell regen- body fluid of some invertebrates VIII. Supplemen- eration in the lingulid Glottidia pyramidata (Brach- tary note on the formed elements in the coelomic iopoda: Inarticulata). Transactions of the American fluid of some Brachiopoda. Scientific Reports of Microscopical Society 108(3):283–289. Tohoku University (series 4, Biology) 11:231–238. Pandian, T. J., & F. J. Vernberg. 1987. Animal Ener- Olsen, G. J., H. Matsuda, R. Hagstrom, & R. Over- getics, Volume 2, Bivalvia through Reptilia. Aca- beek. 1994. fastDNAml: a tool for construction of demic Press. San Diego. 631 p. phylogenetic trees of DNA sequences using maxi- Patel, N. H. 1994. Developmental evolution: insights mum likelihood. Computer Applications in the from studies of insect segmentation. Science Biological Sciences 10:41–48. 266:581–590. Onyia, A. D. 1973. Contribution to the food and Patterson, Colin. 1985. Introduction. In C. Patterson, feeding habit of the threadfin Galeoides decadactylus. ed., Molecules and Morphology in Evolution. Marine Biology 22(4):371–378. Third International Congress of Systematic & Evo- Öpik, A. A. 1930. Brachiopoda protremata der lutionary Biology, University of Sussex, July 1985. estlandischen Ordovizischen kukruse-stufe. Tartu Cambridge University Press. p. 1–22. Ulikooli Geoloogia-Instituudi Toimestuesed Acta et ———. 1989. Phylogenetic relations of major groups: Commentationes Universitatis Tartuensis 17:1– conclusions and prospects. In B. Fernholm, K. 261, pl. 1–22. Bremer, & H. Jörnvall, eds., The Hierarchy of Life: ———. 1933. Über die Plectamboniten. Tartu Molecules and Morphology in Phylogenetic Analy- Ulikooli Geoloogia-Instituudi Toimestuesed Acta et sis. Proceedings from Nobel Symposium 70, 1988. Commentationes Universitatis Tartuensis 28:1–79, Elsevier. Amsterdam. p. 471–488. 6 fig., pl. 1–12. Peck, L. S. 1989. Temperature and basal metabolism in ———. 1934. Über die Klitamboniten. Tartu Ulikooli two Antarctic marine herbivores. Journal of Experi- Geoloogia-Instituudi Toimestuesed Acta et mental Marine Biology and Ecology 127:1–12. Commentationes Universitatis Tartuensis 39:1– ———. 1992. Body volumes and internal space con- 239, 55 fig., pl. 1–48. straints in articulate brachiopods. Lethaia 25:383– Oró, J. & H. B. Skewes. 1965. Free amino acids on 390. human fingertips: the question of contamination in ———. 1993. The tissues of articulate brachiopods microanalysis. Nature 207:1042–1045. and their value to predators. Philosophical Transac- References 519

tions of the Royal Society of London (series B) (Argiope) neopolitana. Arbeiten aus dem 339:17–32. Zoologischen Institute der Universitaet Wien und Peck, L. S., A. Clarke, & L. J. Holmes. 1987a. Size, der Zoologischen Station in Triest, Wien 20:93– shape and the distribution of organic matter in the 108. recent Antarctic brachiopod Liothyrella uva. Lethaia Popov, L. Ye. 1992. The Cambrian radiation of 20:33–40. brachiopods. Topics in Geobiology 10:399–423, 6 ———. 1987b. Summer metabolism and seasonal fig. changes in biochemical composition of the Antarc- Popov, L. Ye., M. G. Bassett, L. E. Holmer, & J. tic brachiopod Liothyrella uva (Broderip, 1833). Laurie. 1993. Phylogenetic analysis of higher taxa Journal of Experimental Marine Biology and Ecol- of Brachiopoda. Lethaia 26:1–5. ogy 114:85–97. Popov, L. Ye., & Yu. A. Tikhonov. 1990. Ranne- Peck, L. S., G. B. Curry, A. D. Ansell, & Mark James. kembriiskie brakhiopody iz iuzhnoi Kirgizii [Early 1989. Temperature and starvation effects on the Cambrian brachiopods from south Kirgizii]. metabolism of the brachiopod Terebratulina retusa Paleontologicheskii Zhurnal 1990(3):33–44, 2 fig., (L.). Historical Biology 2:101–110. pl. 3–4. Peck, L. S., & L. J. Holmes. 1989a. Scaling patterns in Popov, L. Ye., & G. T. Ushatinskaya. 1987. Novyye the Antarctic brachiopod Liothyrella uva (Broderip, dannyye o mikrostrukture rakoviny bezzamkovykh 1833). Journal of Experimental Marine Biology and brakhiopod otryada Paterinida [New data on the Ecology 133:141–150, fig. 1–3. microstructure of the shell in inarticulate brachio- ———. 1989b. Seasonal and ontogenetic changes in pods of the Order Paterinida]. Doklady Akademii tissue size in the Antarctic brachiopod Liothyrella Nauk Soyuza Sovetskikh Sotsialisticheskikh uva (Broderip, 1833). Journal of Experimental Republik [Academy of Sciences of the USSR, Re- Marine Biology and Ecology 134:25–36, fig. 1–2. ports] 293(5):1228–1230. Peck, L. S., D. J. Morris, & Andrew Clarke. 1986a. Popov, L. Ye., O. N. Zezina, & J. Nolvak. 1982. Oxygen consumption and the role of caeca in the Mikrostruktura apikal’noy chasti rakoviny recent Antarctic brachiopod Liothyrella uva bezzamkovykh brakhiopod i eye paleoecologich- notorcadensis (Jackson, 1912). In P. R. Rachebœuf eskoye znacheniye [Microstructure of the apical & C. C. Emig, eds., Les Brachiopodes Fossiles et part of the shell in inarticulate brachiopods and its Actuels, Actes du 1er Congrès International sur les paleoecological significance]. Byuletin’ Moskov- Brachiopodes, Brest 1985. Biostratigraphie du skogo obshchestva ispytatelej prirody, otdel biolo- Paléozoïque 4. p. 349–355. gicheskij [Moscow Society of Natural History, ———. 1986b. The caeca of punctate brachiopods: a Branch of Biology, Bulletin] 87:94–104. respiring tissue not a respiratory organ. Lethaia Poulsen, V. 1971. Notes on an Ordovician acrotre- 19:232, fig. 1. tacean brachiopod from the Oslo region. Bulletin of Peck, L. S., D. J. Morris, A. Clarke, & L. J. Holmes. the Geological Society of Denmark 20:265–278. 1986. Oxygen consumption and nitrogen excretion Powls, R., & G. Britton. 