J. Paleont., 73(4), 1999, pp. 549-570 Copyright ? 1999, The Paleontological Society 0022-3360/99/0073-0549$03.00

A REVISEDCHITINOZOAN CLASSIFICATION

FLORENTIN PARIS, YNGVE GRAHN, VIIU NESTOR, AND ISKRA LAKOVA Laboratoirede Paleontologie,UPR 4661 du CNRS, Universit6de RennesI, 35042 Rennes-cedex,France , Universidadede Estatodo Rio de Janeiro-UERJ,Faculdade de Geologia,20559-900 Rio de Janeiro,Brazil , TallinnTechnical University, Institute of Geology, 7 EstoniaAvenue, 10143 Tallinn,Estonia , and GeologicalInstitute, Bulgarian Academy of Sciences, 1113 Sofia, Bulgaria

ABSTRACT-The successful definitionof chitinozoangenera dependsprimarily on the precisionof the criteriaused. A standardized morphologicalterminology based upon detailsfrom scanningelectron microscope observations of the most representativetaxa bearing these charactersis thereforeproposed. The 143 genera,or subgenera,described so far in the literatureare reviewedin orderto exclude invalidtaxa and obviousjunior synonyms. Particular attention is paid to preventingthe overlapof genericdefinitions of the 56 genera ultimatelyretained. A brief accountof the diagnosticfeatures and stratigraphicrange of selectedgenera is given, andbasic information concerningthe type materialof these genera is listed. Finally, a supragenericclassification of the whole Chitinozoagroup based on diagnosticfeatures whose hierarchyis establishedon statisticaland evolutionarygrounds, is given. One new subfamily,Pogonochitin- inae, three new genera,Baltochitina, Hyalochitina, and Saharochitina,and a new species Baltochitinanolvaki, are defined.The sub- species Fungochitinafungiformis spinifera is elevatedto a specificrank.

INTRODUCTION of all 143 genera described and, after excluding those genera we a HITINOZOANSCONSTITUTE a groupof organic-walledmicro- regarded as invalid by ICZN (third version), propose syn- C list based on the criteria we consider to be essential. fossils, well represented in Early Ordovician (late Trema- onymy the are not suffi- doc) to latest Devonian (latest Famennian) marine sediments Because many of existing generic diagnoses we the definition of selected from nearly all Paleozoic oceans. They are found isolated or in ciently definite, supplement original with their features. we chainlike structures in most sedimentary or low grade metamor- genera respective diagnostic By doing so, to of the definitions, which phic rocks, where their abundance usually ranges from a small hope prevent overlapping generic has to be one of the and most constant number of specimens to several hundred (exceptionally up to proven greatest problems in chitinozoan several thousand) specimens per gram of rock. The length of a taxonomy. chitinozoan vesicle (basic isolated element) ranges from ca. 50 to 2,000 Jim (with an average size of around 150-250 Jim). CHITINOZOAN MORPHOLOGY They constitute an enigmatic group of fossils whose biologic General terminology.-A chitinozoan individual may be de- affinities and significance are still interpreted in a variety of fined as a small organic-walled vesicle with an opening. Well- ways (see review in Miller, 1996), more than 60 years after their preserved chitinozoans (i.e., in full relief and not eroded or dis- first description by Eisenack (1931). Personally, we favor the torted) share three major characteristics (Fig. 1): 1) they are hypothesis of eggs of soft-bodied metazoans (Paris, 1981, p. 84; made up of an organic membrane delimiting a cavity; 2) they Paris and Nolvak, in press) referred as chitinozoophorans by have an aperture sealed with a plug; and 3) they normally dis- Grahn (1981). They have proven to be very efficient stratigraph- play a radial symmetry. The actual chemical composition of the ic tools, and they are extremely useful for paleogeographic re- chitinozoan vesicle remains unknown. Therefore, this definition constructions. However, a unanimously accepted taxonomy is is so broad that various organisms, or part of organisms may be now strongly needed in order to make application of chitinozoan included. Size limits (<2,000 Ijm and >50 jim) or stratigraphic genera for paleogeographic and paleoenvironmental investiga- range restriction (exclusively of Paleozoic age, unless reworked) tions more reliable. restrict this definition and allow various present day cysts, co- There have been many attempts to classify chitinozoans. They coons, or eggs of invertebrates (insects, annelids, molluscs, etc.) were carried out through individual initiatives (Eisenack, 1931, to be excluded from the chitinozoans. When present, the external 1968, 1972; Van Oyen and Calandra, 1963; Jansonius, 1964, ornamentation of a chitinozoan vesicle is fairly distinctive com- 1967, 1970; Tappan, 1966; Taugourdeau,1966, 1981; Paris, pared to the ornament of almost all other organic-walled micro- 1981; Achab et al., 1993) or under the authorityof the chiti- fossils of similar age (i.e., acritarchs, spores, and foraminiferal nozoan Subcommission of the "Commission Internationale de linings). Microfloredu Paleozoique"(CIMP) (Taugourdeau et al., 1967). A detailed morphological terminology was first introduced by In the last 25 years, major improvementsin the knowledge of Combaz and Poumot (1962) and then subsequently completed chitinozoanmorphology have been achieved, primarilydue to by Combaz et al. (1967) under the direction of the Chitinozoa the routineuse of the ScanningElectron Microscope (SEM). In Subcommission of CIMP. Most recent papers dealing with chi- addition,numerous new occurrenceshave greatly increasedthe tinozoan systematics have used either the English adaptation chitinozoanrecord, so that it now includes more than 1,000 taxa proposed by Laufeld (1974) or the French terminology adopted describedfrom almost all continents.We are convincedthat suf- by Paris (1981). These terminologies are still valid and well ficient data exist to propose a standard terminology and a general adapted for the description of the chitinozoans. Only a few mod- classificationof chitinozoansthat integratesthe taxa described ifications or additions are necessary in order to integrate the since the last extensive revision (Taugourdeauet al., 1967; Com- most recent morphological details revealed by SEM observa- baz et al., 1967). Our aim is to integrateall originalmorpholo- tions. One of our aims is to promote a standard morphological gies recorded during the last 30 years into a general classification nomenclature that would be unanimously accepted and used. of the group. Particularattention is paid to the definitionof the The main parts of a chitinozoan vesicle are illustrated, show- morphologiccharacters, and to the establishmentof a hierarchy ing the three principal morphological types of chitinozoans (Fig. among these morphologic elements. A critical review is made 1). The basic element is the vesicle, with a bulging part called 549

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Apertural pole Apertural pole Apertural pole

0 (1 z

shoulder

I-

- E flanks

0

margin

process I apex (=appendix) apex

Antiapertural pole Antiapertural pole FIGURE 1-Main morphological features of the three principal types of chitinozoans (modified from Paris, 1981, fig. 56).

the chamber. The aperture is either situated directly on the cham- that were not accurate enough or were already used with a dif- ber or at the distal end of a tubelike neck. In order to clarify ferent meaning. It also introduces the morphologic terms we use discussion and descriptions, an arbitrary orientation of the ves- for the discrimination of the chitinozoan genera we have select- icle has been adopted. The apertural pole is regarded as repre- ed. In addition, references are made to recently published SEM senting the top of the vesicle. This means that, in a chainlike images which illustrate almost all of these terms. structure, the upper (or last) vesicle has a free aperture. However, 1) antiapertural pole: lower part of the vesicle, opposite the no biological nor physiological significance has to be given to aperture. this orientation, which is exclusively conventional. A collarette, 2) aperture: large opening at the top of the vesicle, delimited frequently corresponding to a thinning of the wall, may be pres- either by the lips of the neck or collarette, or if these two other ent around the aperture. The plug sealing this aperture is either elements are absent, by the border of the chamber (Fig. 1). This term is instead of called an operculum or a prosome depending on its internal or preferred "mouth", which has physiological external position and on its role in inter-vesicle linking (see def- significance. 3) apertural plug: general term designating the simple or initions below). The base, which corresponds to the antiapertural complex plug sealing the aperture, represented either by the end of the vesicle, may display various linkage structures around operculum or by the prosome (see definitions below). the apex. This base is from the flanks the separated by margin. 4) apertural pole: arbitrarily designated as the upper part of This zone proves to be of prime importance for chitinozoan tax- the vesicle. as it onomy, frequently bears peculiar ornamentation (e.g., pro- 5) apex: point of emergence of the axis of symmetry on the cesses or The flanks carina). themselves may be separated from base of the chamber (Fig. 1). the neck by the flexure (e.g., vesicles with well-differentiated 6) apical pit: circular, depressed area surrounded by the rim necks). of the mucron. [N.B. the apertural pit does not open in the ves- Lexicon of the morphologic features and structures.-This icle in well-preserved specimens (see Miller, 1996, pl. 1, fig. 1; lexicon is given in order to restrict some previous definitions Verniers et al., 1995, fig. 6K)].

