1 This article was downloaded by: [ ] On: 04 July 2015, At: 20:54 Publisher: Taylor & Francis Informa Ltd Registered in England and Wales Registered Number: Registered office: 5 Howick Place, London, SW1P 1WG Journal of the Royal Society of New Zealand Publication details, including instructions for authors and subscription information: Kakanuiella (gen. nov.) and Thecidellina: Cenozoic and Recent thecideide brachiopods from New Zealand Daphne E. Lee a & Jeffrey H. Robinson a a Department of Geology, University of Otago, P.O. Box 56, Dunedin, New Zealand Published online: 30 Mar To cite this article: Daphne E. Lee & Jeffrey H. Robinson (2003) Kakanuiella (gen. nov.) and Thecidellina: Cenozoic and Recent thecideide brachiopods from New Zealand, Journal of the Royal Society of New Zealand, 33:1, , DOI: / To link to this article: PLEASE SCROLL DOWN FOR ARTICLE Taylor & Francis makes every effort to ensure the accuracy of all the information (the Content ) contained in the publications on our platform. However, Taylor & Francis, our agents, and our licensors make no representations or warranties whatsoever as to the accuracy, completeness, or suitability for any purpose of the Content. Any opinions and views expressed in this publication are the opinions and views of the authors, and are not the views of or endorsed by Taylor & Francis. The accuracy of the Content should not be relied upon and should be independently verified with primary sources of information. Taylor and Francis shall not be liable for any losses, actions, claims, proceedings, demands, costs, expenses, damages, and other liabilities whatsoever or howsoever caused arising directly or indirectly in connection with, in relation to or arising out of the use of the Content. This article may be used for research, teaching, and private study purposes. Any substantial or systematic reproduction, redistribution, reselling, loan, sub-licensing, systematic supply, or distribution in any form to anyone is expressly forbidden. Terms &
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3 Journal of the Royal Society of New Zealand, Volume 33, Number 1, March 2003, pp Kakanuiella (gen. nov.) and Thecidellina: Cenozoic and Recent thecideide brachiopods from New Zealand Daphne E. Lee 1 and Jeffrey H. Robinson 1 Abstract Small cemented thecideide brachiopods occur sporadically in Cenozoic strata in New Zealand, Australia, and elsewhere in the south-west Pacific, and living species assigned to the genus Thecidellina are widespread in the subtropical and tropical regions of the Pacific Ocean from northern New Zealand to Japan. We propose a new generic name, Kakanuiella, for the species Thecidellina hedleyi which occurs in limestones and tuffs of Eocene and Oligocene age in New Zealand, including the Chatham Islands. Pliocene specimens of Thecidellina from the Chatham Islands, and Pleistocene and Recent examples from the Kermadec Islands, are assigned to Thecidellina maxilla. The type specimen of the Miocene Thecidium australe, from Muddy Creek, Victoria, Australia, is shown to be Lacazella, rather than Thecidellina. The Recent species Thecidellina japonica from Japan is closely related to T. maxilla, but not to the New Zealand Paleogene species as previously suggested. Keywords brachiopods; thecideides; Pacific; Kakanuiella hedleyi; Thecidellina maxilla; Lacazella australis; Thecidellina japonica; Cenozoic; Recent INTRODUCTION Thecideides are distinctive, diminutive cemented articulate brachiopods of uncertain affinities, now accommodated in a separate Order, Thecideida. They have a long and continuous fossil record and are widely distributed in modern oceans in cryptic habitats in fairly shallow, tropical or subtropical environments. Living thecideides are assigned to one of three genera, Thecidellina, Lacazella, or Pajaudina, based on internal characters. No fossil examples of Pajaudina, described from the Atlantic (Logan 1988), are yet known, but several species of Thecidellina and Lacazella are recorded from Paleocene/Eocene to Pleistocene strata in Europe, the Caribbean region, and the Pacific (Pajaud 1970; Cooper 1979, 1988). Thecideide brachiopods were initially recorded from Tertiary strata in Australia by Tate (1880), and the first living examples from the south-west Pacific were described by Hedley (1899). Thomson (1915) named and illustrated a new Tertiary species from New Zealand which he placed in a new genus Thecidellina together with the type, Thecidium barretti Davidson, a Recent species from the Caribbean, and Hedley's T. maxilla. Pajaud (1970), in a Department of Geology, University of Otago, P.O. Box 56, Dunedin, New Zealand. R02049 Received 29 October 2002; accepted 24 January 2003; published 30 April 2003
4 342 Journal of the Royal Society of New Zealand, Volume 33, 2003 major monograph on the Family Thecideidae, reduced T. hedleyi and the Recent Pacific species T. maxilla, T. congregata, and T. blochmanni to subspecies of the poorly known Australian Miocene species Thecidellina australis" based on "Thecidium" australe Tate, Restudy of the type material of "T." australe Tate from Muddy Creek beds, Victoria, by Neil Archbold, shows that this species belongs to the genus Lacazella. In this paper, we redescribe and illustrate the morphology of Kakanuiella hedleyi (Thomson) living and fossil Thecidellina maxilla (Hedley) from the New Zealand region, and T. japonica (Hayasaka) from Japan. In addition, we briefly comment on the type material of Thecidellina barretti, the identity of Thecidium australe, and the ecology and stratigraphic and geographic distribution of thecidellinin brachiopods through the Cenozoic. Table 1 Locality New Zealand. Age list for Kakanuiella hedleyi (Thomson) and Thecidellina maxilla (Hedley) from Fossil Record File no. Kakanuiella hedleyi (Thomson) Mangaorapan L35/f7537 (Early Eocene) Kaiatan- Ch Is/f512 Runangan (Late Eocene) Runangan J41/ (Late Eocene) f8239 Runangan J41/ (Late Eocene) 9644 Runangan (Late J41/ Eocene) 9645 Whaingaroan J42/ (Early Oligocene) f6147 Whaingaroan (Early Oligocene) Whaingaroan (Early Oligocene) Thecidellina maxilla (Hedley) Pliocene ChIs/ f13 Pliocene ChIs/ f472 Opoitian ChIs/ (Early Pliocene) f490 Early Pleistocene KE/f63 Recent Locality White Creek, Eyre River, North Canterbury Te One Limestone, 400 m N of Moreroa, Te Whanga Lagoon, Chatham Island Ototara Limestone, Trig M, Totara, Kakanui Waiareka Volcanic Formation, Cape Wanbrow, Oamaru Waiareka Volcanic Formation, Cape Wanbrow, Oamaru Ototara Limestone, Everett's Quarry, Kakanui Ototara Limestone, Everett's Quarry, Kakanui Ototara Limestone, Everett's Quarry, Kakanui Whenuataru Tuff, west side Moutapu Pt, Pitt Island Onoua Limestone Flowerpot Bay, Pitt Island Motarata Limestone, Pitt Island m elevation, Napier Island, Kermadec Islands 3 m depth, cave, Dayrell Island, Kermadec Islands Collector(s) P. A. Maxwell P. A. Maxwell D. E. Lee D. E. Lee D. E. Lee D. E. Lee D. E. Lee M. J. S. Rudwick J. A. Thomson P. A. Maxwell P. A. Maxwell F. J. Brook F. J. Brook Number of speci- mens Repositoryremarks GS GS OU OU OU DL22, OU DL24, OU OU Sedgwick Museum, University of Cambridge Museum of NZ, Wellington Holotype Paratypes GS GS OU OU 39461, etc
