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Palaeoworld 15 (2006) 135–149

Research paper

Forty years since Boucot, Johnson and Staton’s seminal paper“On some atrypoid, retzioid, and athyridoid Brachiopoda”

Fernando AlvarezDepartamento de Geologıa, Universidad de Oviedo, Oviedo, Spain

Received 30 April 2005; received in revised form 31 March 2006; accepted 7 July 2006Available online 1 September 2006

bstract

The rate at which new spire-bearing brachiopods, exclusive of spiriferids, have been described, their temporal and geographiciversity, and their grade of endemism through time are reviewed. There has been an almost four-fold increase in the number ofpire-bearing brachiopod genera (spiriferids excluded) included in the revised edition of the Treatise compared with those recognizedn the first edition. Many of these genera were based on specimens from previously poorly known geologic successions, especiallyn Russia and China where Chinese and Russian colleagues had undertaken admirable and hard research. Even assuming a relativelyigh proportion of invalid or synonymous genera, the annual rate of creation of genera has been five to seven times that prior toublication of the first edition. Spire-bearing brachiopod genera erected after the first edition of the Treatise show a greater degree ofndemicity than those erected prior to its appearance, with endemism displayed by the athyridids being greater than for the atrypids.o correlation seems to exist between degree of endemicity and generic diversity at any specific time. Endemic spire-bearers aref great value in discriminating biogeographic units. Spire-bearing genera with cosmopolitan tendencies did not evolve rapidly, soheir value for global stratigraphic correlations is generally poor. The large increase in generic-level taxa appears to have resulted

n a cascading expansion in numbers of higher taxa. In addition to the remarkable increase in the number of taxa during the lastour decades, there has been an obvious shift in brachiopod research with increasing integration of other disciplines (biological,alaeontologic, mineralogic), which is improving our understanding of spire-bearers, and of the phylum as a whole in time and space.

2006 Nanjing Institute of Geology and Palaeontology, CAS. Published by Elsevier Ltd. All rights reserved.

Palaeo

eywords: Brachiopoda; Atrypida; Athyridida; Spatio-temporal data;

. Introduction

Forty years have elapsed since Boucot, Johnsonnd Staton’s paper “On some atrypoid, retzioid, andthyridoid Brachiopoda”, a condensed preview, exclu-ive of the spiriferids, of their contribution to Part

(1965) of the Treatise on Invertebrate Paleontol-gy, the reference point for subsequent works on thepire-bearing brachiopods (Boucot et al., 1964, 1965).

E-mail address: fernando@geol.uniovi.es.

871-174X/$ – see front matter © 2006 Nanjing Institute of Geology and Paldoi:10.1016/j.palwor.2006.07.006

zoic

These works, together with the Russian “Osnovy Pale-ontologii” were the first modern attempts to provide acomprehensive classification, rather than listing previ-ously defined genera (Waagen, 1883; Hall and Clarke,1894, 1895; Schuchert, 1896; Zittel, 1903; and the syn-opses of Schuchert, 1929; Muir-Wood, 1955). Boucot’sfive decades of voluminous publication continues relent-lessly.

The present paper is focused on study of the rate atwhich new genera of spire-bearing brachiopods, exclu-sive of the spiriferids, were proposed, their temporaland geographic diversity, and their degree of endemism

aeontology, CAS. Published by Elsevier Ltd. All rights reserved.

world 15 (2006) 135–149

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through time. Rate of proposal of genera, distribution anddegree of endemism are probed for the spire-bearing gen-era revised by Boucot et al. (1965) and those proposedsubsequently. Athyridid and atrypid taxa are analysedseparately and in combination. Problems that remain tobe resolved are highlighted, as are perspectives for futureresearch on spire-bearing brachiopods.