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1 2 3 4

4 5 6 7 8

9 10 11 12 FIGURE2-Different kinds of vesicle wall surfacesof chitinozoans.1, Smooth,scabrate, vermiculate; 2, foveolate;3, feltlike;4, spongy;5, verrucate (granules,tubercules and/or cones less than2 micronshigh); 6, simple spines; 7, simple and branchedhairs or spines;8, bi and multirootedspines; 9, meshlikestructure; 10, crests with verticalrows of free or connectedspines; 11, crests of weblike to discontinuousmembranes; 12, complete or perforated/reticulatedcarina. Stippled = innerlayer; black = outerlayer.

7) apical structure: includes the scar, callus, mucron, copula 12) carina: circular expansion of the wall (outer layer) round and peduncle; the bulb is also regarded as an apical structure. the chamber; it may be located below, on, or above the margin 8) axis: imaginary line joining the center of the aperture to (Fig. 2.12). the apex and representing the axis of symmetry of the vesicle. 13) catenary structure (=chainlike structure): vesicles con- 9) base: part of the chamber opposite to the aperture (=an- nected "aperture to base", along their axis (straight, curved or tiapertural end or chamber bottom). coiled chain) (see Jenkins, 1970b, pl. 6, figs. 4-6; Paris and 10) bulb (=siphon): membranous ampoule extending from Nolvak, in press, pl.1, figs. 1, 3-4), or vesicles connected "neck the base of a vesicle (e.g., in Siphonochitina formosa) (see Jen- to margin", with a coiled pattern (e.g., Lagenochitina navicula kins, 1967, pl. 75, figs. 2-5). in Paris, 1981, pl. 31, figs.1-3). 11) callus: short stublike thickening on the apex (e.g., in Des- 14) central cavity: inner cavity corresponding to the chamber mochitina densa illustrated by Laufeld, 1974, fig. 39D). (Fig. 1) (contained the embryo if one adopts the egg hypothesis). 15) chamber: part of the vesicle (frequently bulging) situated below the neck or below the collarette when present (see Fig. 3 for the different chamber shapes). 16) cluster: group of vesicles belonging to the same species and connected by their flanks (aperture free); they are usually Lenticular arranged in a coiled pattern (see Koslowski, 1963, fig. 3; Miller, 3, 1996, pl. 5, figs. 1-2; Paris and Nolvak, in press, pl. 1, figs. 5- 8). 17) coiled chain (=helicoidal chain): vesicles connected ovoid . - "neck to margin" i.e., with a free aperture, or "aperture to base" i.e., aperture linked to the succeeding vesicle. I cyidia 18) collarette (=collar): thinned cylindrical or flaring part of Spherical / the neck, or of the vesicle wall when the neck is absent. The collarette surrounds the aperture. 19) copula: tubular, membranous expansion surrounding the apex (see Paris, 1981, pl. 1, figs. 7, 13). hemispherical conical claviform 20) crests: vertical rows of free or connected spiny ornamen- tation 2.10), or of weblike to continuous membranes hemispherical conical (Fig. (Fig. 2.11) (see Achab et al., 1993, pl. 1, figs. 1-2; Miller, 1996, pl. FIGURE3-The basic chambershape of the chitinozoans. 2, figs. 2-3).

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21) diameter:refers to the maximumdiameter of the chamber or very long (see Wood and Miller, 1991, pl. 1, figs. 1-5). (N.B. (D), neck (dn), or aperture(da); a coefficientof correction(e.g., the processes never communicatewith the interiorof the cham- 0.7 or 0.8, dependingon the degree of flattening)(see Paris in ber). Babin et al., 1979) must be used to restorethe full-relief mea- 42) prosome:internal plug situatedat the base of the neck; surementwhere the vesicle is flattenedor collapsed. may be simple (disklike)or complex (tubularstructure with sev- 22) equatorialplane: correspondsto the maximumdiameter eral or numeroushorizontal septa, i.e., "accordionpleated pro- of lenticularor sphericalchambers. some" of previous authors;see Miller, 1996, pl. 4, fig. 8); it is 23) external structures:include the carina and apical struc- not involved in vesicle linkage. tures as well as the sleeve. 43) reticulum:reticulated outer layer stronglyattached to the 24) flanks:part of the chambersituated between the margin flanks;may extendbeyond the base of the vesicle as a perforated and the neck or collarette(Fig. 1). carina(e.g., Parisochitinaperforata in Paris, 1996, pl. 2, fig. 2); 25) flexure:concave zone separatingthe flank from the neck reticulumbearing vesicles are regardedas glabrous. (Fig. 1). 44) rib:annular (horizontal) thickening of the wall (e.g., Mar- 26) glabrous:state of the vesicle surfacewhen lacking spiny gachitina catenaria in Paris and Grahn,1996, pl. 3, fig. 2). ornamentation(Fig. 2.1-5). 45) rica: membranous,flanging part of the prosome or oper- 27) inner layer:internal membrane of the wall usually acting culum, lining the upper part of the chamber(see Miller, 1996, as a frameworkfor the vesicle (Figs. 1, 2). pl. 2, fig. 4). 28) laciniated:deeply indentedborders of a carina(e.g., Po- 46) ridge: verticallinear thickeningof the wall (e.g., Laufel- gonochitina spinifera in Paris, 1996, pl. 1, fig. 9). dochitinastentor in Grahnet al., 1996, pl. 2, fig. 6). 29) linearchain: straight, curved, or coiled catenarystructure 47) scar:circular mark (depressed or slightly protruding)cor- where the aperturalpole of one vesicle is fixed to the base of respondingeither to a thinning or to a thickeningat the apex the following one; the apertureis not free (see above). (see Grahnet al., 1996, pl. 1, figs.l, 10); may also exist on the 30) linkage structures:elements intervening in the connection center of the operculum(see Paris and Nolvak, in press, pl. 1, of two successive vesicles in a catenarystructure (e.g., opercu- fig. 2). lum, apical structures,lips, base). 48) septa: horizontalmembranous partition (from a few to 31) lip: distal partof the collarette(or of the neck), surround- more than 20) within a prosome (see Laufeld, 1974, fig. 72B; ing the aperture. Miller, 1996, pl. 2, fig. 4 and pl. 4, fig. 8). 32) margin (=basal edge sensu Laufeld, 1974): transition 49) sheathingprocess: process whose proximalend extends zone between the base and the flanks; may be inconspicuous, horizontally from the margin (e.g., Salopochitina monterrosae; rounded,blunt or sharp.The margin bears the major morpho- see Tekbaliand Wood, 1991, pl. 19, figs. 4-5). logic elements (e.g., carina,processes). 50) shoulder:convex area at the top of the flanks,just below 33) mucron:thickened, raised rim surroundingthe apical pit the flexure;when both shoulderand flexureare present,the up- (e.g., see Laufeld, 1974, fig. 29D). per part of the chamberhas a sigmoid profile (Fig.1l). 34) meshlike: spiny ornamentationmaking up a net-workor 51) sleeve: partiallyor totallyattached membrane (usually the a trellis-workon the surface of the wall (Fig. 2.9) (e.g., Acan- outer layer) covering the vesicle and extendingbeyond the base thochitinabarbata in Nolvak and Grahn,1993, pl. 4, fig. A, or (see Achab et al., 1993, pl. 2, figs. 2-6). Muscochitinamuscosa in Paris, 1981, pl. 30, figs.17-19). 52) spines: all kinds of simple to complex elongate expan- 35) neck (=aperturaltube): tubular structure expanding aper- sions of the outer layer (see below); spine length must be at turalwardfrom the chamber;frequently terminated by a collar- least twice theirwidth and exceed two microns;most frequently ette. hollow (Fig. 2.6-10). (N.B. the hollow spines never communi- 36) operculum:disklike plug sealing the apertureof the ves- cate with the interiorof the vesicle). icles lacking a neck (see Paris and N6olvak,in press, pl. 1, fig. 53) spiny ornamentation:vesicle bearing spines, crests, and/ 2); a membranousexpansion flaring antiaperturalward("rica" or processesrandomly distributed or arrangedin rows or crowns sensu Bockelie, 1981) is common (e.g., see Paris, 1996, pl. 1, (see below for additionalinformation). fig. 4). 54) vesicle: basic chitinozoanindividual, including the wall 37) ornamentation:includes all the spiny, hairlike or linear of the chamberand the neck (and collarettewhen present) as expansions(e.g., crests) of the outer layer (Fig. 2.6-11). well as the aperturalplug (Fig. 1). 38) outer layer: externalmembrane forming the vesicle wall 55) wall (=test): organicenvelope of a chitinozoan;includes togetherwith the inner layer;the vesicle ornaments(carina, pro- both the outer and inner layer. cesses, spiny ornamentation)arise from evaginationsof this out- Shape of the chamber.-The following characteristics (Fig. 3) er layer (Figs. 1, 2), which may also be glabrous(Fig. 2.1-5). concern only vesicles in full relief. In flattenedspecimens the 39) peduncle: solid short (e.g., Margachitina catenaria in collapse of the vesicle is usually perpendicularto the equatorial Paris and Grahn,1996, fig. 2) or elongatedcylindrical structure plane for lenticularand sphericalchamber without a neck. It will extendingfrom the apex (e.g., Urochitinasimplex in Paris,1996, be parallel to the axis for all the other cases. The collaretteis pl. 3, fig. 12). not taken into accountwhen definingthe shape of the chamber. 40) perforated:refers to closely distributedholes occurring 1) lenticular:width significantlylarger than length (D > L); on the carina(e.g., Sagenachitinaoblonga in Paris, 1981, pl. 9, margin rounded; base convex (e.g., Calpichitina, Fungochitina). fig. 8) or on the collarette(e.g., Fungochitinafenestrata in Paris, 2) spherical:width of the chamberequal or close to its length 1996, pl. 1, fig. 1); the ultimate stage is the reticulatepattern (D = -L); margininconspicuous; base rounded(or exception- when holes are only separatedby threadlikeremains (e.g., Par- ally evaginatedsuch as in Margachitinaelegans); shoulderpres- isochitina perforata in Paris, 1996, pl. 3, fig. 2). ent (e.g., Hoegisphaera, Sphaerochitina). 41) processes (=appendices):simple or complex spines (e.g., 3) hemispherical:length close to half the diameter(L= D/2); Ancyrochitinadesmea in Paris, 1996, pl. 2, fig. 6) arrangedin a base flat;margin sharp or blunt;shoulder conspicuous (e.g., Bul- crown on the margin,or near the marginin the case of an ovoid bochitina, Cyathochitina). to claviformchamber; the processes are usually hollow but may 4) ovoid: length of the chambergreater than its diameterbut have a cell-like structure(e.g., Plectochitina);they may be short less than three times this diameter(3D > L >D); margin and