5 Lee & Robinson Thecideide brachiopods from New Zealand S 170 E 175 E 500 km 180 Kermadec Islands ' 175 W S V. North Island Mi %^\ V V -40 S -45 Si / C J / South 7 Island / _ ^ ^ y Z j / Eyre/River ^Kakanui Chatham Islands < Moutapu Point Fig. 1 New Zealand localities mentioned in the text. MATERIAL AND METHODS - Moreroa Island- More than one hundred new specimens of Cenozoic thecideides from New Zealand were obtained by washing and sieving friable limestone and calcareous tuffs from a number of localities in the Oamaru district. P. A. Maxwell collected specimens of tiny thecideides from Early Eocene beds at Eyre Creek, North Canterbury, and material from several localities on the Chatham Islands. Recent specimens of Thecidellina from the Kermadec Islands were provided by A. G. Beu and F. Brook. New Zealand localities mentioned in the text are shown on Fig. 1, and other locality information is presented in Table 1. T. Habe kindly donated specimens of T. japonica for this study, and comparative material of Thecidellina species was examined at The Natural History Museum, London, and the Australian Museum, Sydney. Additional material from Australia, Vanuatu, and New Zealand was made available on loan from the United States National Museum. All figured specimens were mounted on stubs, coated with Au/Pd, and photographed using a digital camera at the Otago Centre for Electron Microscopy, University of Otago. Most of the specimens referred to in this paper are held in the Department of Geology, University of Otago (prefix OU), the United States National Museum (prefix USNM), and the Institute of Geological & Nuclear Sciences (prefix GS), Lower Hutt. New Zealand fossil localities are referred to by the registered New Zealand Fossil Record File locality number, based on the metric NZMS 260 sheets (e.g., J41/9644).
6 344 Journal of the Royal Society of New Zealand, Volume 33, 2003 SYSTEMATIC DESCRIPTION The classification is based on Baker (1990). Phylum Brachiopoda Class Rhynchonellata Williams et al., 1996 Order Thecideida Pajaud, 1970 Superfamily Thecideoidea Gray, 1840 Family Thecidellinidae Elliott, 1958 Subfamily Thecidellininae Elliott, 1953 Genus Kakanuiella gen. nov. ETYMOLOGY: From the locality for the type species. TYPE SPECIES: Thecidellina hedleyi Thomson, AGE: Eocene-Oligocene. DIAGNOSIS: Small endopunctate subtriangular thecidellinin; ventral valve with a flat, welldefined interarea, large attachment scar and blade-like median septum; dorsal valve with long, straight median septum, interior of subperipheral margins ribbed. REMARKS: Williams (1973) described in detail the secretion and structure of the shell of living and fossil Thecidellina. He noted that the skeletal fabrics of living T. barretti (Davidson) and T. australis (Tate) (in fact T. maxilla (Hedley)) were very similar, but that "both were so different from that of the Oligocene T. hedleyi Thomson from New Zealand as to raise doubts about the taxonomic identity of the last-named species". We concur with his judgement that the "anomalous ultrastructure of the teeth and sockets of hedleyi. and the presence of a medial septum in the ventral valve" suggest that the latter might constitute a new genus (Williams 1973, p. 462). Kakanuiella hedleyi (Thomson, 1915) Fig Thecidellina hedleyi Thomson: 643, fig Thecidellina hedleyi Thomson; Allan, p Thecidellina australis hedleyi Thomson; Pajaud, p. 238, fig. 107A Thecidellina hedleyi Thomson; Keyes, p "Thecidellina" hedleyi Thomson; Williams, pp , fig Thecidellina hedleyi Thomson; Dawson, p. 78. TYPE SPECIMENS: The holotype (Br132) and 2 paratypes (1 entire shell and 1 dorsal valve, Br1439) are held in the collection of Museum of New Zealand Te Papa Tongarewa, Wellington, New Zealand. TYPE LOCALITY: Ototara Limestone (McDonald Limestone of Gage 1957), Everett's Quarry, Kakanui, near Oamaru, New Zealand. AGE: Early Whaingaroan (latest Eocene-earliest Oligocene) (Gage 1957; Edwards et al. 1991; Nelson et al. 2001), rather than Miocene as given by Thomson (1915), and quoted by Pajaud (1970). OTHER MATERIAL: Topotypes are held in the Department of Geology, University of Otago; Museum of New Zealand Te Papa Tongarewa, Wellington; Sedgwick Museum, University of Cambridge; Smithsonian Institution, Washington. REMARKS: Kakanuiella is uncommon in its type locality. Thomson's (1915) original description was based on five specimens, including one dorsal valve with the diagnostic features of the
7 Lee & Robinson Thecideide brachiopods from New Zealand 345 Fig Topotypes of Kakanuiella hedleyi (Thomson, 1915), Late Eocene-Early Oligocene (Early Whaingaroan), Everett's Quarry, Kakanui. Fig. 2-8, interior of ventral and dorsal valve of a topotype; Fig. 2-5, interior views of ventral valve (OU 43119a); Fig. 3, oblique view of hemispondylium; Fig. 4, close-up to show interlocking teeth and sockets and cardinal process (broken off the dorsal valve) resting on hemispondylium; Fig. 5, close-up view of interlocking ribs on subperipheral rim; Fig. 6-8, interior views of dorsal valve (OU 43119b); Fig. 6, dorsal valve with tapering median septum, complete bridge and posteriorly directed spur; Fig. 7, oblique view of same; Fig. 8, close-up of serrated auriform brachial lobes. Fig. 9-11, interior of another topotype; Fig. 9, interior of ventral valve (OU 43120a) with sharp, narrow median septum; Fig. 10, dorsal valve (OU 43120b) with robust cardinal process and broken bridge; Fig. 11, close-up view of small "spines" from valve floor adjacent to septum (cf. Williams 1973, pl. 48, fig. 61).