2. Data

The data for this study are derived mainly from Atryp-ida and Athyridida generic databases revised by Boucotet al. (1965) for the first edition of the Treatise, and morerecently by Copper (2002), Alvarez and Rong (2002),and Alvarez and Copper (2002) for the revised edition ofthe Treatise (2002) and the Supplement (in press; see alsoCopper, 2004a; Alvarez, 2004). The data are collectedexclusively from published sources and are of variablequality as to spatial and temporal information (see, forexample, Copper, 2001; Alvarez and Modzalevskaya,2001; Rong et al., 2004; and references therein). Forthe revised edition of the Treatise, special effort wasfocused on revising type species, selecting/illustratingtypes, describing interiors, studying intrapopulationalvariations, and, more important for the present study,updating the chronostratigraphic framework as clearlyolder/original publications often contain imprecise tem-poral/geographic data. The Geological Time Scale isfrom Gradstein and Ogg (2004; see also Ogg, 2004).

3. Suprageneric categories: up-grading andincrease in number

During the period between the first and the revisededitions of the Treatise, the number of genera (see textheading below) and suprageneric taxa increased. Sub-families increased from 21 considered by Boucot et al.(1965) to 46 in the revised edition. Similarly, the numberof families increased from 15 in 1965 to 33 in 2002, andsuperfamilies from 7 in 1965 to 19 in 2002 (Fig. 1).

In the classification of the spire-bearing brachiopodsof Boucot et al. (1964, 1965), Spiriferida was treatedas an order with the Atrypidina, Retziidina, Athyridid-ina, and Spiriferidina as suborders. Based on structureof the brachidium, these authors divided the Atrypid-ina into two superfamilies, Atrypoidea (including theatrypids and the lissatrypids) and Dayioidea (comprisingthe dayiids, anoplothecids, kaiseriids, and leptocoeliids);

they viewed the Uncitidae as being of uncertain position.

The rhynchonelliform retzioids were elevated byBoucot et al. (1964) to the rank of a suborder, Ret-ziidina, including two new superfamilies, Retzioidea,

Fig. 1. Comparison of number of taxa included in both Treatise edi-tions.

for punctate genera, and Athyrisinoidea for impunctateones. The nucleospiridids, athyridids and meristellididswere included, with family rank, in a single superfamily,Athyridoidea, the latter grouped with the koninckinoidsin a new suborder, Athyrididina.

In the Osnovy Paleontologii (1960), Ivanova, Rzhon-snitskaya, Likharev, Makridin, and Nikiforova viewedthe “spiriferids”, and “atrypoids” as different orders(cf. Ivanova, 1960; Rzhonsnitskaya, 1960, respectively),regarding the “athyridoids” as a superfamily incerti ordi-nis (cf. Likharev et al., 1960). Later, Dagys (1974) ele-vated the athyridides to ordinal rank with two suborders,the Athyrididina for the impunctate genera (with meris-telloids and athyridoids as superfamilies) and the Retzi-idina for the punctate ones (superfamily Retzioidea sensuBoucot et al., 1965). The family Uncitidae was includedby Boucot et al. (1965) in the atrypoids as a super-family of uncertain systematic position, but Likharevet al. (1960) placed it in the superfamily Athyridoidea.It was not considered by Dagys (1974) and was omit-ted from the order Athyridida by Grunt (1989). TheNucleospiridae, commonly placed in the superfamilyAthyridoidea (e.g. Boucot et al., 1965; Dagys, 1974;Modzalevskaya, 1985), was elevated to superfamily sta-tus by Grunt (1986) who considered it as embracing threesuperfamilies Meristelloidea, Athyridoidea and Nucle-ospiroidea within the suborder Athyrididina. Dagys(1974) retained the koninckinids in the Strophomenida,but Harper (in Harper et al., 1993), following Brunton

and Mackinnon (1972), included the koninckinids, withsubordinal rank (along with athyrididines, retziidines,dayiidines, thecideidines, and spiriferidines) in the orderSpiriferida. Subsequently, Dagys (1996) referred the

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F. Alvarez / Palaeo

onickinoids to the order Athyridida, together with ret-iidines and athyrididines, all with subordinal rank.aterhouse (1981) considered that the suborder Retzi-

dina (sensu Boucot et al., 1964) differed substantiallyrom “spiriferids”, “atrypids”, and “athyridids”, and pro-osed placing it in a separate order, Retziida. His diag-osis for the order is identical with that given by Boucott al. (1964) for the suborder, but did not discuss itsontents.