This content downloaded on Thu, 7 Mar 2013 06:52:12 AM All use subject to JSTOR Terms and Conditions PARISET AL.-REVISED CHITINOZOANCLASSIFICATION 553 flexureinconspicuous; base convex to rounded(e.g., Desmochi- as a linkage medium with the operculumof the precedingves- tina, Angochitina), more rarely truncated (e.g., Lagenochitina icle. The bulb is also consideredas an apical element, but it is dalbyensis) or ogival (e.g., Lagenochitina conifundus). not clearly involved in intervesiclelinkage. 5) claviform: the chamber length exceeds three times the 3) Sleeve: this correspondsto a partiallyor totally unstuck maximumdiameter (L > 3D), base rounded;margin inconspic- outer layer that may extend beyond the margin and even far uous; shoulder absent (e.g., Clavachitina, Laufeldochitina). beyond the base of the vesicle (e.g., Pellichitina, Cutichitina, 6) conical: straight,tapering flanks; base flat; margin sharp Velatachitinain Achab et al., 1993, pl. 2, figs; 2, 4, 6). This or blunt (e.g., Bursachitina, Euconochitina, Cyathochitina). term is not extended to various types of vertical folding (e.g., 7) cylindrical: length usually several times the diameter; Calpichitinavelata in Paris and Grahn, 1996, pl. 1, figs. 7, 9) flanksstraight and parallel;base flat;margin sharp or blunt (e.g., or to irregulardetachment of the outerlayer (e.g., Desmochitina Rhabdochitina). juglandiformisin Paris, 1981, pl. 17, figs. 16-17). Significant Vesicle characteristics.-The vesicle surface may be glabrous flatteningof the vesicle may be responsiblefor a secondaryde- or spiny; it may also bear a carina or an apical structure.All tachmentof this outer layer mirroringa sleeve (especiallyin the combinationsof these patternsmay exist, with the exceptionof Desmochitinidae). both glabrousand spiny surfacestogether. 1) Surface patternof the wall (Fig. 2; see also Grahnet al., PRINCIPLE AND METHODOLOGY 1996, pl. 1, figs. 1-11, and pl. 2, figs.1-3, 5-6, 9-10): this pat- Since the firstdescription of chitinozoansby Eisenack(1931), tern includes both totally smooth surfaces (Fig. 2.1) as well as two systems have prevailedin the classificationof these organic scabrate, vermiculate (Fig. 2.1), foveolate (Fig. 2.2), feltlike microfossils.One involves the supragenericsubdivisions first in- (Fig.2.3), spongy (Fig. 2.4), or microgranuloussurfaces, includ- troducedby Eisenack(1931) and subsequentlyimproved by this ing tuberclesand cones (Fig. 2.5). When these granulesare less author (1968, 1972). This classification was adopted, and in than two micronshigh, the wall surfaceis regardedas glabrous. some cases furtherdeveloped by other authors(Van Oyen and A glabroussurface may or may not be associatedwith a carina Calandra,1963; Jansonius, 1964, 1967, 1970; Tappan, 1966; or with apical structures.Corrugations (e.g., Armoricochitinare- Taugourdeau, 1966, 1981; Paris, 1981; Schallreuter, 1981; ticulifera in Nolvak and Grahn, 1993, pl. 3, fig. c) as well as Achab et al., 1993; Nestor, 1994; Paris and Grahn,1996; Miller, ridges (e.g., Laufeldochitinastentor in Nolvak and Grahn,1993, 1996). The chitinozoangroup was there by progressivelysub- pl. 1, fig. G) or ribs (e.g., Margachitinacatenaria in Pariset al., divided into Orders,Families, and even into Subfamilies (see 1996, pl. 3, figs. 1-2) may occur on glabroussurfaces. Paris, 1981; Achab et al., 1993; Miller, 1996). 2) Spiny ornamentation(Fig. 2.6-11): this ornamentationin- The second system involves an alphabeticallisting of genera. cludes all kinds of spiny extensions longer than two microns, It was principallyadvocated by Laufeld (1967, 1974) and Jen- occurring on the vesicle, i.e., simple (Fig. 2.6), bifurcatedor kins (1970b). Many supportersof this simpler method argued branchingspines (Fig. 2.7), bi- or multirootedspines (Fig.2.8), that the affinities and the biologic significance of the chitino- anastomosedspines (Fig. 2.10) or those connectedby a meshlike zoans were too poorly known to justify any classification.In structure(Fig. 2.9), spines in rows (Fig. 2.10), crowns or crests reality, when groupingseveral species within a genus on mor- (Fig. 2.11), or randomlydistributed on partor all of the vesicle. phological grounds,a paleontologisthas alreadystarted to clas- The spiny ornament,which is usually hollow, may coexist with sify these taxa. Despite the fact that alphabeticsorting has been any of the otherelements (i.e., carina,apical structures)with the widely used by numerousauthors, it proves to be of no help for exception of a glabrous surface. It is of prime importanceto taxonomicpurposes. The only positive aspectof alphabeticclas- distinguisheroded spines (roundedscars) from a truly glabrous sification is to facilitate a rapid search and location of genera surface. within a long list of taxa. When located on the margin,the spines arrangedinto a crown Utility of a classification for chitinozoans.-Chitinozoan are called processes.They display the same rangeof complexity workershave unanimouslyadopted the Linneanbinominal tax- as the spines occurringon the flanks;in additionthey may have onomy, groupingspecies into genera,and thereforehave already a spongy or a cell-like texture(e.g., Plectochitina). initiateda classificationat a higherrank than the basic biological External structures.-These include the carina, the apical entity, i.e., the species. Consequentlywe see no seriousreasons structures,and the sleeve. to dismiss a supragenericclassification of the chitinozoans.In 1) Carina:this correspondsto an extension (annularevagi- addition, we believe that a formal classification,including su- nation) of the outer layer,perpendicular or not to the axis of the pragenericsubdivisions, may enhancethe knowledge and study vesicle; the carinamay be located eitherbelow, on, or above the of the chitinozoansfor the following reasons: margin;it may be fairly thick (e.g., Laufeldochitina),membra- 1) It encourageschitinozoan workers to adopt a logical tax- nous (e.g., Cyathochitina kuckersiana, Pterochitina perivelata), onomic approach,based on carefulmorphological analysis, rath- extending horizontally (e.g., Cyathochitina vaurealensis), or er than on a superficial survey of the general outline of the flaringantiaperturalward (e.g., Laufeldochitina).The carinamay taxon. also be complete (Fig. 2.12) (e.g., Cyathochitina),perforated, 2) It promotesa clearly defined hierarchyof morphological reticulated (Fig. 2.12) (e.g., Sagenachitina, Baltochitina) or la- charactersused in the chitinozoanidentification, consequently ciniated (e.g., Pogonochitinaspinifera in Paris, 1996, pl. 1, fig. preventingoverlap of generic diagnoses. 9). The carinamay be associatedeither with a glabrousor spiny 3) It provides a level of identificationcompatible with the wall and/orapical structures. state of preservationof the material,while still remaininginfor- A carina does not coexist with processes (homologous ele- mative for broad stratigraphicalpurposes. ments); however, the threadlikeexpansions extending from in- 4) It gives a frameworkfor cladistic analysis. sertionzones partiallyenveloping the marginof some taxa (e.g., 5) It gives a betterground for phylogeneticanalysis, even if "Conochitinafilifera" in Tekbaliand Wood, 1991, pl. 19, figs.1, such approachesare still very tentativein an enigmatic group 4-6) are regardedhere as the ultimate case of processes (i.e., such as the Chitinozoa. sheathingprocesses). 6) It preventsredundancy in generic diagnosis because it re- 2) Apical structures:they include the scar, callus, mucron, quires direct referenceto family or subfamilycharacteristics. copula and peduncle (see definitionsabove); they usually serve 7) A supragenericclassification also provides an excellent