8 346 Journal of the Royal Society of New Zealand, Volume 33, 2003 genus. Allan (1960, p. 238) commented that the species is so rare at its type locality that he "searched for it for twenty years" before he succeeded in obtaining specimens. However, sieving of friable limestone from the upper section of Everett's Quarry has yielded more than 30 entire shells and separated valves. Most specimens are detached from the substrate, but they are well preserved and not infilled with matrix. In addition to the topotypes, about 100 entire shells and separated, mostly dorsal, valves of K. hedleyi have been collected from thin calcareous horizons interbedded in a thick sequence of unfossiliferous tuffs at Cape Wanbrow, Oamaru, 10 km north of Kakanui. The following descriptions are based on the topotypes, and these marginally older Late Eocene examples from Oamaru. STRATIGRAPHIC AND GEOGRAPHIC DISTRIBUTION: The oldest specimens of K. hedleyi are minute individuals (maximum length 2 mm) from the Early Eocene Eyre Sands, White Creek, Eyre River, North Canterbury. Most specimens are from Late Eocene-Early Oligocene bioclastic limestones and calcareous tuffs in the Oamaru district with a few, rather worn individuals from the Late Eocene of the Chatham Islands (Table 1). DESCRIPTION: The shell outline is commonly subtriangular (Fig. 2, 18, 25), but may be somewhat asymmetric (Fig. 26). None of the specimens examined in the present study exceeds 4 mm in length, which is commonly greater than width. Thomson (1915) gave the dimensions of the type specimens (Table 2). Table 2 Dimensions (mm) of type of Kakanuiella hedleyi (Thomson). Holotype Paratype Paratype Length Width specimens Thickness The measurements are correct but, as Pajaud (1970) noted, the scale provided on Thomson's (1915) figure does not correspond with the measurements given. Ventral valve: The attachment scar occupies between 30 and 90% of ventral valve exterior, depending on initial orientation (i.e., whether plane of commissure is parallel to or at an angle to substrate). In some instances, the scar may reveal the nature of the substrate (e.g., a bryozoan colony in Fig. 26). The interarea is flat and triangular in shape, marked only by subparallel growth lines. There is no trace of a pseudodeltidium or a delthyrial notch. The cardinal margin is straight (Fig. 18, 25) in comparison to the notched margin of Thecidellina maxilla (Fig. 28). The teeth are robust and subtriangular. The hemispondylium consists of slightly concave vertical plates (Fig. 3,4). The pallial cavity is deep, and divided in two by a high, blade-like median septum, extending from the valve floor just anterior of the hemispondylium almost to the anterior border (Fig. 2, 9, 18). Tiny acicular crystallites are present on the floor of the ventral valve on either side of the median septum of some specimens (e.g., Fig. 18, 20, 21) (cf. Williams 1973, fig ). Dorsal valve: The dorsal valve varies in outline from subcircular in small individuals to subtrigonal to subquadrate (Fig. 14, 12, 19). The umbonal region of the dorsal valve is somewhat convex (Fig. 16). The cardinal process is relatively broad, rectangular in outline,
9 Lee & Robinson Thecideide brachiopods from New Zealand µm µm Fig Topotypes of Kakanuiella hedleyi (Thomson, 1915), Late Eocene-Early Oligocene (Early Whaingaroan), Everett's Quarry, Kakanui. Fig , dorsal valve interior of OU 43121, and close-up of ribs on subperipheral rim (note bifurcation); Fig , dorsal valve interior of another topotype (OU 43122) showing large, broad cardinal process, serrated auriform brachial lobes and close-up of marginal ribs (aligned tubercles) in anterior region of median septum; Fig. 16, exterior view of dorsal valve showing convex posterior region and?gastropod drillhole(s); Fig. 17, interior of another dorsal valve (OU 43124) with complete bridge and posteriorly directed spur.
10 348 Journal of the Royal Society of New Zealand, Volume 33, 2003 and projects strongly beyond the cardinal margin (Fig. 10, 19, 22, 27). The median septum is variable in width, but is commonly relatively wide anteriorly, extending about half the valve length and tapering posteriorly. It consists of a central shallow sinus flanked by two anteriorly diverging ridges which carry small spiny/serrated processes. The bridge is narrow and delicate, with a small posteriorly directed spur (marsupial notch) (Fig. 6, 17, 24). Two welldeveloped auriform brachial lobes with serrated margins unite at the posterior end of the septum (Fig. 6-8). The anterior margins of the brachial valves are characteristically ribbed or fluted, with the ribs at times bifurcating (Fig. 12, 13). SHELL MICROSTRUCTURE: Williams (1973) described in detail the secretion and structure of the shells of living and fossil thecideidine brachiopods, and commented on the apparently "anomalous ultrastructure of the teeth and sockets of hedleyi as a species of Thecidellina". Williams (1973, p. 461) noted that although the shells had undergone some recrystallisation, the original fabric was not obliterated, and "consisted of a primary layer composed mainly of acicular crystallites about 700 (Am thick.. Interlocking granules of calcite up to 12 µm in size also occur especially in the teeth, the core of the cardinalia and the areas of muscle attachment". He noted that the absence of secondary shell from the teeth and sockets of hedleyi was unlikely to be due to size differences, as smaller, immature examples of Thecidellina barretti had well-developed secondary fibres. DISCUSSION: Kakanuiella differs from modern species of Thecidellina in possessing a welldefined blade-like median septum in the ventral valve and in having marginal ribs rather than individual scattered tubercles in the subperipheral margin. Kakanuiella shares many morphological characters with Eothecidellina imperfecta (Nekvasilova) from the Upper Cretaceous of Czechoslovakia (Baker 1991). It differs in its overall subtriangular shape, the presence of the ventral septum, the shape of the brachial lobes, and the marginal ribbing. While we found no canopied brachial lobes as reported for Eothecidellina and Bifolium (Baker 1991) (and illustrated in Thecidellina minuta by Cooper 1981) in any of the specimens of Kakanuiella which we have examined, the serrated margins on the brachial lobes (e.g., Fig. 6-8) may represent the remnants of these structures which provided further support for the lophophore. Genus Thecidellina Thomson, 1915 (= Thecidellella Hayasaka, 1938 (type species Thecidellella japonica Hayasaka, 1938: 9).) TYPE SPECIES: Thecidium barretti Davidson, 1864 (original designation). AGE: Recent. DIAGNOSIS: Small endopunctate subrounded to subtriangular thecidellinin; ventral valve with a flat, triangular interarea, small to medium attachment scar, no median septum in ventral valve; dorsal valve with long straight median septum, interior of subperipheral margin strongly tuberculate; lophophore schizolophous. REMARKS: The genus Thecidellella was said to differ from Thecidellina in the possession of a "reversed spondylium" (Hayasaka 1938, p. 10) but Cooper (1954) pointed out that this feature was present in all species of Thecidellina. DISCUSSION: NO review of species assigned to Thecidellina has been carried out since the comprehensive study by Pajaud (1970), apart from a brief account by Zezina (1985) of the living species. Since information on the type material of several species (including T. barretti) was lacking, and this paper provides new illustrations of T. maxilla and T. japonica, we provide a brief chronological survey of species assigned to the genus Thecidellina.