1976. A carotenoprotein in the Antarctic brachiopod Liothyrella uva (Jack- violaxanthin, isolated from scenedesmus obliquus son, 1912) under simulated winter conditions. D3. Biochimica et Biophysica Acta 453:270–276. Journal of Experimental Marine Biology and Ecol- Prenant, M. 1928. Notes histologiques sur Terebra- ogy 104:203–213. tulina caput-serpentis L. Bulletin de la Société Zoo- Peck, L. S., & K. Robinson. 1994. Pelagic larval devel- logique de France 59:113–125. opment in the brooding Antarctic brachiopod Pross, A. 1980. Untersuchungen zur Gliederung von Liothyrella uva. Marine Biology 120:279–286. Lingula anatina (Brachiopoda). Archimerie bei Pedersen, Fl. Bo. 1978. A brief review of present theo- Brachiopoden. Zoologische Jahrbücher, abteilung ries of fjord dynamics. In C. J. Nihoul, ed., Hydro- für Anatomie und Ontogenie der Tiere 103:250– dynamics of Estuaries and Fjords. Elsevier Scientific 263. Publishing Company. Amsterdam. p. 407–422. Punin, M. Y., & M. V. Filatov. 1980. The epithelium Percival, E. 1944. A contribution to the life-history of organisation in the digestive gland of the articulate the brachiopod Terebratella inconspicua Sowerby. brachiopod Hemithyris psittacea. Citologija Transactions of the Royal Society of New Zealand 22(3):277–286. 74:1–23, pl. 1–7. ———. 1960. A contribution to the life history of the In Russian. brachiopod Tegulorhynchia nigricans. Quarterly Racheboeuf, P. R. 1973. Données nouvelles sur le Journal of Microscopical Science 101:439–457. développement des épines chez certains Brachio- Philippe, Hervé, A. Chenuil, & A. Adoutte. 1994. Can pods Chonetacea du Massif Armoricain. Compte- the Cambrian explosion be inferred through mo- Rendus de l’Académie des Sciences de Paris lecular phylogeny? In M. Akam, P. Holland, P. 277(D):1741–1744. Ingham, & G. Wray, eds., The Evolution of Devel- Racheboeuf, P. R., & Paul Copper. 1990. The opmental Mechanisms. Development, 1994 mesolophe, a new lophophore type for chonetacean Supplement. Company of Biologists. Cambridge. p. brachiopods. Lethaia 23(4):341–346, 5 fig. 15–25. Raff, R. A., C. R. Marshall, & J. M. Turbeville. 1994. Plenk, H. 1913. Die Entwicklung von Cistella Using DNA sequences to unravel the Cambrian 520 Brachiopoda

radiation of the animal phyla. Annual Review of ———. 1981c. Recent brachiopods from New Ecology and Systematics 25:351–375. Zealand. New Zealand Journal of Zoology 8:133– Ramamoorthi, K., K. Venkataramanujam, & B. 248. Srikrishnadhas. 1973. Mass mortality of Lingula ———. 1981d. Distribution and orientation of six anatina (Lam.) (Brachiopoda) in Porto-Novo wa- articulate species from New Zealand. New Zealand ters, S. India. Current Science 42(8):285–286. Journal of Zoology 8:189–196. Rand, D. M. 1993. Endotherms, ectotherms and mi- ———. 1986. Brachiopods. Scientific American tochondrial genome-size variation. Journal of Mo- 254:100–106. lecular Evolution 37:281–295. ———. 1987. Brachiopods from carbonate sands of Reed, C. G. 1987. Phylum Brachiopoda Coast. In M. the Australian shelf. Proceedings of the Royal Soci- Strathmann, ed., Reproduction and Development ety of Victoria 99:37–50. of Marine Invertebrates of the Northern Pacific. ———. 1991. Australasian Tertiary Brachiopoda. The University of Washington Press. Seattle. p. 486– subfamily Anakineticinae nov. Proceedings of the 493. Royal Society of Victoria 103(1):29–45, fig. 1–5. Reed, C. G., & R. A. Cloney. 1977. Brachiopod ten- ———. 1994. Origins and dispersal of a brachiopod tacles: ultrastructure and functional significance of family—the systematics, biogeography and evolu- the connective tissue and myoepithelial cells in tion of the Family Terebratellidae. Proceedings of Terebratalia. Cell and Tissue Research 185:17–42. the Royal Society of Victoria 106:17–29. Regnault, M. 1987. Nitrogen excretion in marine and Richardson, J. R., & R. J. Mineur. 1981. Differentia- fresh-water Crustacea. Biological Reviews 62:1–24. tion of species of Terebratella (Brachiopoda: Reynolds, W. A., & H. M. McCammon. 1977. As- Terebratellinae). New Zealand Journal of Zoology pects of the functional morphology of the lopho- 8:163–168. phore in articulate brachiopods. American Zoolo- Richardson, J. R., I. R. Stewart, & Liu Xixing. 1989. gist 17:121–132. Brachiopods from Chinese Seas. Chinese Journal of Rhodes, M. C. 1990. Extinction patterns and com- Oceanology and Limnology 7:211–219, pl. 1–4. parative physiology of brachiopods and bivalves. Richardson, J. R., & J. E. Watson. 1975. Form and PhD. thesis, Department of Geology, University of function in a recent free living brachiopod Maga- Pennsylvania. Philadelphia. 152 p. dina cumingi. Paleobiology 1:379–387. Rhodes, M. C., & C. W. Thayer. 1991. Effects of tur- Ricker, W. E. 1971. Methods for assessment of fish bidity on suspension feeding: Are brachiopods bet- production in fresh waters. I. B. P. Handbook No. ter than bivalves? In D. I. MacKinnon, D. E. Lee, 3. Blackwell Scientific Publications. Oxford & & J. D. Campbell, eds., Brachiopods Through Edinburgh, United Kingdom. 326 p. Time, Proceedings of the 2nd International Bra- Rickwood, A. E. 1968. A contribution to the life his- chiopod Congress, University of Otago, Dunedin, tory and biology of the brachiopod Pumilus New Zealand, 1990. Balkema. Rotterdam. p. 191– antiquatus Atkins. Transactions of the Royal Society 196. of New Zealand (Zoology) 10:163–182, 10 fig. Rhodes, M. C., & R. J. Thompson. 1992. Clearance ———. 1977. Age, growth and shape of the intertidal rate of the articulate brachiopod Neothyris len- brachiopod Waltonia inconspicua Sowerby, from ticularis (Deshayes, 1839). Journal of Experimental New Zealand. American Zoologist 17:63–73, fig. Marine Biology and Ecology 163:77–89. 1–9. ———. 