This content downloaded on Thu, 7 Mar 2013 06:52:12 AM All use subject to JSTOR Terms and Conditions 554 JOURNAL OF PALEONTOLOGY,V. 73, NO. 4, 1999 framework for a computer-assisted system of identification, in- volving large and very informative chitinozoan databases which plug neck [ chamber 0 carina include photos (Paris and Bernard, 1994). 0 E E ! may digitalized z spn The most common classifica- 2 | glabrous | linkage processes argument against suprageneric 35- tion is the poor knowledge of the biological significance of the chitinozoans. We, however, are less pessimistic than the 30 - = 14 sup- cJ porters of the alphabetic classification. Indeed, converging ar- 30 0 25 -2 guments have progressively emerged that convincingly support the hypothesis (first advocated by Koslowski, 1963) that chiti- 20-20 i- nozoans represent eggs of soft-bodied marine metazoans (Paris, 1981; Paris and Nolvak, in press), i.e., of the "chitinozoophor- 15- ans" sensu Grahn (1981). This explanation, however, is not yet unanimously accepted, and other diverse hypotheses (see dis- 5-5 I cussion in Miller, 1996) have been proposed, e.g., a possible I relationship with cysts of tintinids (Reid and John, 1981), fungi Characters theory (Locquin, 1976), or rhizopod affinities (e.g., Cashman, 1990). Because the two latter hypotheses are based either on FIGURE4-Histogram of frequency of the characters recorded in the 56 unreliable arguments or on obvious misinterpretations of chiti- selected chitinozoan genera. nozoan morphology, they are not considered here. Nevertheless, regardless of the affinities or the biological significance of chi- tinozoans, these microfossils are in any case related to fossil characters shared by the greatest number of genera and those organisms. The morphology of the chitinozoans is certainly ge- restricted to a single genus (Fig. 4). A general database was set netically controlled, as demonstrated by the occurrence of pop- up as a table of the morphologic characters regarded as the "var- ulations, that are statistically homogenous during time intervals iables" and reported in columns. In all calculations, genera were ranging from several hundred thousand to a few million years. referred to as "individuals" and reported on rows. These populations display diagnostic morphologies, indicating A better approach would have been to apply these sorting that they belong to the same species. Because the time intervals procedures to all known chitinozoan species and subspecies; involved exceed the usual duration of the life of any individual however this was not undertaken because too many species are by numerous orders of magnitude, they clearly represent nu- poorly described. merous successive generations of individuals. The stability of Weight of the different morphologic characters.-The mor- their morphology through time is controlled by genetic mixing phologic characters we selected were generally of the same (i.e., interfecundity of the organisms producing each "species" "weight" for each calculation when they were taken separately; of chitinozoans) that was efficient enough to prevent speciation all were of Boolean type (i.e., on the general table they are processes for periods of time corresponding to the total-range of quoted by "1" if present, and by "0" if absent). However, some the involved species. In addition, the existence of evolutionary criteria were directly subordinate to the presence of another, processes is unquestionably demonstrated by the short range of which was therefore considered to be of a higher magnitude, many chitinozoans species, making the group an outstanding e.g., the characteristics "spines in rows" or "spine in crown" biostratigraphic tool. Therefore, we are convinced that the mor- (second order characters) were subordinate to the existence of a phology of chitinozoans reflects evolutionary processes just as "spiny wall" (first order character). Similarly, the criterion in any complete organism, even if the tempo might be slightly "anastomosed" (third order character) was subordinate to the different. Consequently, the usual rules, especially those of clas- existence of "processes", themselves depending on the exis- sification used for "around" true organisms, should also be ap- tence of a "spiny wall". For taxa sharing this feature, the or- plied to the chitinozoans. namentation of the surface of the vesicle had a "higher weight" Sorting of the available morphologic characters.-In order to (three levels of answers) than the character "glabrous" shared select the most suitable characters for generic and suprageneric by other taxa (only one level of answer). Other characters were classification of the chitinozoans, we list the criteria used by totally dependent on each other, e.g., "operculum" and "neck previous authors for the definition of the 143 previously pub- absent". Criteria related to the shape of the chamber, e.g., "cy- lished chitinozoan genera. We eliminated from this general sur- lindrical," "claviform", and to a lesser extent "conical", were vey the redundant characters, i.e., those which correspond with closely tied (see Fig. 6) to the character "neck inconspicuous" fragmentary or dissociated descriptors of a single morphologic because in this type of chamber, neither the flexure nor the feature (e.g., "rounded base," "rounded flanks," and "rounded shoulder were well distinct enough to indicate precisely the po- margin" for a spherical chamber). We excluded dubious ele- sition of the boundary between the chamber and the neck. ments or features obviously related to misinterpretation or to Some discrepancies may have been introduced by a few fea- artifacts of preservation (e.g., crystal casts, folding of the vesicle tures satisfying several states. This concerned exclusively the wall). After this preliminary sorting we ran a number of statis- shape of the chamber and results both from some old definitions tical analyses on the remaining data set in order to control the (not conforming with the sorting procedure adopted here) and "weight" of each character and to identify the most definitive from the existence of intrageneric, and even intraspecific varia- criteria. The ultimate aims were to detect morphological features tions of this parameter. Therefore, several genera may have in- that allow the separation of all genera by at least one character cluded up to three states for the "shape of the chamber" (e.g., and can be used to cluster large groups of genera (e.g., subfam- "lenticular," "conical," and "ovoid" for Ancyrochitina). These ilies, families, and order). multiple answers influence the results of the multivariate anal- We utilized different techniques and software for simple sta- ysis (Fig. 6). tistical analyses (histograms), multivariate analysis [e.g., Facto- Frequency histogram.-The histogram (Fig. 4) of all of the rial Analysis of Correspondence (FAC), Ascendant Hierarchic 36 characters finally selected reveals that only 5 variables are Classification (AHC)], or cladistic approach. These sorting tech- restricted to only one genus ("widened base" in Pistillachitina; niques were employed in order to identify both the morphologic "bulblike ampoule" in Siphonochitina; "cell-like processes" in