11 Lee & Robinson Thecideide brachiopods from New Zealand Fig Kakanuiella hedleyi (Thomson, 1915), Late Eocene (Runangan), Waiareka Volcanic Formation, Cape Wanbro w, Oamaru. Fig. 18, interior of ventral valve (OU 43125a) showing transverse growth lines of interarea, straight cardinal margin, and narrow median septum; Fig. 19, dorsal valve interior (OU 43125b) showing wide cardinal process; Fig , close-up view of acicular crystallites on ventral valve floor of previous specimen; Fig , dorsal valve interior (OU 43126) and close-up of aligned tubercles; Fig. 24, dorsal interior of slightly asymmetric specimen (OU 43127) with complete bridge and posteriorly directed spur; Fig. 25, ventral valve interior (OU 53128); Fig. 26, large attachment scar of specimen which was cemented to a bryozoan colony (OU 43129); Fig. 27, dorsal valve with broad septum (OU 43130).
12 350 Journal of the Royal Society of New Zealand, Volume 33, 2003 Thecidellina barretti (Davidson, 1864) 1864 Thecidium barretti Davidson: 17, pl. 2, fig Thecidellina barretti (Davidson); Thomson, p Thecidellina barretti (Davidson); Pajaud, pp , pl. VIII, fig. 8; fig. 19, Thecidellina barretti (Davidson); Cooper, p. 132, pl. 3, fig , Thecidellina barretti (Davidson); Zezina, p TYPE SPECIMENS AND TYPE LOCALITY: The specimen collected by Lucas Barrett from the north-east coast of Jamaica at a depth of 60 fathoms (c. 110 m), figured by Davidson (1864, pl. II, fig. 1a) and preserved as two separated valves (I 6801) in the collections of the Department of Zoology, University of Cambridge, England, is here designated as lectotype. The second syntype (ZB4260) in the Davidson Collection, The Natural History Museum, London, has not been located. These brachiopods had a tragic provenance. Barrett arrived in Jamaica in In March 1862 he returned to England and gave S. P. Woodward Recent brachiopods from Jamaica. In December 1862, Barrett died in a diving accident in Jamaica, while collecting further brachiopod specimens (Sarah Long pers. comm. 2000). Subsequently, Woodward asked Davidson to describe the brachiopods collected by Barrett, and accordingly Thecidium barretti was named by Davidson (1864) with illustrations of the dorsal view of a Recent specimen, and the interior view of one Recent and one fossil example of T. barretti (the latter presumably from the "newest Pliocene beds" of Jamaica (Davidson 1864, pl. II, fig. 1-3)). Note that these figures are all reversed. The lectotype (I6801) consists of the separated valves of one individual with dimensions of 4.1 x 4.1 mm, and is the specimen drawn by S. P. Woodward and figured by Davidson (1864) as pl. II, fig. 1. Both syntypes were redrawn and refigured in correct orientation by Davidson (1887, pl. 23, fig. 10). DISTRIBUTION: Thecidellina barretti is widespread in the Caribbean region (Jackson et al. 1971; Cooper 1977). STRATIGRAPHIC RANGE: Pliocene-Recent. DESCRIPTION: Thecidellina barretti is described and illustrated by Pajaud (1970) and Cooper (1977). Thecidellina maxilla (Hedley, 1899) Fig Thecidea maxilla Hedley: 508, fig Thecidellina maxilla (Hedley); Dall, p Thecidellina maxilla (Hedley); Thomson, p Thecidellina maxilla (Hedley); Cooper, p. 317, pl. 81, fig Thecidellina cf. T. maxilla (Hedley); Cooper, p. 1118, pl. 301, fig , Thecidellina australis maxilla (Hedley); Pajaud, p. 240, fig. 22, 107B Thecidellina maxilla (Hedley); Zezina, p TYPE SPECIMENS AND TYPE LOCALITY: Australian Museum, Sydney. The holotype (C.6202) and paratypes (C ex C.6202, C68588) were collected by Charles Hedley in 1896, from west slope of Funafuti Atoll, Tuvalu [formerly Ellice Islands], South Pacific Ocean, from a sheet of dead coral, fathoms ( metres). (Other specimens labelled "cotypes" by Hedley are now regarded as paratypes (Ian Loch pers. comm. 2003).) DISTRIBUTION: Thecidellina maxilla is widespread in the Pacific Ocean. It is reported from Tuvalu (Ellice Islands); Torres Strait in 5-8 fathoms (9-15 m) (Thomson 1927); Vanuatu (New Hebrides), m (USNM NHE185A); Bikini Atoll, Gambier, Tuamotu Island (Pajaud 1970); and as far south as the Kermadec Islands (Fig. 1) in a cave at c. 3 m depth, and
13 Lee & Robinson Thecideide brachiopods from New Zealand Fig Thecidellina maxilla (Hedley, 1899), Recent, 3 m, Kermadec Islands. Fig. 28, interior of ventral valve of adult specimen (OU 43131a) showing notched cardinal margin, hemispondylium, absence of median septum, finely punctate and tuberculate valve floor, and tuberculate subperipheral margin; Fig. 29, dorsal valve of same specimen (OU 43131b), showing trilobed cardinal process with bevelled edges; small posteriorly directed spur, and dried, bilobed schizolophous lophophore; Fig. 30, close-up of anterior region of median septum showing randomly arranged tubercles (cf. Fig. 15); Fig. 31, interior of another dorsal valve (OU 43132) with most of lophophore removed, showing large auriform brachial lobe processes and row of lophophore muscle scars; Fig , interior of dorsal (OU 43133b) and ventral valve (OU 43133a) of juvenile specimen; Fig. 32, note relatively large marginal tubercles, and lophophore attachment scars; Fig. 33, note short interarea, the twin prongs of the hemispondylium, lophophore attachment scars, and few subperipheral tubercles.