1993. Comparative physiology of suspension Roberts, John. 1968. Mantle canal patterns in feeding in living brachiopods and bivalves: evolu- Schizophoria (Brachiopoda) from the Lower Car- tionary implications. Paleobiology 19:322–334. boniferous of New South Wales. Palaeontology Richards, J. R. 1952. The ciliary feeding mechanism of 11(3):389–405, pl. 74–75. Neothyris lenticularis (Deshayes). Journal of Mor- Robertson, J. D. 1989. Physiological constraints upon phology 90:65–91, fig. 1–6. marine organisms. Transactions of the Royal Soci- Richardson, J. R. 1973. Studies on Australian ety of Edinburgh, Earth Sciences 80:225–234. Cainozoic Brachiopods 3. The subfamily Bouchar- Rokop, F. J. 1977. Seasonal reproduction of the brach- diinae (Terebratellidae). Royal Society of Victoria iopod Frieleia halli and the scaphopod Cadulus Proceedings 86(1):127–132, pl. 7. californicus at bathyal depths in the deep sea. Ma- ———. 1975. Loop development and the classification rine Biology 43:237–246. of terebratellacean brachiopods. Palaeontology Rong, J.-Y., & L. R. M. Cocks. 1994. True Stro- 18(2):285–314, 4 fig., pl. 44. phomena and a revision of the classification and evo- ———. 1979. Pedicle structure of articulate brachio- lution of strophomenoid and “strophodontoid” pods. Journal of the Royal Society of New Zealand brachiopods. Palaeontology 37:651–694. 9:415–436. Rosenburg, G. D., & W. W. Hughes. 1991. A meta- ———. 1980. Studies on Australian Cainozoic brach- bolic model for the determination of shell compo- iopods 5. The genera Victorithyris Allan and Died- sition in the bivalve mollusc, Mytilus edulis. Lethaia rothyris nov. Memoirs of the National Museum of 24:83–96. Victoria 41:43–52. Rosenberg, G. D., W. W. Hughes, & R. D. Tkachuck. ———. 1981a. Brachiopods in mud: resolution of a 1988. Intermediatory metabolism and shell growth dilemma. Science 211:1161–1163. in the brachiopod Terebratalia transversa. Lethaia ———. 1981b. Brachiopods and pedicles. Paleo- 21:219–230. biology 7:87–95. Rouse, G. W., & B. G. M. Jamieson. 1987. An ultra- References 521

structural study of the spermatozoa of polychaetes vertebrates. Nature 266:533–535. Eurythoe complanata (Amphinomidae), Clymnella Sandy, M. R. 1994. Triassic–Jurassic articulate brachio- sp. and Micromaldane sp. (Maldanidae), with pods from the Pucará group, central Peru, and de- definition of sperm types in relation to reproductive scription of the brachidial net in the spiriferid biology. Journal of Submicroscopic Cytology Spondylospira. Palaeontographica A233:99–126. 19:573–584. Sandy, M. R., & M. R. Langenkamp. 1992. The Rowell, A. J. 1960. Some early stages in the develop- brachidial net—a unique internal support structure ment of the brachiopod Crania anomala (O. F. in the Brachiopoda? Geological Society of America, Müller). Annals and Magazine of Natural History Abstracts with Programs (Boulder) 24(7):A226. (series 13) 3:35–52. Sass, D. B., E. A. Monroe, & D. T. Gerace. 1965. Shell ———. 1961. Inhalant and exhalant feeding current structure of recent articulate Brachiopoda. Science systems in recent brachiopods. Geological Magazine 149:181–182. 98(3):261–263. Satake, K., M. Yugi, M. Kamo, H. Kihara, & A. ———. 1977. Valve orientation and functional mor- Tsugita. 1990. Hemerythrin from Lingula unguis phology of the foramen of some siphonotretacean consists of two different subunits, alpha and beta. and acrotretacean brachiopods. Lethaia 10:43–50. Protein Sequences and Data Analysis 3:1–5. ———. 1982. The monophyletic origin of the Savazzi, E. 1991. Burrowing in the inarticulate brach- Brachiopoda. Lethaia 15:299–307. iopod Lingula anatina. Palaeogeography, Palaeo- Rowell, A. J., & N. E. Caruso. 1985. The evolutionary climatology, Palaeoecology 85:101–106. significance of Nisusia sulcata, an early articulate Sawada, N. 1973. Electron microscope studies on ga- brachiopod. Journal of Paleontology 59:1227– metogenesis in Lingula unguis. Zoological Magazine 1242, fig. 1–9. 82:178–188. Rowley, A. F., & P. J. Hayward. 1985. Blood cells and Schaeffer, C. 1926. Untersuchungen zur verg- coelomocytes of the inarticulate brachiopod Lingula leichenden Anatomie und Histologie der Brach- anatina. Journal of Zoology, London (series A) iopodengattung Lingula. Acta Zoologica, 205:9–18. Stockholm 7:329–402. Rubenstein, D. I., & M. A. R. Koehl. 1977. The Schägger, H., & G. Von Jagow. 1987. Tricine-sodium mechanisms of filter feeding: some theoretical con- dodecyl sulfate-polyacrylamide gel electrophoresis siderations. American Naturalist 111:981–994. for the separation of proteins in the range from 1 to Rudall, K. M. 1955. The distribution of collagen and 100 kDa. Analytical Biochemistry 166:368–379. chitin. Symposium of the Society for Experimental Scheid, M. J., & J. Awapara. 1972. Stereospecificity of Biology 9:49–71. some invertebrate lactic dehydrogenases. Compara- ———. 1969. Chitin and its association with other tive Biochemistry and Physiology 43B:619–626. molecules. Journal of Polymer Science (Part C) Scheul, H. 1978. Secretory functions of egg cortical 28:83–102. granules in fertilisation and development. A critical Rudwick, M. J. S. 1959. The growth and form of review. Gamete Research 1:299–382. brachiopod shells. Geological Magazine 94:1–24. Schmidt, Herta. 1937. Zur Morphogenie der ———. 1960. The feeding mechanisms of spire- Rhynchonelliden. Senckenbergiana 19:22–60, fig. bearing fossil Brachiopoda. Geological Magazine 1–56. 97:369–383, 8 fig. Schmidt-Nielsen, K. 1979. Animal Physiology: Adap- ———. 1961. ‘Quick’ and ‘catch’ adductor muscles in tation and environment, second edition. Cam- brachiopods. Nature 191:1021. bridge University Press. Cambridge. 