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Alpenachitina Axis 2 Armigutta Ancyrochitina (29.1%) Eisenackitina Angochitina Kalochitina Clathrochitina Orbichitina Fungochitina Muscochitina .-o0 Ordochitina - neck spinyspi - 0.98 Plectochitina Salopochitina present Ramochitina Vinnalochitina Sommerochitina 0 Anthochitina Cyathochitina Lagenochitina Parisochitina operculum Pellichitina m 0 Prosome neck absent i* --- Sagenachitina g u . --I I Saharochitina o 0 050 1.0 Axis 1 Sphaerochitina < (50.2%) Urochitina

Acanthochitina glabrous Belonechitina Hercochitina Pogonochitina Spinachitina O

neck * l inconspicuous - -1.1 Baltochitina Laufeldochitina Clavachitina Pistillachitina Conochitina Rhabdochitina Eremochitina Siphonochitina Euconochitina Tanuchitina Hyalochitina Velatachitina

FIGURE5-Factorial Analysis of Correspondence(FAC) of the chitinozoangenera (open circles) and of seven characterstates (open stars)of the aperturalplug, neck, and wall surface.

Plectochitina; "anastomosed hollow processes" in Clathrochi- would allow us to separate all of the selected genera by at least tina; and "crown of spines on the shoulder" in Alpenachitina). one character. In contrast, 12 variables among the 36 selected are present in at Based on the Factorial Analysis of Correspondence (FAC) least 10 genera, and two of them ("prosome" and "glabrous diagrams, we tested different combinations of characters (Figs. wall") are shared by 35 genera among the 56 accepted herein. 5-7). Some are totally independent from each other (e.g., neck In the construction of this diagram (Fig. 4) we dissociated some differentiation/ornamentation, chamber shape/occurrence of a morphologic elements into their two mutually exclusive char- carina). On the other hand, the FAC diagram of the variables acteristics. The apertural plug was therefore divided into "oper- dealing with the apertural plug, the neck differentiation, and the culum" and "prosome", while the wall surface was defined by shape of the chamber indicates clear relationships between these its two exclusive states "glabrous" and "spiny wall". In both variables (Fig. 6). cases, the sum of the positive ("1") and negative ("0") answers The most discriminate combination involves only four char- is 56, i.e., the total number of accepted chitinozoan genera. In acters, i.e., "operculum," "neck present," "neck inconspicu- some other cases (e.g., the chamber shape) the sum of the dis- ous," and "spiny wall". It yields a perfect clustering of the sociated answers (i.e., "spherical," "lenticular," "hemispheri- genera into six groups with 43.7 percent of the variance ex- cal," "ovoid," "claviform," "cylindrical," and "conical") is plained by the first axis and 40.9 percent by the second axis. It much greater than the total number of the selected genera, be- is worth noting that a similar clustering is obtained when using cause some of them have several states (see above). characters complementary to those chosen for this combination Multivariate analysis.-We used the multivariate analysis (the answer "operculum present" is identical to "prosome ab- programs ANALMUL program for Factorial Analysis of Cor- sent" and the lack of a "spiny wall" implicates a "glabrous respondence (FAC) on Macintosh (Febvay and Bonnot, 1991), wall"). Consequently, the introduction of all the variables de- and STAT ITCF program for the construction of Ascendant Hi- scribing the first order characters (apertural plug, characteristics erarchic Classification (AHC using Euclidean distances). We ran of the neck, surface of the vesicle wall) does not improve the these successively with different combinations involving some, distinctiveness of the clusters. In the corresponding FAC, using or all, of the characters quoted in the 56 selected genera in order seven variables, the main axis explains 50.2 percent of the var- to distinguish the most diagnostic and discriminating characters. iance whereas the second axis explains only 29.1 percent of this Our ultimate goal was to detect morphological features that variance (Fig. 5).

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Axis 2 Angochltina Atmorncocht'tna the morphologicalchanges of the chitinozoan vesicle through M,'scochtirna (19.4 %) Armaguta Par!soch:iti;na Sa iopoch,tina C,utlc.Itir..a time, we the first occurrenceof successive major inno- neck plotted C.nga. .h;ina Desmtoct;tna Pe'iichitrna present 1.4 Sommerochr;na Eisenackh tina vations recordedfrom the appearanceof the group in the Tre- Hamocm,itvina , Ancyrochitina Utr.och,ti,na madoc to its extinctionin the Famennian 9.1-2). Urmch:itma topmost (Fig. ?!?na,oc.h:

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Axis 2 (9.3%) 5 hi 1.00 Marga.

Vinrnalo. Eisena.

Urno. Desmo. 1rmigutta e~ Cuti. Kalo. Salopo. Bursa. g

Lino. - Uro. i Ordo. Orbi.

Armorico. - Cingulo. Sphaero. o-- Ango. Ramo.

.:::. ";:::::::::.:: . : :?::::::: . .I. i- Sommero. -1.00 Saharo. Axis 1 - Pseudoclathro. "f]Pelli. Ancyro. (10.7%) Eremo. 0 Lageno.Lageno. '-'"Plecto. I ll Rha,bdo. Alpena. Belone. -- Clathro.

\ Musco. Siphono. 9. Cono. LVelata. / Acantho. Laufeldo. Pistilla. Balto. Pogono.

FIGURE7-Factorial Analysisof Correspondence(FAC) of the chitinozoangenera (open circles) and of the 36 morphologicalvariables of the vesicle. The subfamiliesare surrounded.Dark grey = Desmochitinidae;light grey = Lagenochitinidae;open = Conochitinidae.Rectangles indicate monogenericsubfamilies.

identified to genus and sometimes even to species level, based diachronism of the appearance of apomorphic characters in sev- exclusively on the external aspect of their eggs exclusively. eral chitinozoan families. Since the characteristics of the eggs of living metazoans can be Some apomorphic characters occur earlier in the Conochitin- used in generic identification, the morphology of the egg may idae and the Lagenochitinidae than in the Desmochitinidae, at be fairly strictly tied to the processes of speciation. Hence, the least up to the global faunal crisis at the Ordovician-Silurian relationships existing between different metazoan genera should boundary (see above). This is illustrated in northern Gondwana also be reflected by their eggs. Based on this argument, we have (Fig. 9.1) by the apomorphic character "carina below margin," tried through a cladistic approach to depict the relationships be- which is already present in the middle Arenig in the Conochi- tween chitinozoan genera (Fig. 10). The application of the cla- tinidae (e.g., Tanuchitina), whereas it appears for the first time distic principle to the chitinozoans is relatively straightforward in the late Abereiddian in the Desmochitinidae (e.g., Armori- as long as the procedure is applied to taxa within the same fam- cochitina). Conversely, after the topmost Ordovician faunal ily. However, difficulties arise when we attempt to include all event, the Desmochitinidae show the best potential of innova- the chitinozoan genera in a single cladogram. This is due to a tion, e.g., the apomorphic character "reticulum with perforated