In the 4th volume of Part H (revised) of the Trea-ise on Invertebrate Paleontology, Copper (2002) recog-ized four suborders, Atrypidina, Anazygidina, David-oniidina, and Lissatrypidina within the Atrypida. Someembers of the superfamily Davidsonioidea, includ-

ng Davidsonia, were removed from Strophomenata andlaced, as suborder Davidsoniidina in the order AtrypidaCopper, 1996; cf. Copper, 2002; Williams and Brunton,993). Alvarez and Rong (2002; cf. Alvarez et al., 1998)ncluded three suborders, Athyrididina, Retziidina, andoninckinidina in an apparently broadly paraphyleticrder Athyridida. The koninckinidins have a very dis-inctive morphology, different from typical Athyridid-

na and Retziidina. They are characterized by a plano-r concavoconvex lateral profile, a strophic hinge line,aving tips of the spiralia dorsomedially directed, andaving the umbonal blades curving laterally from the

ig. 2. Annual rate of erection of genera: (a) cumulative number when alliscarding possibly synonyms and those that requires revision ( ); (b) cumonsidered.

5 (2006) 135–149 137

crura. They may be closely related to thecideidines (e.g.Carlson and Leighton, 2001). The punctate rhynchonelli-form retziidines appear as the most derived group of theathyridids (Alvarez et al., 1998; Alvarez and Carlson,1998; Alvarez and Rong, 2002), although they wereconsidered as a suborder, and not a new order (e.g.Waterhouse, 1981) to avoid increase of paraphyleticgroupings (see discussion in Alvarez and Carlson, 1998).The Dayiidae, Anoplothecidae, Kayseriidae, and Unci-tidae, excluded from the Atrypida by Copper (1973a,b,1986; cf. Copper in Alvarez and Copper, 2002) are diffi-cult to accommodate in the Athyridida; they are regardedas uncertain in origin (cf. Alvarez and Carlson, 1998;Alvarez et al., 1998; Alvarez in Alvarez and Copper,2002). The Leptocoeliidae, despite similarities with theAnoplothecidae, were considered rhynchonelloids byCocks (1978) because of absence of evidence for a spi-ralial apparatus. This was accepted by Harper et al.(1993) and Savage (1996, 2002).

4. Rate of proposal of new genera

The rate of description of new genera of spire-bearingbrachiopods (spiriferids excluded) was rather variablefollowing their initial discrimination in the early 1800s(Fig. 2a and b). This was not random but was closely con-

spire-bearing genera (spiriferids excluded) are considered ( ), andulative number of atrypid ( ) and athyridid genera ( ) separately

138 F. Alvarez / Palaeoworld 15 (2006) 135–149

wing th

Fig. 3. Number of authors vs. number of genera erected by author shoCopper, 2002; Alvarez and Rong, 2002; Alvarez and Copper, 2002).

nected with monumental work undertaken by a very few,highly prolific palaeontologists (Fig. 3), and reflected thepolitical and economic state of the world. For example,there was a noticeable increase in the number of taxa dur-ing the second half of the 18th century (Figs. 2 and 4)

coinciding with the remarkable work produced by JamesHall, alone or in collaboration with John M. Clarke, inNew York State (e.g. Hall, 1859a,b, 1860a,b, 1861, 1863,1867, 1894; Hall and Clarke, 1892, 1893, 1894, 1895),