560 p. ———. 1962a. Filter feeding mechanisms in some ———. 1984. Scaling: Why is animal size so impor- brachiopods from New Zealand. Journal of the Lin- tant? Cambridge University Press. Cambridge. 241 nean Society, Zoology 44:592–615. p. ———. 1962b. Notes on the ecology of brachiopods Schram, F. R. 1983. Method and madness in phylog- in New Zealand. Transactions of the Royal Society eny. In F. R. Schram, ed., Crustacean Issues I: Crus- of New Zealand, Zoology 1:327–335. tacean Phylogeny. Balkema. Amsterdam. p. 331– ———. 1965a. Ecology and paleoecology. In R. C. 350. Moore, ed., Treatise on Invertebrate Paleontology. ———. 1991. Cladistic analysis of metazoan phyla Part H, Brachiopoda. Geological Society of and the placement of fossil problematica. In A. M. America & University of Kansas Press. New York & Simonetta & Simon Conway Morris, eds., The Lawrence. p. 199–214. Early Evolution of the Metazoa and the Significance ———. 1965b. Sensory spines in the Jurassic brachio- of Problematic Taxa. Cambridge University Press. pod Acanthothiris. Palaeontology 8:604–617, pl. Cambridge. p. 35–46. 84–87. Schuchert, Charles, & G. A. Cooper. 1932. Brachio- ———. 1970. Living and Fossil Brachiopods. pod genera of the suborders Orthoidea and Hutchinson and Co. Ltd. London. 199 p., 99 fig. Pentameroidea. Peabody Museum of Natural His- Runnegar, B., A. Scheltema, L. Jackson, M. Jebb, J. tory Memoir 49(1):1–270, pl. 1–29. McC. Turbeville, & C. R. Marshall. In preparation. Schulgin, M. A. 1885. Argiope kowalewskii (ein A molecular test of the hypothesis that Aplacophora Beitrag zur Kenntniss der Brachiopoden). are progenetic aculifera. Zeitschrift für wissenschaftliche Zoologie 41:116– Russell, G. R., & J. H. Subak-Sharpe. 1977. Similar- 141. ity of the general designs of protochordates and in- Schumann, Dietrich. 1969. “Byssus” - artige 522 Brachiopoda

Stielmuskel - Konvergenzen bei artikulaten Brach- the 2nd International Brachiopod Congress, Uni- iopoden. Neues Jahrbuch für Geologie und versity of Otago, Dunedin, New Zealand, 1990. Palaeontologie, Abhandlungen 133:199–210. Balkema. Rotterdam. p. 159–165. ———. 1970. Inäquivalver Schalenbau bei Crania Smith, A. B. 1994. Rooting molecular trees: problems anomala. Lethaia 3:413–421. and strategies. Biological Journal of the Linnean ———. 1973. Mesodermale Endoskelette Society 51:279–292. terebratulider Brachiopoden. I. Palaeontologische Smith, M. J., A. Arndt, S. Gorski, & E. Fajber. 1993. Zeitschrift 47(1–2):77–103, 10 fig., pl. 12–15. The phylogeny of echinoderm classes based on mi- ———. 1991. Hydrodynamic influences in brachio- tochondrial gene arrangements. Journal of Molecu- pod shell morphology of Terebratalia transversa lar Evolution 36:545–554. (Sowerby) from the San Juan Islands, USA. In D. I. Soota, T. D., & K. N. Reddy. 1976. On the distribu- MacKinnon, D. E. Lee, & J. D. Campbell, eds., tion and habitat of the brachiopod Lingula in India. Brachiopods Through Time, Proceedings of the 2nd Newsletter of the Zoological Survey of India International Brachiopod Congress, University of 2(6):235–237. Otago, Dunedin, New Zealand, 1990. Balkema. Steele, K. P., K. E. Holsinger, R. K. Jansen, & D. W. Rotterdam. p. 265–272. Taylor. 1991. Assessing the reliability of 5S rRNA Scott, M. P. 1994. Intimations of a creature. Cell sequence data for phylogenetic analysis in green 79:1121–1124. plants. Molecular Biology and Evolution 8:240– Senn, E. 1934. Die Geschlechtsuerhaeltnisse der 248. Brachiopoden im besonderen die Spermato—und Steele-Petrovic, H. M. 1976. Brachiopod food and Oogenese der Lingula. Acta Zoologica 15:1–154. feeding processes. Palaeontology 19:417–436. Sewell, R. B. S. 1912. Note on the development of the Stehli, F. G. 1956. Evolution of the loop and lopho- larva of Lingula. Records of the Indian Museum phore in terebratuloid brachiopods. Evolution 7:88–90. 10:187–200, 9 fig. Shiells, K. A. G. 1968. Kochiproductus coronus sp. nov. ———. 1965. Paleozoic Terebratulida. In R. C. from the Scottish Viséan and a possible mechanical Moore, ed., Treatise on Invertebrate Paleontology. advantage of its flange structure. Transactions of the Part H, Brachiopoda. Geological Society of Royal Society of Edinburgh 67:477–510, 20 fig., 1 America & University of Kansas Press. New York & pl. Lawrence. p. 730–762, fig. 594–621. Shimizu, N., & K-I. Miura. 1972. Studies on nucleic Steinich, G. 1965. Die articulaten Brachiopoden der acids of living fossils. I. Isolation and characteriza- ruegener Schreibkreide (Unter-Maastricht). Geo- tion of DNA and some RNA components from the logische und Palaeontologische Abhandlungen brachiopod Lingula. Animal ribosomes: Experimen- (Abt. A) 2:220 p., 21 pl. tal studies of the last five years. MSS Information Stewart, I. R. 1975. Morphological variation within Corporation. New York. p. 19–25. the recent brachiopod genus Magellania. Unpub- Shipley, A. E. 1883. On the structure and development lished thesis submitted for Geology Fellowship Di- of Argiope. Zoologie Stazione Neapoli, Mittei- ploma. Royal Melbourne Institute of Technology. lungen 4:494–520. 13 p. Shipley, A. E., & E. W. MacBride. 1920. Zoology. 4th ———. 1981. Population structure of articulate bra- edition. Cambridge University Press. Cambridge. chiopod species from soft and hard substrates. New 752 p. Zealand Journal of Zoology 8:197–208. Short, G., & S. L. Tamm. 1991. On the nature of St. Joseph, J. K. S. 1938. The Pentameracea of the paddle cilia and discocilia. Biological Bulletin, Ma- Oslo region. Norsk Geologisk Tidsskrift 17:255– rine Biological Laboratory, Woods Hole, Massachu- 336, pl. 1–8. setts 180:466–474. Storch, Volker, & Ulrich Welsch. 