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558 JOURNAL OF PALEONTOLOGY,V. 73, NO. 4, 1999

Acanthochitina - 4) or more sophisticatedmultivariate analysis (Figs. 5-8), re- Belonechitina vealed which charactersare shared by the greatest numberof Hercochitina taxa. The subdivisionis the Pogonochitina highest providedby aperturalplug, Spinachitina - i.e., either a prosomeor an operculum.Together with the cham- Alpenachitina ber, aperture,neck and glabrouswall, the prosome was one of Ancyrochitina the basic characteristicsof a chitinozoanvesicle when the . very Clathrochitina group appearedduring the Tremadoc.The neck and the surface Sommerochitina - Plectochitina i of the walls themselves showed modificationsas early as those Angochitina of the aperturalplug (Fig. 8). However,for the stability of the Ramochitina nomenclature,as recommendedby the InternationalCode of Muscochitina Zoological Nomenclature,we preferthe preservationof the ple- Fungochitina siomorphic characters "prosome" and "operculum" for the Siphonochitina Eremochitina | highest supragenericsubdivision of the group of the chitino- Baltochitina T0 zoans. The basic reason for this is because Eisenack(1972) had Tanuchitina - alreadyused them, as the diagnosticelements of his widely ac- Laufeldochitina - cepted Prosomatiferaand Operculatiferaorders respectively. Hyalochitina - o Clavachitina- This scheme has been used for more than 20 years by almost Conochitina - 2 all the chitinozoanworkers who have accepted and used a su- Rhabdochitina - pragenericclassification (see discussion in Paris, 1981; Achab Pistillachitina- et al., 1993), and there are no serious reasons to change it. Euconochitina CD Other featuresshared several Velatachitina - highly diagnostic by genera(see Pellichitina - Figs. 5-6) are the various characterstates applied to the differ- Anthochitina --- entiationof the neck (i.e., "neckpresent," "neck absent," "neck Cyathochitina - inconspicuous"),the surfaceof the vesicle wall (i.e., "glabrous" Sagenachitina - and "spiny wall") and the shape of the chamber(i.e., "lentic- Parisochitina - Sphaerochitina - - - ular," "hemispherical," "spherical," "ovoid," "claviform," Saharochitina "conical," "cylindrical").The most distinctivecombinations of Lagenochitina charactersare those that relate neck differentiationto apertural Urochitina plug. These combinationsallow the division of the Chitinozoa Vinnalochitina into three main here as families, i.e., the Eisenackitina categories regarded Ordochitina " Conochitinidae,the Lagenochitinidae,and the Desmochitinidae Kalochitina (Fig. 11). Armigutta Salopochitina CLASSIFICATION OF THE ACCEVI'ED GENERA Orbichitina Cutichitina Discussion.-In numerouspublications, original generic di- Bulbochitina ( agnoses are too vague or introduceinconstancies in application Bursachitina of terminology.Consequently, under the heading "diagnosis", Calpichitina c: and in order to promote a more standardizedterminology, we Desmochitina have indicated the discriminatecharacter(s) (=diagnostic fea- Ollachitina l. we Hoegisphaera tures) for each genus accept. Armoricochitina The emendeddiagnoses proposedby Paris (1981, p. 110) re- Pterochitina spectively for the Desmochitinidae,the Conochitinidaeand the Pseudoclathrochitina Lagenochitinidaeare strictlyadopted here. Cingulochitina of Ordoviciantaxa is referenceto Urnochitina Age assignment given by Margachitina the recently modified British chronostratigraphy(Fortey et al., Linochitina 1995).

FIGURE8-Hierarchic AscendantClassification (HAC) using Euclidean SYSTEMATIC PALEONTOLOGY distancesfor the 56 selectedgenera and the characteristics of the ap- erturalplug, neck, surface pattern, and vesicle structures (12 states). Order PROSOMATIFERAEisenack, 1972 Darkgrey = Operculatifera;light grey = Prosomatifera. Family CONOCHITINIDAEEisenack, 1931 emend. Paris, 1981 Subfamily CONOCHMTININAEParis, 1981 Genus CLAVACHITINATaugourdeau, 1966 expansion" occurs as early as the Wenlock(Al-Hajri and Paris, Type species.-Rhabdochitina claviformis Taugourdeau, 1998) in the Desmochitinidae(e.g., Pseudoclathrochitina)but is 1961. Holotype in Taugourdeau,1961, p. 150, pl. 4, fig. 69, not reported in the Lagenochitinidaebefore the late Givetian Avensac 101 borehole, 2051 m, Avensac Formation,Llandeili- (e.g., Parisochitina).Similar observations are made for the apo- an, south-westernFrance, collections of the Museum National morphic character"sheathing processes" which occur for the d'HistoireNaturelle de Paris, France. first time in Salopochitina,a Late Llandovery-EarlyWenlock Diagnosis.-Conochitinidae with a glabrousclaviform cham- Desmochitinidae,but only in the late Frasnian-earlyFamennian ber and withoutmucron. for Sommerochitina,a genus belonging to the Lagenochitinidae. Occurrence.-Ordovician. This diachronismof some apomorphiccharacters leads to poorly resolved cladogramswhen all chitinozoangenera are an- Genus CONOCHmNAEisenack, 1931 alyzed together.This may be relatedto a possible polyphyletism emend. Paris, Grahn,Nestor, and Lakova of the Chitinozoa. Type species.-Conochitina claviformis Eisenack, 1931. Lost Hierarchy of the used criteria.-Mathematical treatmentsof holotype in Eisenack, 1931, p. 84, pl. 1, fig. 17, from erratic all the morphologicaldata, using eithera simple histogram(Fig. graptoliticrocks ("Graptolithengestein");neotype: in Eisenack,

This content downloaded on Thu, 7 Mar 2013 06:52:12 AM All use subject to JSTOR Terms and Conditions Eisenau kilina ORDOVICIAN SILURIAN DEVONIAN Vinnialohlitifll r - -n Chronostratigraphy Ordoch itina (0 . Innovations ITI C. '

O/'bicliitiitt cT1 .-. .------.---. ------claviformchamber 0 ....------neck present Salopo'hitina * ------prosome ------0 a 0 .. ----. ---o ------glabrous Ariuiigiitltt ------cylindricalchamber A rinorlcochiiinui ------ovoidchamber ------.----0------.------mucron .-----* ------o------conicalchamber PArnetochitil, t -. ------o ------copula O i ------0 ------sleeve N) Ptfeochitilua -..-- .---- .------o ------.------carina II ------o ------carina on margin Cihl '/itilna ------o ------carina below margin ..------o ------neck absent rT- n0 Mttf^/;f)t'/ ifli// ------operculum Urnochlfiil ...------. ..------.------o------perforatedcarina It -...... -.----- o ------spines Li)wno hitinat ------...... ------.------chamberhemispheric 9<0 Z.------. ..------bulb Ctilichitilla a ..------0 -..------lenticularchamber r.i --o--? ------* ------sphericalchamber ------0D ....-. ------.------laciniated carina ------* o ------crests con 1hi'/i1sch/ lil ------.-.------o -----o ------processes .. ) .------_ ------. ------mesh likeornament BifilochIififii7 a0 ------anastomosedprocesses ------cell likeprocesses Hoegisphaura C ------collaretteabsent ------o ------operculum/pedunclefixed Calpichitilua rio ..--- ....------o ------peduncle o1 - - ..------sheathingprocesses 0'llitilitifind ..... ----.----.------carina round the aperture I ...... ------reticulum . ------o ------evaginatedchamber ------* ------ornaments in 3crowns c 00 tIT ------claviformchamber -.------neckpresent -...- . ------prosome -. 0 ------glabrous .------.------. ------cylindricalchamber I1 -----.------.----.------ovoid chamber rI 0De -----o ------0o ------mucron P. ------conical chamber 0T tTi ------0ri c -----.------.--- 0------copula -C ------sleeve -- . o ------carina - -- - .------.----.------.------o------carinaon margin (A Q ------* ---- -o -.----.------carina below margin ? -. 0 - . -.o ------neckabsent o~~~~ ~ Ct -----o------.------operculum 0Q m-Ce -----...------..------.------perforatedcarina i) z7 ..-----.. o ------...... spines .. ------chamber hemispheric -- W(D 1-3 ------0 ------bulb 0x ...---- .--- *- -.------lenticularchamber io ------...... ------..... ---...... --- o ------spherical chamber 0 ------laciniatedcarina ------.------crests r-EL -----.....------.-- .------.------.------processes 0 0k4 .------..------mesh like ornament -. . 0 ------anastomosed 4W L w..-----W----:------.------* ------processes ..el.r0 ------cell like processes p Oe .. ------.------1-o ------o ------collaretteabsent ON~ 0 ----- . ------.--.--.------operculum/peduncle fixed 0n ------o------peduncle ------sheathingprocesses ------.------o------carinaround the aperture ------reticulum ------evaginatedchamber ------.------ornamentsin 3 crowns