Fig. 4. Annual rate of erection of genera: (a) considering all spire-bearing genthose that requires revision ( ); (b) considering the number of atrypid ( ) a

e very small number of authors that erected more than five genera (cf.

and studies by Alexander Bittner in the Alpine Triassic(e.g. Bittner, 1888, 1890, 1893) (Fig. 5a). After 1895,coinciding with publication of Hall’s last collaborativework with Clarke, the number of genera increased at alow, but rather steady rate until the decade 1956–1965,

with a small increase just prior to each of the two WorldWars, followed by small declines (Figs. 2 and 4). Bycontrast, during the 1950s and early 1960s, just priorto publication of the first edition of Part H of the Trea-

era (spiriferids excluded) ( ), and discarding possibly synonyms andnd athyridid genera ( ) separately.

F. Alvarez / Palaeoworld 15 (2006) 135–149 139

F type speC erids exg

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ig. 5. Annual rate of erection of genera vs. area of which types ofhina ( ), other ( )]: (a) considering all spire-bearing genera (spirifenera separately.

ise, about 30 new genera were proposed (Figs. 2a, 5a).part from an extraordinary peak in the mid-1980s, this

nnual rate was maintained until 2002—publication datef volume 4, with spire-bearers, of Part H of the revisedreatise (Figs. 2 and 4). During that time, significant

umbers of new genera were proposed based on mate-ial from previously poorly known geologic sequences,specially in China and Russia (Fig. 5), and often fromemote areas which were difficult to access (e.g. Dagys,

cimens came from [Europe (©), North America ( ), Russia ( ),cluded) together; (b and c) considering the atrypid and the athyridid

1974; Grunt, 1980, 1991; Chen, 1983; Modzalevskaya,1985, 1997; Wang and Rong, 1986; Rong et al., 1994,1995; Rzhonsnitskaya et al., 1998; Popov et al., 1999;and references quoted therein). Even allowing for a rela-tively high proportion of invalid or synonymous names,

the annual rate of production has been five to seven timesthat observed prior to publication of the first edition(Fig. 2). Revision of Treatise Part H was thus timelyas there has been an almost four-fold increase in the

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number of spire-bearing brachiopod genera (spiriferidsexcluded) to nearly 470 compared with those acceptedby Boucot et al. (1965).

5. Spatio-temporal generic distribution

5.1. Temporal diversity

Generic data available when the first edition of theTreatise was published (Boucot et al., 1965) indicatedthat the diversity of spire-bearing brachiopods, exclu-sive of spiriferids, peaked in the Ludlow, Lower Devo-nian and Upper Triassic (Fig. 6). It has been noted(e.g. Copper, 2001) that atrypids were the only abun-dant spire-bearers in the Caradoc and Ashgill and that,apart from a few reworked Frasnian shells, no Atrypidaare known from the Famennian. Atrypids are thus impor-tant as regards Ordovician diversity, but post-Famenniandiversity of non-spiriferid spire-bearers is concernedexclusively with athryidid lineages that survived the

end-Frasnian global extinction events. It is especiallynoteworthy, considering the number of genera erectedbetween the dates of publication of the two editions ofthe Treatise, that as well as the high diversity observed in

Fig. 6. (a) Total number of spire-bearing genera (spiriferids excluded) for eac( ); genera erected after the first edition was published ( ); all genera inclrepresent percentage of genera. Abbreviations: O2, Middle Ordovician; O3, LaD1, Early Devonian; D2, Middle Devonian; Fr, Late Devonian (Frasnian); FmP1, Cisuralian; P2, Guadalupian; P3, Lopingian; T1, Early Triassic; T2, MiddJ3, Late Jurassic.

5 (2006) 135–149

the Lower Devonian Realm of diversity are seen in theLlandovery, the Mississippian, the Upper Triassic andthe Lopingian (Fig. 6a). When all genera in volume 4 ofPart H (revised) published in 2002 are considered, theresulting pattern is very similar to that resulting fromconsidering the database presented in the first editionof Part H, except that the peak in the Silurian is lesspronounced, having widened to include the Llandovery,Wenlock and Ludlow (Fig. 6).