1972. Über Bau und Shumway, S. E. 1982. Oxygen consumption in brach- Entstehung der Mantelrandstacheln von Lingula iopods and the possible role of punctae. Journal of unguis L. (Brachiopoda). Zeitschrift für Wissen- Experimental Marine Biology and Ecology 58:207– schaftliche Zoologie (Leipzig) 183:181–189. 220. ———. 1974. Epitheliomuscular cells in Lingula un- Simon, J. L., & D. M. Dauer. 1977. Reestablishment guis (Brachiopoda) and Branchiostoma lanceolatum of a benthic community following natural de- (Acrania). Cell and Tissue Research 154:543–545. faunation. In B. C. Coull, ed., Ecology of Marine ———. 1975. Elektronenmikroskopische und enzym- Benthos. University of South Carolina Press, Co- histochemische Untersuchungen über die lumbia, Belle W. Baruch Library in Marine Science Mitteldarmdrüse von Lingula unguis L. (Brach- 6:139–154. iopoda). Zoologische Jahrbücher, abteilung für Singer, T. P. 1965. Comparative biochemistry of succi- Anatomie und Ontogenie der Tiere 94:441–452. nate dehydrogenase: forms and functions. In T. E. ———. 1976. Elektronenmikroskopische und enzym- King, H. S. Mason, & M. Morrison, eds., Oxidases histochemische Untersuchungen über Lophophor and Related Redox Systems. Wiley. New York. p. und Tentakeln von Lingula unguis L. (Brach- 448–481. iopoda). Zoologische Jahrbücher, abteilung für Smirnova, T. N., & E. Popiel-Barczyk. 1991. Charac- Anatomie und Ontogenie der Tiere 96:225–237. teristics of the shell ultrastructure in Terebratellacea. Stoyanova, R. Z. 1984. Alkane and fatty acid content In D. I. MacKinnon, D. E. Lee, & J. D. Campbell, and composition in Palaeozoic Brachiopoda fossils. eds., Brachiopods Through Time, Proceedings of Comptes-rendus de l’Académie Bulgare des Sci- References 523

ences 37(6):771–774. Sinauer Associates, Inc. Sunderland, Massachusetts. Strathmann, R. R. 1973. Function of lateral cilia in p. 3–27. suspension feeding of lophophorates (Brachiopoda, Thayer, C. W. 1975. Size-frequency and population Phoronida, Ectoprocta). Marine Biology 23:129– structure of brachiopods. Palaeogeography, Palaeo- 136. climatology, Palaeoecology 17:139–148. Strathmann, R. R., & D. J. Eernisse. 1994. What ———. 1977. Recruitment, growth, and mortality of molecular phylogenies tell us about the evolution of a living articulate brachiopod, with implications for larval forms. American Zoologist 34:502–512. the interpretation of survivorship curves. Paleo- Stricker, S. A., & C. G. Reed. 1985a. The ontogeny of biology 3:98–109. shell secretion in Terebratalia transversa (Brach- ———. 1986a. Are brachiopods better than bivalves? iopoda, Articulata). I. Development of the mantle. Mechanisms of turbidity tolerance in articulates Journal of Morphology 183:233–250. and their interaction with feeding in articulates. ———. 1985b. The ontogeny of shell secretion in Paleobiology 12:161–174. Terebratalia transversa (Brachiopoda, Articulata). II. ———. 1986b. Respiration and the function of brach- Formation of the protegulum and juvenile shell. iopod punctae. Lethaia 19:23–31. Journal of Morphology 183:251–271, fig. 1–45. Thayer, C. W., & R. A. Allmon. 1990. Evolutionary ———. 1985c. Development of the pedicle in the ar- refugia? Oligotrophic marine caves of Micronesia. ticulate brachiopod Terebratalia transversa (Brachio- Abstract. Fourth International Congress of System- poda, Terebratulida). Zoomorphology 105:253– atic and Evolutionary Biology. p. A323. 264. Thayer, C. W., & H. M. Steele-Petrovic. 1975. Bur- Suchanek, T. H., & J. Levinton. 1974. Articulate bra- rowing of the lingulid brachiopod Glottidia pyra- chiopod food. Journal of Paleontology 48:1–5. midata: its ecologic and palaeoecologic significance. Summers, R. G. 1970. The fine structure of the sper- Lethaia 8:209–221. matozoon of Pennaria tiarella (Coelenterata). Jour- Thomas, G. A. 1958. The Permian Orthotetacea of nal of Morphology 131:117–130. Western Australia. Bureau of Mineral Resources, Sundarsan, D. 1968. Brachiopod larvae from the west Geology and Geophysics, Bulletin 39:1–159. coast of India. Proceedings of the Indian Academy Thompson, R. R., & W. B. Creath. 1966. Low mo- of Science (section B) 68:59–68. lecular weight hydrocarbons in recent and fossil ———. 1970. On a lingulid larva from Coondapur shells. Geochimica et Cosmochimica Acta (Mysore State), India. Journal of the Marine Bio- 30:1137–1152. logical Association of India 12:97–99. Thompson, R. J., E. Ward, & M. C. Rhodes. 1992. In Surlyk, Finn. 1972. Morphological adaptations and vivo observations of feeding in an articulate bra- population structures of the Danish chalk brachio- chiopod. Abstract. Annual Marine Benthic Ecology pods (Maastrichtian, Upper Cretaceous). Kongelige Meeting, Newport, RI, March 1992. Danske Videnskabernes Selskab, Biologiske Skrifter Thomson, J. A. 1918. Brachiopoda. Australasian Ant- 19(2):2–57, 24 fig., pl. 1–5. arctic Expedition, 1911–1914, Scientific Reports Swedmark, B. 1967. Gwynia capsula (Jeffreys), an ar- (series C) 4:1–76, pl. 15–18. ticulate brachiopod with brood protection. Nature ———. 1927. Brachiopod morphology and genera (re- 213:1151–1152. cent and Tertiary). New Zealand Board of Science ———. 1971. A review of Gastropoda, Brachiopoda, and Art, Manual 7:338 p., 103 fig., 2 pl. and Echinodermata. Smithsonian Contributions to Tkachuck, R. D., G. D. Rosenberg, & W. W. Hughes. Zoology 76:41–45. 1989. Utilization of free amino acids by mantle tis- Swofford, D. L. 1993. PAUP: phylogenetic analysis sue in the brachiopod Terebratalia transversa and the using parsimony. Computer program distributed by bivalve mollusc, Chlamys hastata. Comparative Bio- the author. Smithsonian Institution. Washington, chemistry and Physiology 92B:747–750. D.C. Tommasi, L. R. 1970a. Sôbre o braquiopode Taddei Ruggiero, E. 1991. A study of damage evidence Bouchardia rosea (Mawe, 1823). Boletim do in brachiopod shells. In D. I. MacKinnon, D. E. Instituto Oceanografico, Sao Paulo 19:33–42. Lee, & J. D. Campbell, eds., Brachiopods Through ———. 