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Order Family Sub-family Genera Plug Neck differentiation Chambersurface Chambershape and arrangement of ornamentation lenticular...... Calpichitina spherical ------...... -- ---.... ---... Hoegisphaera Desmochitininae hemispherical...... Bulbochitina (glabrous) conical --...... Bursachitina ovoid ...... Desmochitina cylindrical --...------...------.....-- - Ollachitina Cutichitininae ovoid------Cutichitina (sleeve) 4 lenticularto spherical...... Pterochitina Pterochitininae ovoid (below margin)-...... Armoricochitina - (carina) conical (reticulum,perforated) ...... Pseudoclathrochitina I DESMOCHITINIDAE conical to ovoid (on margin) ---.....------..--Cingulochitina ^; $-4 (no neck) lenticularto spherical(peduncle) ...... Margachitina Margachitininae ovoid...... Urnochitina (copula) claviformto cylindrical ...... Linochitina 0 hemispherical...... Vinnalochitina Eisenackitininae conical ...... Kalochitina (spiny) conical (with crests)------..------Ordochitina ovoid -..------...------..---Eisenackitina conical ...... Orbichitina Orbichitininae ovoid...... Armigutta (processes) I conical to ovoid (sheathing)-.------.- Salopochitina conical-----.---.------..--.--...-.- - Euconochitina conical to claviform (with mucron)...... Conochitina Conochitininae claviform------Clavachitina (glabrous) cylindrical (with widened base) ------Pistillachitina cylindrical ------...... Rhabdochitina Velatachitininae claviform --...-----...------(sleeve) I Velatachitina Eremochitininae claviform...... Eremochitina (copula) claviformto cylindrical(bulb) ----. ..-----Siphonochitina CONOCHITINIDAE conical to cylindrical(on margin)...... Hyalochitina ( flexure Tanuchitininae claviform (below margin)------Laufeldochitina unconspicuous) (carina) conical to cylindrical(perforated) ------Baltochitina cylindrical(below margin)...... ------Tanuchitina Pogonochitininae *1conical (laciniated,on margin)------Pogonochitina a! carina/spiny) conical...... Belonechitina -i Belonechitininae conical to cylindrical(mesh-like) ------Acanthochitina (spiny) conical ? I (with crests)...... Hercochitina Spinachitininae conical to cylindrical ------Spinachitina o; 0o (processes) 0O rt./_ lenticularto conical------Saharochitina CA? 1- Lagenochitininae spherical...... Sphaerochitina (glabrous) t ovoid to cylindrical------...... ------Lagenochitina i conical to hemispherical(laciniated) -. - Anthochitina Cyathochitininae \I conical to hemispherical(perforated) --....- Sagenachitina (carina) conical to hemispherical(complete) ...... Cyathochitina ovoid (reticulum,perforated) ------. ....---Parisochitina Pellichitininae LAGENOCHITINIDAE (sleeve) ovoid------Pellichitina (flexureconspicuous) Urochitininae ovoid (peduncle).------Urochitina (copula) lenticularto conical...... Fungochitina ovoid------Angochitina Angochitininae ovoid (mesh-like) ---...------..-.. - Muscochitina (spiny) ovoid (with crests)-. ---. - .-.------Ramochitina lenticularto conical -.------.------.------Ancyrochitina conical (anastomosed).------Clathrochitina , Ancyrochitininae conical to ovoid (cell-like) ------Plectochitina (processes) ovoid to cylindrical(3 crowns) ------Alpenachitina claviform(sheathing) ..-...... ------Sommerochitina

FIGURE11-Suprageneric and genericclassification of the chitinozoans.