The above diversity changes could be accentuated bygaps in the record (cf. Savage et al., 1976; Bambachet al., 2004) and heterogeneity of available informa-tion. Data is always “biased” in palaeontology due to“limited availability of raw data on a global scale, orbecause the taxonomy of the group is in a state of fluxthat hampers understanding the distribution patterns” (cf.Gray and Boucot, 1976b). Nonetheless, I believe that themajor changes described correspond to real changes indiversity of spire-bearing brachiopods (cf. Copper, 2001;Alvarez and Modzalevskaya, 2001).

If only atrypid genera are considered, there is a clearpeak in the Lower Devonian. “The late Emsian-Eifelianmarked a peak in atrypoid diversity, reaching a maxi-mum of 47 taxa specifically adapted to and confined to

h age period (considering only the genera included in the first editionuded in the revised edition ( ); (b) as previously, but vertical scalete Ordovician; S1, Llandovery; S2, Wenlock; S3, Ludlow; S4, Prıdoli;, Late Devonian (Famennian); C1, Mississippian; C2, Pennsylvanian;le Triassic; T3, Late Triassic; J1, Early Jurassic; J2, Middle Jurassic;

F. Alvarez / Palaeoworld 15 (2006) 135–149 141

Fig. 7. (a) Total number of atrypid genera for each age period; considering only the genera included in the first edition ( ); genera erected after thefirst edition was published ( ); all genera included in the revised edition ( ); (b) as previously, but vertical scale represent percentage of genera.Abbreviations as in Fig. 6.

Fig. 8. (a) Total number of athyridid genera for each age period; considering only the genera included in the first edition ( ); genera erected afterthe first edition was published ( ); all genera included in the revised edition ( ); (b) as previously, but vertical scale represent percentage of genera.Abbreviations as in Fig. 6.

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142 F. Alvarez / Palaeo

reefal carbonates” (Copper, 2001). If only atrypid gen-era in the first edition of the Treatise are considered,there is another peak in the Wenlock, but this smallerpeak has shifted to the Llandovery when genera erected

Fig. 9. Number (a, c, e) and percentage of genera (b, d, f) vs. distribution; cerected after the first edition was published ( ); all genera included in the revgenera (spiriferids excluded). (c–d) Considering only the Atrypida. (e–f) Consof limited distribution (in two to three listed areas); genera of wide distributionthan nine areas). Areas: 1, North Europe (Britain, Baltic, Norway); 2, CentralSpain); 4, North Africa (Morocco, Mauritania); 5, Iran; 6, Urals; 7, Siberia; 8Greenland; 13, North America; 14, south America; 15, Australia and New Ze

5 (2006) 135–149

between the two editions of the Treatise, or all genera inthe revised edition are considered (Fig. 7). For diversity,radiations and extinctions of atrypid genera see Copper(2001).

onsidering only the genera included in the first edition ( ); generaised edition ( ); (a–b) considering the total number of spire-bearingidering only the Athyridida. Endemics, distributed in one area; genera(in four to nine listed areas); “cosmopolitan” genera (present in moreEurope (Bohemia, Germany, Poland); 3, south Europe (France, Italy,, Kazakhstan; 9, Central Asia; 10, North China; 11 South China; 12,aland.