1970b. Observaçoes sôbre a fauna bêntico do Time, Proceedings of the 2nd International Brach- complexo estuarino-lagunar de Cananéia. Boletim iopod Congress, University of Otago, Dunedin, do Instituto Oceanografico, Sao Paulo 19(1):43– New Zealand, 1990. Balkema. Rotterdam. p. 203– 56. 210. Tortell, P. 1981. Notes on the reproductive biology of Tanaka, S., K. Anno, & N. Seno. 1982. A novel sul- brachiopods from southern New Zealand. New fated glycosaminoglycan, lingulin sulfate, composed Zealand Journal of Zoology 8:175–182. of galactose and N-acetylgalactosamine from Lin- Towe, K. M. 1980. Preserved organic ultrastructure: an gula unguis. Biochimica et Biophysica Acta unreliable indicator for paleozoic amino acid bio- 704:549–551. geochemistry. In P. E. Hare, T. C. Hoering, & K. J. Teichert, Curt. 1958. Cold and deep-water coral King, eds., Biogeochemistry of Amino Acids. John banks. Bulletin of the American Association of Pe- Wiley & Sons. New York. p. 65–74. troleum Geologists 42:1064–1082. Towe, K. M., & C. W. Harper, Jr. 1966. Pho- Templeton, Alan. 1989. The meaning of species and lidostrophiid brachiopods: origin of the nacreous speciation: a genetic perspective. In D. Otte & J. A. lustre. Science 154:153–155. Endler, eds., Speciation and Its Consequences. Trueman, E. R., & T. M. Wong. 1987. The role of the 524 Brachiopoda

coelom as a hydrostatic skeleton in lingulid brach- intracrystalline proteins and amino acids. Ph.D. iopods. Journal of Zoology 213(2):221–232, 3 fig. thesis. University of Glasgow. 237 p. Tsuchiya, M., & C. C. Emig. 1983. Macrobenthic as- Walton, Derek, & G. B. Curry. 1991. Amino acids semblages in a habitat of the recent lingulid brach- from fossils, facies and fingers. Palaeontology iopod Lingula anatina Lamarck at Asamushi, 34:851–858. Northern Japan. Bulletin of the Marine Biological Walton, Derek, Maggie Cusack, & G. B. Curry. 1993. Station of Asamushi, Tohoku University Implications of the amino acid composition of re- 17(3):141–157. cent New Zealand brachiopods. Palaeontology Tunnicliffe, Verena. 1981. High species diversity and 36:883–896. abundance of the epibenthic community in an oxy- Wang, Yu, Paul Copper, & Jia-Yu Rong. 1983. Distri- gen-deficient basin. Nature 294:354–356. bution and morphology of the Devonian brachio- Tuross, Noreen, & L. W. Fisher. 1989. The proteins in pod Punctatrypa. Journal of Paleontology 57:1067– the shell of Lingula. In R. E. Crick, ed., Origin, 1089, 11 fig. Evolution, and Modern Aspects of Biomineral- Watabe, Norimitsu, & C.-M. Pan. 1984. Phosphatic ization in Plants and Animals. Plenum Press. New shell formation in atremate brachiopods. American York. p. 325–328. Zoologist 24:977–985. Tuross, Noreen, & P. E. Hare. 1990. A 40 kDa protein Webb, G. R., A. Logan, & J. P. A. Noble. 1976. Oc- in modern and fossil Lingula. In The Sixth Interna- currence and significance of brooded larva in a re- tional Symposium on Biomineralization, Confer- cent brachiopod, Bay of Fundy, Canada. Journal of ence Abstracts. Kyoritsu Shuppan Co. Ltd. Tokyo, Paleontology 50:869–871. Japan. p. 76. Weiner, S. 1983. Mollusk shell formation: isolation of Ulrich, E. O., & G. A. Cooper. 1938. Ozarkian and two organic matrix proteins associated with calcite Canadian Brachiopoda. Geological Society of deposition in the bivalve Mytilus californianus. Bio- America Special Paper 13:viii + 323 p., 14 fig., 58 chemistry 22:4139–4145. pl. Westbroek, Peter. 1967. Morphological observations Uribe, M. E., & A. P. Larrain. 1992. Estudios with systematic implications on some Palaeozoic biológicos en el enteropneusto Ptychodera flava Rhynchonellida from Europe, with special empha- Eschscholtz, 1825, de Bahia Conceptión, Chile. I: sis on the Uncinulidae. Thesis. Leidse Geologische Aspectos morphológicos y ecológicos. Gayana, Mededelingen 41:1–82, 14 pl. Zoologia 56(3–4):141–180. Westbroek, Peter, J. Yanagida, & Y. Isa. 1980. Func- Ushatinskaya, G. T. 1988. Obolellidy (Brakhiopody) s tional morphology of brachiopod and coral skeletal zamkovym sochleneniem stvorok iz nizhnego structures supporting ciliated epithelia. Paleo- kembriia Zabaikal’ia [Obolellids (Brachiopoda) biology 6:313–330. with a hinge from the Lower Cambrian of the Widdows, J. 1985. Physiological procedures. In B. L. Transbaikal area]. Paleontologicheskii Zhurnal Bayne, ed., The Effects of Stress and Pollution on 1988(1):34–39, 1 fig., pl. 3–4. Marine Animals. Praeger Scientific. New York. p. Vader, W. 1970. The amphipod, Aristias neglectus 161–178. Hansen, found in association with brachiopods. Wilkens, J. L. 1978a. Adductor muscles of brachio- Sarsia 43:13–14. pods: activation and contraction. Canadian Journal Valentine, J. W., & F. J. Ayala. 1975. Genetic variation of Zoology 56:315–323. in Freileia halli, a deep sea brachiopod. Deep-Sea ———. 1978b. Diductor muscles of brachiopods: ac- Research 22:37–44. tivation and very slow contraction. Canadian Jour- Vallentyne, J. R. 1964. Biogeochemistry of organic nal of Zoology 56:324–332. matter II: Thermal reaction kinetics and transfor- ———. 1987. Tonic free maintenance after the decay mation products of amino compounds. Geochimica of active state in brachiopod smooth adductor et Cosmochimica Acta 32:1353–1356. muscle. Journal of Comparative Physiology (series Vandercammen, A., & M. Lambiotte. 1962. Observa- B) 157:651–658. tions sur les sarcoglyphes dans Atrypa reticularis (C. Williams, Alwyn. 