14 352 Journal of the Royal Society of New Zealand, Volume 33, 2003 as dead shells in shell gravel at 30 m and 40 m depth at the base of submarine cliffs on Raoul Island, Kermadec Group (Fred Brook pers. comm. 1989, 2002) (Table 1). STRATIGRAPHIC RANGE: Miocene to Recent. Cooper (1964) recorded Thecidellina cf. T. maxilla from post-miocene drill cores on Eniwetok and Rongelap, Marshall Islands. The specimens of Thecidellina spp. figured by Cooper (1978) from the Early Miocene of Java, the Miocene of Fiji, and the Pleistocene of Vanuatu (New Hebrides) all resemble Thecidellina maxilla. Specimens of Pliocene age from Whenuataru Tuff, Moutapu Point, Pitt Island (GS 12163) are here assigned to Thecidellina maxilla (Hedley). Specimens attached to in situ hermatypic corals occur in uplifted Pleistocene beds on Raoul Island, Kermadecs (Brook 1998a). DESCRIPTION: The specimens described and figured herein were collected by SCUBA from about 3 m depth in a cave, west side of Dayrell Island, Kermadec Islands (Fred Brook pers. comm. 1989). About 150 specimens ranging in size from 0.6 mm to 4.1 mm are cemented to a small basalt boulder (120 x 60 x 70 mm). T. maxilla may reach 6 mm in length (Hedley 1899). Typical dimensions are given in Table 3. Juveniles are nearly circular in outline (Fig. 32, 33) and are cemented to the substrate by the entire ventral valve. As animals grow, the ventral valve Table 3 Dimensions (mm) of ma y lift off the substrate, and the shape of the specimens of Thecidellina maxilla (Hedley) from Kermadec Islands. Length Width adult shell will vary depending on the initial orientation and the irregularity of the underlying rock or shell. Young specimens are semitranslucent, and the cemented ventral valve floor even in adults is thin and transparent. Adult shells are white or semitranslucent in colour, and subcircular to subpentagonal in outline, although considerable variation is present. The anterior commissure is rectimarginate. The cardinal margin is straight in juvenile specimens (Fig. 33), and generally so in adults, but may appear to be notched due to damage as valves are prised apart (Fig. 28). The interarea is short and triangular, with transverse striations. There is no trace of a pseudodeltidium. The marginal border is wide and finely tuberculate. The ventral cavity is deep, with no median septum. The hemispondylium is cup-shaped with two slender prongs extending anteriorly just past the cardinal margin (Fig. 33), as shown by Hedley (1899, fig. 57). Dorsal valve is subcircular, cardinal process is trilobed, with bevelled corners (Fig. 29, 31, 32). Tubercles on subperipheral rim extend onto anterior median septum. Posterior projecting spur is conspicuous on juvenile specimens (Fig. 32). Lophophore is schizolophous (Fig. 29, 32). Pliocene specimens from the Chatham Islands (Fig. 34, 35) closely resemble T. maxilla from Kermadec Islands.
15 Lee & Robinson Thecideide brachiopods from New Zealand " * Fig Thecidellina maxilla (Hedley, 1899), Pliocene, Chatham Islands. Fig. 34, interior of dorsal valve showing trilobed cardinal process with bevelled corners, worn/ damaged brachial lobes, and conspicuous randomly arranged subperipheral tubercles; Fig. 35, close-up of anterior region of median septum. Fig Thecidellina japonica (Hayasaka, 1938), Recent, 183 m, Tosa Bay, Shikoku, Japan (OU 11974a). Fig. 36, interior of ventral valve, showing notched cardinal margin, transverse growth lines of interarea, absence of median septum, parallel-sided hemispondylium, and finely tuberculate subperipheral rim; Fig. 37, interior of dorsal valve, showing small, trilobed cardinal process with bevelled corners, posteriorly directed spur, dried schizolophous lophophore with long, delicate paired tentacles; Fig. 38, close-up of anterior region of median septum showing small punctae and randomly arranged subperipheral tubercles.