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quently (Fig. 10b). This is also evident when consideringthe atrypids (Fig. 11a–c) or the athryidids (Fig. 11d–f)alone. Among the atrypids, genera with wide or cos-mopolitan distribution are common in the Ordovician,

F. Alvarez / Palaeo

Similarly, when athyridid genera are consideredlone, diversity peaked in the Lower Devonian withmaller peaks in the Ludlow, Mississippian and Upperriassic, and much lower peaks in the Cisuralian and theopingian. The latter two, appearing as plateaus between

he Mississippian and the Lower Triassic, give a steppedppearance to the curve when only athyridid genera in therst edition of the Treatise are plotted. If only genera pro-osed between the two editions of the Treatise are con-idered, the peak in the Ludlow shifts to the Wenlock, andhe peak in the Cisuralian disappears, but the peaks in theower Devonian, Mississippian, Lopingian and Upperriassic remain, clearly (Fig. 8). Athyridide diversityynamics through the Palaeozoic and early Mesozoicas considered by Alvarez and Modzalevskaya (2001).

.2. Endemism

Because geographic range is involved in definingndemism, apparent levels of endemism may vary fromuthor to author (cf. Alvarez and Modzalevskaya, 2001;arper and Sandy, 2001; Tychsen and Harper, 2004).enera restricted to a specific area, generally to par-

icular plates or terranes, are here termed “endemic”.ost of the spire-bearing brachiopod genera, spiriferids

xcluded, are endemic (as here defined) or have a lim-ted distribution. A few genera have wider distributionrepresented in four to nine areas); about 30 could be con-idered “cosmopolitan” (found in more than nine areas)Fig. 9).

It is an obvious truism that not all regions have beenqually studied. “In poorly sampled biogeographic units,he apparent percentage of cosmopolitan taxa is alwaysreater than in units where collecting has been moredequate, since endemic taxa are invariably more rarelyepresented, the small sample thus introduces a signif-cant sampling artifact” . . . “the chances are thus greathat where only a few specimens are available, the veri-able traveller’s cigarbox collection, that they will be ofosmopolitan taxa” (Gray and Boucot, 1976b). So, withhe heterogeneous data currently available, the resultsrovide only a tentative picture of the spire-bearingiversity.

.2.1. General endemismFig. 9a shows a great increase in endemic genera

roposed between the dates of publication of the two edi-ions of the Treatise compared with those accepted in the

rst edition. In both editions, the number of genera withide or cosmopolitan distribution is small (Fig. 9a and). When only atrypid genera are considered, the curveshow the number of endemic genera is greater among

5 (2006) 135–149 143

those proposed between 1965 and 2002 than amongthose proposed prior to 1965. Conversely, the number ofgenera with limited distribution increases greatly if onlydata from the first edition are considered (Fig. 9d). Otherportions of the curves are rather similar (Fig. 9c and d).With only athyridid genera the curves show endemismto be greater for genera proposed between the two edi-tions of the Treatise. The number of genera having wideor cosmopolitan distribution is always very small (halfthan in the Atrypida) (Fig. 9e and f).

5.2.2. Endemism through timeConsidering the degree of endemism through time,

the number of spire-bearing brachiopod genera (spir-iferids excluded) with wide or cosmopolitan distributionis greater among those included in the first edition ofthe Treatise (Fig. 10a) than among those erected subse-

Fig. 10. Percentage (vertical scale) of endemic and cosmopolitan spire-bearing genera (spiriferids excluded) during the Phanerozoic. Con-sidering (a) only the genera included in the first edition; (b) generaerected after the first edition was published; (c) all genera included inthe revised edition. Abbreviations as in Fig. 6.

144 F. Alvarez / Palaeoworld 15 (2006) 135–149

pid (a–ected af

Fig. 11. Percentage (vertical scale) of endemic and cosmopolitan atryand d) only the genera included in the first edition; (b and e) genera errevised edition. Abbreviations as in Fig. 6.

especially Middle Ordovician (Fig. 11a–c), and thePrıdoli (Fig. 11a and b). On the other hand, Llandovery,Ludlow, and Lower Devonian genera display greaterendemicity (Fig. 11a–c). The degree of endemism in theathyridids is greater than in the atrypids (cf. Fig. 11a–cwith d–f). The Prıdoli, Famennian, Pennsylvanian andGuadalupian show an increase in athyridid genera withwide to even cosmopolitan distribution (Fig. 11d–f).Degrees of endemism versus cosmopolitanism, and gen-eral diversity, extinction, origination and turnover pat-terns among athyridides were studied by Alvarez andModzalevskaya (2001); this was focused on general ten-dencies in evolution at generic and subfamilial levels.