1953. North American Linné, 1767). Bulletin de l'Institut Royal des Sci- stropheodontids: their morphology and systematics. ences Naturelles de Belgique, Bruxelles 38:1–15, pl. Geological Society of America Memoir 56:67 p., 13 1. pl. Van Kleef, F. S. M., W. W. de Jong, & H. J. Hoenders. ———. 1956. The calcareous shell of the Brachiopoda 1975. Stepwise degradation of the eye lens protein and its importance to their classification. Biological α-crystallin in aging. Nature 258:264–266. Reviews 31:243–287, fig. 1–7. Vargas, J. A. 1988. Community structure of macro- ———. 1957. Evolutionary rates of brachiopods. Geo- benthos and the results of macropredator exclusion logical Magazine 94:201–211. on a tropical intertidal mud flat. Revista de Biología ———. 1962. The Barr and lower Ardmillian series Tropical 36(2A):287–308. (Caradoc) of the Girvan district, southwest Ayr- Walker, A. O. 1909. Amphipoda Gammaridea from shire, with descriptions of the Brachiopoda. Geo- the Indian Ocean, British East Africa, and the Red logical Society of London Memoir 3:267 p., 25 pl. Sea. Transactions of the Linnean Society of London ———. 1963. The Caradocian brachiopod faunas of (series 2, Zoology) 12:323–344, pl. 42–43. the Bala District, Merionethshire. Bulletin of the Walton, Derek. 1992. Biogeochemistry of brachiopod British Museum (Natural History) Geology, Lon- References 525

don 8:327–471, 13 fig., 16 pl. International Brachiopod Congress, University of ———. 1965. Stratigraphic Distribution. In R. C. Otago, Dunedin, New Zealand, 1990. Balkema. Moore, ed., Treatise on Invertebrate Paleontology. Rotterdam. p. 133–140, fig. 1–4. Part H, Brachiopoda. Geological Society of Williams, Alwyn, Maggie Cusack, & Sarah Mackay. America & University of Kansas Press. New York & 1994. Collagenous chitinophosphatic shell of the Lawrence. p. 237–250, fig. 148–154. brachiopod Lingula. Philosophical Transactions of ———. 1966. Growth and structure of the shell of liv- the Royal Society of London (series B) 346:223– ing articulate brachiopods. Nature 211:1146–1148. 266. ———. 1968a. Evolution of the shell structure of ar- Williams, Alwyn, & R. A. Hewitt. 1977. The del- ticulate brachiopods. Special Papers in Palaeon- thyrial covers of some living brachiopods. Proceed- tology 2:1–55, 27 fig., 24 pl. ings of the Royal Society of London (series B) ———. 1968b. Significance of the structure of the 197:105–129, pl. 1–7. brachiopod periostracum. Nature 218:551–554. Williams, Alwyn, & L. E. Holmer. 1992. Ornamenta- ———. 1968c. Shell structure of the billingsellacean tion and shell structure of acrotretoid brachiopods. brachiopods. Palaeontology 11:486–490. Palaeontology 35:657–692. ———. 1968d. A history of skeletal secretion among Williams, Alwyn, & Sarah Mackay. 1978. Secretion articulate brachiopods. Lethaia 1:268–287. and ultrastructure of the periostracum of some ———. 1970a. Origin of laminar-shelled articulate terebratulide brachiopods. Proceedings of the Royal brachiopods. Lethaia 3:329–342. Society of London (series B) 202:191–209. ———. 1970b. Spiral growth of the laminar shell of ———. 1979. Differentiation of the brachiopod the Brachiopod Crania. Calcified Tissue Research periostracum. Palaeontology 22:721–736. 6:11–19. Williams, Alwyn, Sarah Mackay, & Maggie Cusack. ———. 1971a. Comments on the growth of the shell 1992. Structure of the organo-phosphatic shell of of articulate brachiopods. In J. Thomas Dutro, Jr., the brachiopod Discinisca. Philosophical Transac- ed., Paleozoic Perspectives: A Paleontological Trib- tions of the Royal Society of London (series B) ute to G. Arthur Cooper. Smithsonian Contribu- 337:83–104. tions to Paleobiology 3:47–67. Williams, Alwyn, & A. J. Rowell. 1965a. Brachiopod ———. 1971b. Scanning electron microscopy of the anatomy. In R. C. Moore, ed., Treatise on Inverte- calcareous skeleton of fossil and living Brachiopoda. brate Paleontology. Part H, Brachiopoda. Geologi- In V. H. Heywood, ed., Scanning Electron Micros- cal Society of America & University of Kansas copy. Systematic and Evolutionary Applications. Press. New York & Lawrence. p. 6–57, fig. 1–58. Academic Press. London & New York. p. 37–66. ———. 1965b. Morphology. In R. C. Moore, ed., ———. 1973. The secretion and structural evolution Treatise on Invertebrate Paleontology. Part H, of the shell of thecideidine brachiopods. Philo- Brachiopoda. Geological Society of America & sophical Transactions of the Royal Society of Lon- University of Kansas Press. New York & Lawrence. don (series B) 264:439–478. p. 57–138, fig. 59–138. ———. 1974. Ordovician Brachiopoda from the Williams, Alwyn, A. J. Rowell, D. V. Ager, G. F. Shelve District, Shropshire. Bulletin of the British Elliott, R. E. Grant, H. M. Muir-Wood, & F. G. Museum (Natural History) Geology, Supplement Stehli. 1965. Morphological terms applied to 11:1–163, 11 fig., 28 pl. Brachiopods. In R. C. Moore, ed., Treatise on In- ———. 1977. Differentiation and growth of the vertebrate Paleontology. Part H, Brachiopoda. Geo- brachiopod mantle. American Zoologist 17:107– logical Society of America & University of Kansas 120. Press. New York & Lawrence. p. 139–155. ———. 1984. Lophophorates. In J. Bereiter-Hahn, A. Williams, Alwyn, & A. D. Wright. 1961. The origin of G. Matoltsy, & K. S. Richards, eds., Biology of the the loop in articulate brachiopods. Palaeontology Integument, Volume 1, Invertebrates. Springer- 4:149–176, fig. 1–13. Verlag. Berlin, Germany. p. 728–745. ———. 1963. The classification of the “Orthis ———. 1990. Biomineralization in the lophophorates. testudinaria Dalman” group of brachiopods. Journal In J. G. Carter, ed., Skeletal Biomineralization: Pat- of Paleontology 37:1–32, fig. 1–10, pl. 1–2. terns, Processes and Evolutionary Trends. Volume I ———. 