16 354 Journal of the Royal Society of New Zealand, Volume 33, 2003 Thecidellina blochmanni Dall, Thecidellina blochmanni Dall: Thecidellina australis blochmanni Dall; Pajaud, p. 245, fig. 110, pl. II, fig Thecidellina blochmanni Dall; Zezina, p Thecidellina blochmanni Dall; Cooper, p. 8, pl. 8, fig TYPE SPECIMENS AND TYPE LOCALITY: The holotype in the United States National Museum (USNM ) was figured by Cooper (1973). The specimen was collected from a depth of 84 m from Christmas Island, Indian Ocean (10 S, 'E), not Christmas Island, Pacific Ocean as indicated by Pajaud (1970). STRATIGRAPHIC RANGE: Recent. DESCRIPTION: Dall (1920) differentiated his new species from T. maxilla by stating that it "is much more regular, the internal arrangements symmetrical, bilaterally identical, and elegant". Pajaud (1970), mistakenly believing that the type of T. blochmanni had been collected from the Pacific Ocean, may have inadvertently described specimens which represent T. maxilla sens. str. Thecidellina japonica (Hayasaka, 1938) Fig Thecidellella japonica Hayasaka: 9-13, fig Thecidellella japonica Hayasaka; Hatai, pp , fig Thecidellina japonica (Hayasaka); Elliott, H Thecidellina japonica (Hayasaka); Pajaud, pp , fig Thecidellina japonica (Hayasaka); Zezina, p TYPE LOCALITY AND DISTRIBUTION: Pajaud (1970) noted that the type specimens, collected from the Hatizyo-zyma region (c. 33 N, 140 E) (Hayasaka 1938), were lost during World War II. Tadashige Habe (pers. comm. 1981) mentioned that other material has been collected by deep-sea trawl from a few localities off southern Japan including Sagami Bay at about 150 m, and Tosa Bay, Shikoku, in 183 m (100 fathoms), and these specimens are held in the collections of the National Science Museum, Tokyo. Three specimens from the former locality were presented to the University of Otago, and are catalogued as OU 11974a-c. DESCRIPTION: Shell small (maximum length observed 5.1 mm). The dimensions of the specimens held by the University of Otago are given in Table 4. Ventral valve thick and geniculate, colour greyish white, growth lines well marked, outline nearly circular, with transversely striated triangular interarea; cardinal margin with small notch (Fig. 36); teeth large, triangular. Ventral cavity deep, with no medium septum (in contrast to Hayasaka 1938, fig. 2b). Hemispondylium consisting of anteriorly diverging flat, vertical plates. Dorsal valve subcircular, posterior cardinal border short, straight; cardinal process trilobed, strongly projecting, with bevelled corners; marginal border wide, finely Table 4 Dimensions (mm) of specimens of Thecidellina japonica (Hayasaka) from Japan. Length Width Thickness
17 Lee & Robinson Thecideide brachiopods from New Zealand 355 tuberculate (Fig. 37, 38), tubercles larger on inner margin, and interspersed with punctae. Median septum long; lophophore filaments are very long. REMARKS: The illustration provided by Hayasaka (1938) gives the impression of a poorly defined "septum" in the ventral valve, but this structure is not present in the specimens we have seen. Williams (1973) noted that the presence of a septum in the ventral valve of the Recent T. japonica might constitute a link between this genus and T. hedleyi. Our study demonstrates that the Japanese species is morphologically similar to other Pacific Thecidellina, and is closely related to T. maxilla, with which it may be conspecific. Thecidellina congregata Cooper, Thecidellina congregata Cooper: 316, pl. 80, fig Thecidellina australis congregata Cooper; Pajaud, p. 241, fig. 108, 109, pl. XIII, fig. 5, Thecidellina congregata Cooper; Zezina, p TYPE SPECIMENS AND LOCALITY: The holotype from Bikini Atoll, Marshall Islands, is held in the United States National Museum (USNM ). DISTRIBUTION: Central Pacific Ocean, Marshall Islands (Cooper 1954); Guam, Saipan (Thayer & Allmon 1991). STRATIGRAPHIC RANGE: Recent. DESCRIPTION: Cooper (1954) distinguished T. congregata from T. maxilla which he recorded from the same localities on the basis of colour, smaller size, and shape of the median septum. Thecidellina minuta Cooper, Thecidellina minuta Cooper: 61-62, pl. 6, fig Thecidellina minuta Cooper; Zezina, p TYPE SPECIMENS AND TYPE LOCALITY: The holotype from south of Madagascar is held in the Muséum National d'histoire Naturelle, Paris (MNHN-BRA-78-74c). DISTRIBUTION: Indian Ocean, around Madagascar (Cooper 1981). STRATIGRAPHIC RANGE: Recent. DESCRIPTION: Cooper (1981) distinguished T. minuta from T. blochmanni which he recorded from the same localities on the basis of shape, smaller size, and lack of a broad, interior peripheral margin. Family Thecideidae Gray, 1840 Subfamily Lacazellinae Backhaus, 1959 "Thecidium australe" Tate, 1880 = Lacazella australis (Tate, 1880) 1880 Thecidium australe Tate: 166, pl. ix, fig. 3a-c Thecidellina australis australis (Tate); Pajaud, p REMARKS: The earliest record of a thecideide brachiopod from the Cenozoic of the Southern Hemisphere was the description of Thecidium australe by Tate (1880) from the "Superior beds", Muddy Creek, Victoria, Australia. Tate's (1880) description and poor lithographs which appear to show an incomplete foramen were based on external features of one complete shell and four ventral valve interiors. Thomson (1915, 1927) and Crespin & Chapman (in Thomson 1927) suggested that Tate's specimens were likely to be Lacazella. Pajaud (1970) doubted this attribution since the former genus was apparently restricted to the Mediterranean region, and suggested instead that it was likely to belong to the genus
18 356 Journal of the Royal Society of New Zealand, Volume 33, 2003 Thecidellina, and might be identical to T. hedleyi from the Miocene of New Zealand. Unfortunately, based on this assumption and without further study of Thecidium australe, Pajaud (1970) then reduced all living and fossil Pacific species of Thecidellina to subspecies of "Thecidellina australis". Tate's type material, held in the South Australian Museum (Adelaide), was re-examined by Neil Archbold, who noted that most of the specimens were somewhat abraded and the interiors of some ventral valves were encrusted with minute serpulid worms. Ventral valve length varied from mm. "Shape variable, subcircular in smaller specimens, elongate in larger specimens. Attachment scar well-developed, irregular in shape. Ventral valve with raised pseudodeltidium. Outer margins of ventral interarea poorly defined. Cardinal teeth robust. Hemispondylium well developed. No median septum. Cardinal process broad, projects strongly anteriorly" (N. Archbold pers. comm. 1989). Examination of other Muddy Creek material borrowed from the United States National Museum by the senior author in 1991 caused further confusion, as examples of both Thecidellina (USNM ) and Lacazella (USNM ) were present. However, N. Archbold (pers. comm. 1992) succeeded in opening the double-valved lectotype of Thecidium australe and confirmed that this species is indeed Lacazella. Further work on better preserved Muddy Creek material is needed to establish the relationships of the species of Thecidellina present in the same fauna. AGE: The Muddy Creek beds are now regarded as "marginal members of the Port Campbell Limestone" (Abele et al. 1976), and are of early-middle Miocene (Balcombian) age. HOW MANY SPECIES OF THECIDELLINA ARE PRESENT IN MODERN OCEANS? To date, six species of Thecidellina have been described from the Caribbean, Pacific, and Indian Oceans. The geographic distributions of at least two of these putative species overlap completely (T. maxilla with T. congregata; T. blochmanni with T. minuta), and the main points of difference between these species pairs are based either on adult size, shape, and size of attachment scar, or on features which change during ontogeny such as presence of a welldefined internal subperipheral margin (compare the interior characters of a young specimen of T. maxilla (Fig. 32, 33) with an adult example (Fig. 28, 29)). It is valid to use size as a taxonomic discriminant only when complete ontogenetic stages are known for the species in question. Given that morphological variation within the living (and fossil) species of Thecidellina is considerable, and the features on which some species have been defined (colour, size, shape) are minor, and/or dependant on the relief of the attachment substrate, we suggest that morphological and ontogenetic studies of large populations, coupled with molecular genetic methods, are needed to elucidate the true relationships of these little brachiopods. It is possible that all Indo-Pacific species are variants of Thecidellina maxilla. ECOLOGY AND PALEOECOLOGY Water depth and substrate Most species of Thecidellina occupy nearshore reefal environments at depths ranging from subtidal (1-3 m) down to a maximum of about 150 m (there is a single record from c. 480 m (Cooper 1954), although this latter record may represent a redeposited block of coral). Density appears to be greatest in dark, cryptic environments (caves, tunnels, deep recesses, crevices, undersides of foliaceous corals) between 10 and 60 m (Jackson et al. 1971; Grant 1987; Thayer & Allmon 1991) although this may partly reflect the fact that most specimens are collected by diving. Substrates include rock and coral.