6. Conclusions and perspectives

There has been an almost four-fold increase in thenumber of spire-bearing brachiopod genera (spiriferidsexcluded) included in the revised edition of the Trea-tise (to nearly 470; Figs. 1 and 2; Copper, 2002; Alvarezand Rong, 2002; Alvarez and Copper, 2002) comparedwith those recognized by Boucot et al. (1965). Manyof these genera were based on specimens from pre-viously poorly known geologic successions, especially

in Russia and China (Fig. 5) where Chinese and Rus-sian colleagues had undertaken admirable and difficultresearch, frequently in very remote areas that are dif-ficult to access. These ambitious new collections and

c) and athyridid (d–f) genera during the Phanerozoic. Considering (ater the first edition was published; (c and f) all genera included in the

descriptions “diluted at least somewhat the fairly strongNorth American and European bias in the 1965 Treatise”(Carlson and Leighton, 2001, p. 27). Even assuming arelatively high proportion of invalid or synonymous gen-era, the annual rate of creation of genera has been five toseven times that prior to publication of the first edition(Fig. 2).

Spire-bearing brachiopod genera (spiriferidsexcluded) erected after the first edition of the Treatiseshow a greater degree of endemicity than those erectedprior to its appearance (cf. Boucot et al., 1965), withendemism displayed by the athyridids being greater thanfor the atrypids (Fig. 11a–f). No correlation seems toexist between degree of endemicity and generic diversityat any specific time (Fig. 12). Endemic spire-bearersare of great value in discriminating biogeographic units(cf. Boucot et al., 1969; Boucot and Johnson, 1973;Johnson and Boucot, 1973; Savage et al., 1976; Ronget al., 1995; Talent et al., 2001). Spire-bearing generawith cosmopolitan tendencies did not evolve rapidly(e.g. Nucleospira, Composita), so their value for globalstratigraphic correlations is generally poor.

The large increase in generic-level taxa appears tohave resulted in a cascading expansion in numbers of

higher taxa. During the period between the first andrevised editions of the Treatise, subfamilies increasedfrom 21 discriminated by Boucot et al. (1965) to 46 inthe revised edition. Similarly, the number of families

F. Alvarez / Palaeoworld 1

Fig. 12. Percentage (left vertical scale) of spire-bearing genera (spir-iferids excluded) for each age period [considering only the generaincluded in the first edition ( ); genera erected after the first editionwas published ( ); all genera included in the revised edition ( )]vt

ib

bailit

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s. rate (right vertical scale) of endemic distribution (wide distribu-ion/restricted distribution). Abbreviations as in Fig. 6.

ncreased from 15 to 33, and superfamilies from 7 to 19etween 1965 and 2002.

In addition to the remarkable increase in the num-er of taxa during the last four decades, there has beenn obvious shift in brachiopod research with increasingntegration of other disciplines (biological, palaeonto-ogic, mineralogic), which is improving our understand-ng of spire-bearers, and of the phylum as a whole inime and space (cf. Williams, 2001).

Advances in plate tectonics and in knowledge ofnvironmental changes in marine environments willreatly help paleobiogeographic interpretations (e.g.cotese’s PALEOMAP Project: http://www.cscotese.om/earth.htm). Therefore, and as Gray and Boucotlready predicted in 1976a: “. . . the next generationf taxonomic palaeontologists will publish many moreiogeographic syntheses that, in turn, will begin to influ-nce the attitudes of practising taxonomists concernedith the classification and evolution of their specialroups”.