1965. Orthida. In R. C. Moore, ed., Treatise & II. Van Nostrand Reinhold. New York. p. 67–82. on Invertebrate Paleontology. Part H, Brachiopoda. Williams, Alwyn, & C. H. C. Brunton. 1993. Role of Geological Society of America & University of Kan- shell structure in the classification of the ortho- sas Press. New York & Lawrence. p. 299–359, fig. tetidine brachiopods. Palaeontology 36:931–966. 188–228. Williams, Alwyn, S. J. Carlson, C. H. C. Brunton, L. ———. 1970. Shell structure of the Craniacea and E. Holmer, & L. E. Popov. 1996. A supra-ordinal other calcareous inarticulate brachiopods. Special classification of the Brachiopoda. Philosophical Papers in Palaeontology 7:1–51, 15 pl. Transactions of the Royal Society of London Willmer, P. G., & P. W. H. Holland. 1991. Modern B351:1171–1193. approaches to metazoan relationships. Journal of Williams, Alwyn, & G. B. Curry. 1991. The micro- Zoology 224:689–694. architecture of some acrotretide brachiopods. In D. Wilson, G. D. F. 1987. Speciation in the deep sea. I. MacKinnon, D. E. Lee, & J. D. Campbell, eds., Annual Review of Ecology and Systematics 18:185– Brachiopods Through Time, Proceedings of the 2nd 207. 526 Brachiopoda

Wilson, H., & R. K. Cannon. 1937. The glutamic Young, J. 1884. Notes on the shell structure of acid-pyrrolidone carboxylic acid system. Journal of Eichwaldia capewelli. Geological Magazine 1(series Biological Chemistry 119:309–331. 3):214–218. Winberg, G. G. 1956. Rate of metabolism and food Zabi, S. G. 1984. Rôle de la biomasse dans la requirements of fishes. Trudy Belorusskogo détermination de l’“importance value” pour la mise gosudarstvennogo universiteta Minske. 253 p. en évidence des unités de peuplements benthiques en Lagune Ebrié (Côte d’Ivoire). Documents Scien- Translated from Russian by Fisheries Research tifiques du Centre de Recherches Océano- Board of Canada, Translation Series 194, 1960. graphiques, Abidjan 15(1/2):55–87. Winnepenninckx, B., T. Backeljau, & R. De Wachter. Zezina, O. N. 1961. Distribution of the deepwater 1995. Phylogeny of protostome worms derived brachiopod Pelagodiscus atlanticus (King). from 18S rRNA sequences. Molecular Biology and Okeanology 5(2):354–358. Evolution 12:641–649. Wisely, B. 1969. Preferential settlement in concavities In Russian. (rugophilic behaviour) by larvae of the brachiopod ———. 1970. Brachiopod distribution in the recent Waltonia inconspicua (Sowerby, 1846). New ocean with reference to problems of zoogeographic Zealand Journal of Marine and Freshwater Research zoning. Paleontologicheski Zhurnal 2:3–17. 3:237–280. Witman, J. D., & R. A. Cooper. 1983. Disturbance In Russian. and contrasting patterns of population structure in ———. 1975. Deep-sea brachiopods from the south- the brachiopod Terebratulina septentrionalis (Couth- east Pacific and Scotia Sea. Trudy Instituta ouy) from two subtidal habits. Journal of Experi- Okeanologii 103:247–258. mental Marine Biology and Ecology 73:57–79. Worcester, W. 1969. On Lingula reevii. Unpublished In Russian. Master of Science thesis. University of Hawaii. 49 ———. 1981. Recent deep-sea Brachiopoda from the p. western Pacific. Galathea Report 15:7–20. Wourms, J. S. 1987. Oogenesis, p. 49–187. In A. C. ———. 1985. Sovremennye brakhiopody i problemy Giese, J. S. Pearse, & V. B. Pearse, eds., Reproduc- batial noi zony okeana [Living brachiopods and tion of Marine Invertebrates. Volume 9. General problems of the north bathyal oceans]. Nauka. aspects: seeking unity in diversity. Blackwell Moscow. 247 p. Scientific Publications. California. 712 p. ———. 1987. Brachiopods collected by BENTHEDI- Wright, A. D. 1966. The shell punctation of Dicoelosia Cruise in the Mozambique Channel. Bulletin du biloba (Linnaeus). Geologiska Foreningens I Stock- Muséum Nationale d'Histoire naturelle de Paris holm Forhandlingar 87:548–556. (série 4) 9:551–563. ———. 1981. The external surface of Dictyonella and Zhang, J-H., & D. M. Kurtz. 1991. Two distinct sub- of other pitted brachiopods. Palaeontology 24:443– units of hemerythrin from the brachiopod Lingula 481, pl. 62–71. reevii: an apparent requirement for cooperativity in Yamada, M. 1956. Notes on discinid larvae (Brach- oxygen binding. Biochemistry 30:9121–9125. iopoda) from Osyro, West coast of Hokkaido. An- Zittel, K. A. von. 1880. Handbuch der Palaeontologie, notated Zoology of Japan 29:165–167. Vol. 1, No. 4. R. Oldenbourg. München & Leipzig. Yano, Hiroyuki, K. Satake, Y. Ueno, K. Kondo, & A. p. 641–722. Tsugita. 1991. Amino acid sequence of the Zuckerkandl, Emil, & L. Pauling. 1965. Molecules as haemerythrin α subunit from Lingula unguis. Jour- documents of evolutionary history. Journal of nal of Biochemistry 110:376–380. Theoretical Biology 8:357–366. Yano, Hiroyuki, K. Satake, Y. Ueno, & A. Tsugita. Zuker, Michael. 1989. On finding all suboptimal 1991. The amino acid sequence of the β chain of foldings of an RNA molecule. Science 244:48–52. hemerythrin from Lingula unguis. Protein Se- Zwaan, A. de, M. Leopold, E. Marteyn, & D. R. quences and Data Analysis 4:87–91. Livingstone. 1982. Phylogenetic distribution of Yatsu, Naukidé. 1902a. On the development of Lingu- pyruvate oxidoreductases, arginine kinase and la anatina. Journal of the College of Science, Impe- aminotransferases. In A. D. F. Addink & N. rial University, Tokyo, Japan 17(4):1–112, pl. 1–8. Spronk, eds., Exogenous and Endogenous Influ- ———. 1902b. On the habits of the Japanese Lingula. ences on Metabolic and Neural Control, Volume 2. Annotationes Zoologicae Japonenses 4(2):61–67. Pergamon Press. Oxford. p. 136–137.