19 Lee & Robinson Thecideide brachiopods from New Zealand 357 Most of the specimens of T. maxilla from the Kermadecs, collected from a dark cave at 3 m depth, are cemented directly, in random orientations, on to a basalt cobble, although a few are attached to serpulids or to both live and dead eroded shells of conspecifics. Kakanuiella hedleyi occupied similarly shallow water depths, and is assumed to have attached to either rock or shell as a few specimens are still cemented to larger brachiopod shells. Temperature Thecidellina is a well-documented warm-water indicator, and all living records are from the tropical and subtropical Indo-Pacific or Caribbean. Pajaud (1970) suggested that the "valeurs moyennes pour les eaux où vivent les actuels Thecidellina sont situées autour de 23 C". The sea surface temperature at the northern end of the Kermadec Islands (c. 29 S) ranges from 16 to 26 C (mean c C): this is the most southern (and coolest) site from which Thecidellina species have been reported. The fauna associated with Thecidellina includes a range of subtropical molluscs such as Spondylus, Cypraea, Chama, Bursa, Morum, Conus, and Distorsio (A. G. Beu pers. comm. 1989; Brook 1998b) and about 15 genera of hermatypic (zooxanthellate) corals, although there is no reef as such (Brook 1999). The New Zealand records of Kakanuiella in the Early and Late Eocene-earliest Oligocene, and Thecidellina in the Early Miocene of Northland (Hiller 2000), and Pliocene of Chatham Islands correspond closely with periods of subtropical sea temperatures in the New Zealand region (Hornibrook 1993). The Pliocene record from the Chatham Islands (at 44 S, some 14 further south than the Kermadec Islands) may coincide with a major global warming episode, and we suggest here that the 20 C isotherm may have migrated as far south as the Chatham Islands during Pliocene times. Reproduction and dispersal mechanisms Little has been published on reproduction and dispersal in Thecidellina species. According to James et al. (1992), T. barretti is probably a hermaphrodite, its larvae are lecithotropic, and the species possesses a brood chamber. The latter characters are shared by the other very small reef-dwelling brachiopods which commonly occur with Thecidellina in both modern oceans and in the fossil record, i.e., Lacazella and Argyrotheca, although Lüter (pers comm. 2002) noted that Lacazella exhibits sexual dimorphism. Jackson et al. (1971) considered that distinct bimodality in populations of Thecidellina from both the Caribbean and Pacific suggested a single annual spawning. Our small population sample from the Kermadec Islands is less conclusive, with all size ranges present. Thecideide brachiopods may occur in very dense numbers in suitable environments. Logan (1977) reported a density for T. barretti on Grand Cayman Island equivalent to 4600 m 2 in a deep recessed cave at 12 m depth with almost no light or water movement. Jackson et al. (1971) noted similar densities of live and dead Thecidellina in dark caves at shallow depths in the Pacific. Biotic associations The cemented thecideides with very similar external form, Thecidellina and/or Lacazella, and one or more very small megathyrid genera such as Argyrotheca, and/or small cemented craniids "seem to constitute a suite of brachiopods that characterizes the shallow waters of tropical islands" (Grant 1987, p. 79), and are often found together in the fossil record (e.g., Meile & Pajaud 1971), as far back as the Eocene in the Pacific (Cooper 1971). Grant (1987, p. 79) commented that "the cryptic environment produced by foliaceous corals has been exploited [by these small brachiopods] for a long time".
20 358 Journal of the Royal Society of New Zealand, Volume 33, 2003 Thecidellina and Lacazella occur together today in the Caribbean (Jackson et al. 1971) and in the Pacific in Palau (Thayer & Allmon 1991) and Vanuatu (this paper), and as fossils in the Paleocene and Eocene of Alabama (Cooper 1979, 1988), the Eocene and Miocene of Cuba, the Miocene of Australia (this paper) and Java (Cooper 1978), and the Pleistocene of Vanuatu (Cooper 1978). However, Lacazella has not yet been recorded from any New Zealand localities and Thecidellina no longer occurs in the Mediterranean Sea, although it has a long history in the region (Pajaud & Tambareau 1970). The reasons for their absence are uncertain but may be related to sea temperatures and/or historical factors. In the New Zealand Late Eocene-earliest Oligocene, Kakanuiella hedleyi is moderately common in a shallow-water "brachiopod coquina" near Kakanui and in several thin calcareous tuffaceous horizons within a thick unfossiliferous volcanic succession at Oamaru. Only a few individuals are still in life position, cemented to larger brachiopods, but attachment scars (e.g., Fig. 26) suggest that bryozoans and echinoid spines also provided substrates. Numerous other small brachiopods (<10 mm), including Argyrotheca, juveniles of Terebratulina, Tegulorhynchia, and Novocrania are present, together with large foraminifera, bryozoans, echinoids, and some larger brachiopods and molluscs (including Conus). A few specimens show evidence of predation, probably by small gastropods (e.g., Fig. 16). These communities probably represent shallow, warm-water communities living on the flanks of small submarine volcanoes, but which were subsequently redeposited in somewhat deeper water (Lee et al. 1997). Distribution of Thecideide brachiopods Kakanuiella is known only from the New Zealand region where it ranges from Early Eocene to Early Oligocene. Thecidellina has a cosmopolitan distribution, being recorded from Europe (Spain and France) during the Paleocene (Pajaud & Plaziat 1972; Pajaud & Tambareau 1970), Alabama (Toulmin 1940; Cooper 1979), and from the Caribbean (Cuba) (Cooper 1979) and the Pacific ('Eua, Tonga) (Cooper 1971) in the Eocene. Forms closely related to modern species appear in Cuba (Cooper 1979), and in the south-west Pacific, including Australia and New Zealand, during the Miocene. CONCLUSIONS We have redescribed and illustrated a number of fossil and living thecideide brachiopods from the New Zealand and Indo-Pacific region. Kakanuiella hedleyi from the Paleogene of New Zealand shares some morphological characters with Late Cretaceous thecideides from Europe and some with modern species of Thecidellina. Both Thecidellina and Lacazella are reported from the Miocene of Australia, although the latter genus is not yet recorded from New Zealand. Thecidellina japonica from Japanese seas is shown to be closely related to T. maxilla from the Pacific. Although rarely reported, these small cryptic thecideide brachiopods are widespread in the Pacific and can be extremely abundant on hard substrates in shallow, cryptic environments. The long history of these warm-water brachiopods and the close associations between small thecideides, megathyrid, and craniid brachiopods point to ecological stability of these environments for at least the past 50 million years, and perhaps back to the Cretaceous and Jurassic (Pajaud 1970, 1974a,b). We suggest that the geographic distribution of thecideides such as Thecidellina and Kakanuiella during the Cenozoic was controlled largely by fluctuations in sea temperature and that, conversely, the presence of these tropothermal taxa in the fossil record can be used to postulate mean sea surface temperatures of at least 20 C for the sites at which they occur.