Advances in computer science, information technol-gy, and the World-Wide Web make new applicationsossible. For example, on completion of the reviseddition of the Treatise, several initiatives have beenaunched to provide databases that should result in excel-ent overviews of the current state of our knowledge ofhe taxonomy of brachiopod faunas (Curry et al., 2001;urry, personal communication, 2003) and their palaeo-iogeography (e.g. Talent et al., 2001; Harper and Sandy,001; and reviews in Jablonski et al., 1985; Ebach and

dgecombe, 2001; Kaesler et al., 2001). This togetherith new phylogenetic software will surely help evo-

utionary studies (Alvarez and Carlson, 1998; Alvarez,003, and discussion therein). New, more precise bios-

5 (2006) 135–149 145

tratigraphic data will provide a better standard againstwhich phylogenetic trees may be tested, essentiallyindependent of traditional stratigraphy (e.g. Adrain andWestrop, 2001, and references therein). Studies on bra-chiopod genetics and molecular systematics may providean independent test of shell-morphology-based taxon-omy of spire-bearers and other groups of brachiopods(cf. Cohen, 2001); this area is expanding.

New techniques for understanding biomineraliza-tion depend to a great extent on progress madeby brachiopod workers (e.g. Cusack, 2001). Isotopicstudies, particularly using low-magnesium brachio-pod shells as environmental indicators, are impactingimportantly on specification of palaeotemperatures (e.g.Buening, 2001; and discussion in Curry and Fallick,2002).

The recently updated glossary of morphological andanatomical terms applied to brachiopods (e.g. Bruntonet al., 1996; Williams and Brunton, 1997) has helpedgreatly in reducing unnecessary proliferation of termi-nology – often a spinoff from “geographic, linguistic,geopolitical, or stratigraphic boundaries or specialisa-tions” (cf. Brunton et al., 1996) – and in assemblingdatabases. It is hoped that new ways for reconstructingshell interiors will help in study of the cardinalia and bra-chiojugal system of spire-bearers. At present, the config-uration of such an important structure as the brachiojugalsystem is known, albeit superficially, in less than 50% ofthe genera commonly considered to be athyridides (seeAlvarez et al., 1998; Alvarez and Carlson, 1998; Alvarez,1999; Alvarez and Rong, 2002).

Changes in editorial policies, new publication pos-sibilities (e.g. electronic journals) and referee’s eval-uations may stimulate the submission of manuscriptsfocused on revision of poorly known taxa, and theimplications of systematics to biostratigraphy, paleoe-cology, paleogeography and evolution (e.g. Posenato,1989, 1998, 2001; Alvarez et al., 1996; Alvarez andBrime, 2000; Alvarez and Brunton, 2000, 2005; Talent etal., 2001; Chen et al., 2003; Botquelen and Gourvennec,2003; Rong et al., 2004; Shen et al., 2004; Sun et al.,2004; Copper, 2004b). It is hoped that these changes willdecrease the frequent tendency of elevating taxa fromone rank to another “with relatively little substantivechange in the amount of morphological diversity theycircumscribe” (cf. Carlson and Leighton, 2001). Thefuture lies, as Harper and Sandy wrote (2001, p. 207; seealso “current challenges” and “anticipated future direc-

tions” in Carlson, 2001) “in the continued taxonomicstudy of new and existing faunas together with the refine-ment of our age and environmental constraints on thesefaunas”.

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Acknowledgements

This paper is dedicated to Art Boucot, passionateaddict to palaeontology and stratigraphy, a “friendlygiant” with good friends all around the globe. I amindebted to Rong Jia-yu for providing me with the oppor-tunity to collaborate in this special-issue of Palaeoworldin his honour. Rong Jia-yu and John A. Talent providedconstructive comments and kindly improved the styleof this paper. Helpful discussions and support for vari-ous aspects of this study from Covadonga Brime is alsogratefully acknowledged. Finally, I thank the two refer-ees A.J. Boucot and S.L. Long for their valuable reviewsand Chen Siwei for her editorial work.

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