21 Lee & Robinson Thecideide brachiopods from New Zealand 359 We conclude that although some aspects of thecideide systematics and relationships and ecology/paleoecology are reasonably well understood, there is need for further research to clarify relationships of modern and fossil species and establish dispersal mechanisms. ACKNOWLEDGMENTS This paper is dedicated to two people with a passionate interest in geology, Doug Campbell and Shelagh Gorringe, both members of the Department of Geology, University of Otago, who died in July 2001 and August 2001, respectively. Daniel Pajaud gave considerable assistance to the senior author during a visit to Paris in Neil Archbold helped with information on Thecidium australe; Phil Maxwell collected all of the specimens from Eyre Creek, and much of the Chatham Island material. Alan Beu and Fred Brook provided the specimens of Thecidellina from the Kermadecs. Sarah Long provided information on the type material of T. barretti and Hilary Potter provided access to specimens held in the Department of Zoology, University of Cambridge. The late Richard Grant and Mark Florence made it possible for DEL to borrow thecideides from Australia, Vanuatu, and New Zealand from the USNM. Ian Loch and Janet Waterhouse of the Australian Museum, Sydney, provided information on the type specimens of T. maxilla, and Bruce Marshall, Museum of New Zealand Te Papa Tongarewa, confirmed details on the types of T. hedleyi. We also wish to thank Carsten Lüter for providing a preview copy of the paper describing a new thecideide genus, and commenting on a draft of this paper. JHR wishes to acknowledge assistance from Mark Gould and Liz Girvan at the Otago Centre for Electron Microscopy, University of Otago. Funding for this study was provided by the Division of Sciences, University of Otago. REFERENCES Allan, R. S. 1960: The succession of Tertiary brachiopod faunas in New Zealand. Records of the Canterbury Museum 7(3): Abele, C.; Kenley, P. R.; Holdgate, G.; Ripper, D. 1976: Otway Basin. In: Douglas, J. D.; Ferguson, J. A. ed. Geology of Victoria. Geological Society of Australia Special Publication 5: Baker, P. G. 1990: The classification, origin and phylogeny of thecideidine brachiopods. Palaeontology 33: Baker, P. G. 1991: Morphology and shell microstructure of Cretaceous thecideidine brachiopods and their bearing on thecideidine phylogeny. Palaeontology 34(4): Brook, F. J. 1998a: Stratigraphy and paleontology of Pleistocene submarine volcanic sedimentary sequences at the northern Kermadec Islands. Journal of the Royal Society of New Zealand 28: Brook, F. J. 1998b: The coastal molluscan fauna of the northern Kermadec Islands, Southwest Pacific Ocean. Journal of the Royal Society of New Zealand 28: Brook, F. J. 1999: The coastal scleractinian coral fauna of the Kermadec Islands, southwestern Pacific Ocean. Journal of the Royal Society of New Zealand 29: Cooper, G. A. 1954: Recent brachiopods in Bikini and nearby atolls, Marshall Islands. United States Geological Survey Professional Paper 260G: Cooper, G. A. 1964: Brachiopods from Eniwetok and Bikini Drill Holes. United States Geological Survey Professional Paper 260FF: Cooper, G. A : Eocene brachiopods from Eua, Tonga. United States Geological Survey Professional Paper 640F: F1-F9. Cooper, G. A. 1973: New Brachiopoda from the Indian Ocean. Smithsonian Contributions to Paleobiology p. Cooper, G. A. 1977: Brachiopods from the Caribbean Seas and adjacent waters. Studies in Tropical Oceanography 14: Cooper, G. A. 1978: Tertiary and Quaternary brachiopods from the Southwest Pacific. Smithsonian Contributions to Paleobiology p. Cooper, G. A. 1979: Tertiary and Cretaceous Brachiopods from Cuba and the Caribbean. Smithsonian Contributions to Paleobiology p. Cooper, G. A. 1981: Brachiopods from the Southern Indian Ocean. Smithsonian Contributions to Paleobiology p. Cooper, G. A. 1988: Some Tertiary Brachiopoda of the East Coast of the United States. Smithsonian Contributions to Paleobiology p.
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23 Lee & Robinson Thecideide brachiopods from New Zealand 361 Thayer, C. W.; Allmon, R. A. 1991: Unpalatable thecideid brachiopods from Palau: Ecological and evolutionary implications. In: MacKinnon, D. I.; Lee, D. E.; Campbell, J. D. ed. Brachiopods through time. Proceedings of the Second International Brachiopod Congress, University of Otago, Dunedin, New Zealand. Pp Thomson, J. A. 1915: On a new genus and species of the Thecidiinae. Geological Magazine Decade VI, 2: Thomson, J. A. 1927: Brachiopod morphology and genera (Recent and Tertiary). New Zealand Board of Science and Art Manual p. Toulmin, L. D. 1940: Eocene brachiopods from the Salt Mountain limestones of Alabama. Journal of Paleontology 14: Williams, A. 1973: The secretion and structural evolution of the shell of thecideidine brachiopods. Philosophical Transactions of the Royal Society of London B, Biological Sciences 264: Zezina, O. N. 1985: Living brachiopods and problems of the bathyal zone in the oceans. Akademiia Nauk SSSR. 247 p. (In Russian)
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