This content downloaded on Thu, 7 Mar 2013 06:52:12 AM All use subject to JSTOR Terms and Conditions PARISET AL.-REVISEDCHITINOZOAN CLASSIFICATION 561 formationfrom the RheinischenSchiefergebirges, Llanvimrn-ear- turning reticulated distally and extending more or less perpen- ly Caradoc,Germany, collections of the SenckenbergMuseum, dicularto the axis. Frankfurtam Main, Germany. Description.-Long and slenderchitinozoan (length of the ho- Diagnosis.-Conochitinidae with a glabrous cylindrical lotype: 440 microns,chamber diameter: 60 microns)with a sub- chamberended by a lenticularto hemisphericalwidening. cylindricalto slightly conical chamber;cylindrical neck ended Occurrence.-Middle-Upper Ordovician. with straight lips; chamber bottom flat; wall surface entirely Discussion.-It must be stressedthe transitionbetween cham- smooth; perforatedto reticulatedcarina erected on the margin; ber and neck (positionof the prosome)is within the tubularpart this fairly short membranouscarina extends more or less per- of the vesicle. pendicularlyto the flanks. Etymology.-Dedicated to our colleague Jaak N1olvak(Esto- Genus RHABDOCHITINAEisenack, 1931 nian Academy of Sciences, Tallinn,Estonia) for his outstanding Type species. -Rhabdochitina magna Eisenack, 1931. Lost investigationson Ordovicianchitinozoans from the Baltic area. holotypein Eisenack, 1931, p. 90-91, pl. 3, fig. 18, erraticBaltic Holotype.-fig. A, pl. 5 in N1olvak and Grahn (1993), Ch. limestones ("Ostseekalk");neotype in Eisenack, 1962, p. 292- 1416/7631, collections of the Institute of Geology, Estonian 293, pl. 14, fig. 1, erraticBaltic limestones,Ordovician, collec- Academy of Sciences, Tallinn,Estonia; late Abereiddian,Las- tions of the Geologisch-PalaontologischenInstitute der Univer- namagi Stage, Voo Formation,Rapla borehole (depth 179.2 m), sitat, Tiibingen,Germany. south of Tallinn,Estonia. Diagnosis.-Conochitinidae with a glabrouselongated cylin- Occurrence.-Late Abereiddian. vesicle. drical Discussion.-No other chitinozoan species has a subcylindri- Occurrence. -Ordovician. cal vesicle and a reticulated carina. Discussion.-Mucron may occur. Subfamily EREMOCHITININAEParis, 1981 Genus HYALOCHITINAParis and Grahn new genus Genus EREMOCHITINATaugourdeau and de Jekhowsky,1960 Type species.-By original designation: Cyathochitina hyalo- baculata and de Type species.-Eremochitina Taugourdeau Eisenack, 1959. Holotype in Eisenack, 1959, p. 11-12, pl. 1960. Lost in and de Jek- phrys Jekhowsky, holotype Taugourdeau 2, fig. 6, Cincinnati Group, Cincinnati, Ohio, late Maysvillian- 1960, 1228, 8, 107, Or-I borehole,core sam- howsky, p. pl. fig. Richmondian (early Ashgill), collections of the Geolo- from 2930 m, Sahara, Algeria; neotype: in Tau- early ple Arenig, Institute der Universitat, Tubingen, gourdeau,1967, pl. 1, fig. 19, (DD. 52), same referencesas for gisch-Palaontologischen the collections of the MuseumNational d'Histoire Na- Germany. holotype, with a conical to turelle de Paris, France. Diagnosis.-Conochitinidae cylindrical a membranous carina on the Diagnosis.-Conochitinidae with claviformglabrous chamber chamber provided with complete with a tubularcopula. margin. Occurrence.-Lower Ordovician. Occurrence.-Upper Ordovician. Discussion.-This new genus is separated from Tanuchitina Genus SIPHONOCHITINAJenkins, 1967 Jansonius, 1964, which has a carina below the margin, and from Type species.-Siphonochitina formosa Jenkins, 1967. Holo- Cyathochitina Eisenack, 1955b emend. Paris, Grahn, Nestor, and type in Jenkins, 1967, p. 469-471, pl. 75, fig. 2, Shelve area, Lakova herein, which has a conspicuous flexure. upper Hope Shale, early Abereiddian,Shropshire, England, S 17593, collections of the Centrefor PalynologicalStudies, Uni- Genus LAUFELDOCHITINAParis, 1981 versity of Sheffield, England. Diagnosis.-Conochitinidae with a claviform to cylindrical Type species.-Cyathochitina stentor Eisenack, 1937 (lost ho- chamberwith a membranousbulb. lotype in Eisenack, 1937, p. 221-222, pl. 15, fig. 2, grayish Occurrence. -Lower-Middle Ordovician. Ordovicianlimestones; neotype in Eisenack, 1962, p. 300, pl. 14, fig. 10, allochthonousKukruse equivalents, Halde bei Met- Subfamily TANUCHITININAEParis, 1981 zingen, Wurtemberg, Germany, collections of the Geologisch- Genus BALTOCHITINAParis and Grahn new genus Palaontologischen Institute der Universitat, Tiibingen, Germa- Type species.-By original designation: Baltochitina nolvaki ny). Paris and Grahn new species. Holotype: Sagenachitinasp. in Diagnosis.-Conochitinidae with a claviform, glabrous cham- Nolvak and Grahn, 1993, pl. 5, figs. A and B, Rapla borehole, ber and a complete, flaring membranous carina, below the mar- Voo Formation,Lasnamagi Stage, late Abereiddian,south of Tal- gin. linn, Estonia; collections of the Instituteof Geology, Estonian Occurrence. -Ordovician. Academy of Sciences, Tallinn,Estonia. with a conical to Diagnosis.-Conochitinidae cylindrical Genus TANUCHITINAJansonius, 1964 carinaon the chamberwith a perforated margin. emend. Paris, Grahn,Nestor, and Lakova Etymology.-from Baltic, its type area. Occurrence.-Middle Ordovician. Type species.-Tanuchitina ontariensis Jansonius, 1964. Ho- Discussion.-This genus differs from Sagenachitina Jenkins, lotype in Jansonius, 1964, p. 910-911, pl. 1, fig. 6, cuttings from 1970a and from Parisochitina Boumendjel, 1985, which have a Imperial-Calvan Anderson no. 9-6 borehole, depth 738 m (2,420 well-definedneck. ft), Anderson Township, Essex County, Ontario, Canada, collec- tions of Oil Limited, Alberta, Canada. BALTOCHITINANOLVAKI Paris and Grahnnew species Imperial Calgary, Diagnosis.-Conochitinidae with a cylindricalchamber and a Sagenachitina sp. NOLVAK AND GRAHN, 1993, pl. 5, figs. A and B complete membranouscarina below the margin. Diagnosis.-Conochitinidae with an elongate subcylindrical Occurrence.-Ordovician-Silurian. vesicle (ratio of vesicle length to chamberdiameter >five), a Discussion.-The conical forms with a carina on the margin smooth wall surface, straightlip, sharpmargin bearing a carina are transferred into Hyalochitina Paris and Grahn, n. gen.

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SubfamilyVELATACHITININAE Achab, Asselin and SubfamilySPINACHITININAE Paris, 1981 Soufiane, 1993 Genus SPINACHITINASchallreuter, 1963 Genus VELATACHITINAPoumot, 1968 emend. Paris, Grahn,Nestor, and Lakova Type species. -Velatachitina nebulosa Poumot, 1968. Holo- Type species.-Conochitina cervicornis Eisenack, 1931. Lost type in Poumot, 1968, p. 50-51, pl. 1, fig. 9, borehole SN-1, holotype in Eisenack, 1931, p. 89, pl. 2, fig. 12, erraticcalcar- core sample from 3,082-3,083 m (referredto "Llandeilo"by eous sandy siltstones; proposed neotype: Spinachitina cervicor- the author,but likely Abereiddian),Tunisia, CZ 244, collections nis in Nolvak and Grahn, 1993, pl. 3, fig. A, Rapla borehole, of the Service Palynoplanctologiquede la SNPA-Elf,France. core sample from 142.70 m, Kahula Formation,Keila Stage Diagnosis.-Conochitinidae with a claviformchamber envel- (early Caradoc),south of Tallinn,Estonia, collections of the In- oped within a membranoussleeve. stitute of Geology, EstonianAcademy of Sciences, Tallinn,Es- Occurrence.-Lower-Middle Ordovician. tonia. Discussion.-The membranoussleeve may extendbeyond the Diagnosis.-Conochitinidae with a conical to cylindrical margin. chamber bearing a crown of processes. Occurrence.-Upper Ordovician-LowerSilurian. SubfamilyBELONECHITININAE Paris, 1981 Discussion.-In additionto the emendationof Laufeld(1967), Genus ACANTHOCHITINAEisenack, 1931 the genus is here restrictedto formswithout conspicuous flexure. CoronochitinaEisenack, 1965 is here regardedas a junior syn- Type species.-Acanthochitina barbata Eisenack, 1931 (Lost onym of Spinachitina. holotypein Eisenack, 1931, p. 82-83, pl. 1, fig. 10, erraticlime- stone from the Baltic ["Ostseekalk"];proposed neotype: in N61- Hullo 21.0 late Vormsi Family LAGENOCHITINIDAEEisenack, 1931 vak, 1980, pl. 29, fig. 1, borehole, m, emend. 1981 Ch. 701/5670, collections of the Paris, Stage (early Ashgill), Estonia, LAGENOCHITININAE 1981 of of the Estonian of Science, Tal- Subfamily Paris, Institute Geology Academy 1931 Estonia. Genus LAGENOCHITINAEisenack, linn, emend. Paris, Grahn,Nestor, and Lakova Diagnosis.-Conochitinidae with a conical to cylindrical chamberand raisedmeshlike ornamentation; may be surrounded Type species.-Lagenochitina baltica Eisenack, 1931. Lost with a membranoussleeve. holotype in Eisenack, 1931, p. 80-81, pl. 1, fig. 1, erraticOr- Occurrence.-Middle Ordovician(?)-Upper Ordovician. dovicianBaltic limestones("Ostseekalk"); neotype in Eisenack, 1959, p. 2, pl. 3, fig. 6, from Ordovicianerratic limestones of Genus BELONECHITINAJansonius, 1964 the Baltic shore ("Ostseekalk"),collections of the Geologisch- Instituteder Universitat,Tuibingen, Germany. micracantha robusta Ei- Palaontologischen Type species.-Conochitina subsp. Diagnosis.-Lagenochitinidae with an ovoid to cylindrical senack, 1959. in Eisenack 1959, p. 9-10, pl. 3, fig. 4, Holotype glabrous chamber. Koppelmann,Saku Member of the WasalemmaFormation, Oan- Occurrence.-Lower Devonian. collections of the Senck- Ordovician-Upper du Stage, D3 (late Caradoc),Estonia, Discussion.-Forms with spherical chambers are Frankfurtam glabrous enberg Museum, Main, Germany. transferredinto emend. Paris, with a conical chamberand ran- SphaerochitinaEisenack, 1955a, Diagnosis.-Conochitinidae Grahn, Nestor, and Lakova, herein. domly distributedspines. Occurrence.-Lower Ordovician-Silurian. Genus SAHAROCHITINAParis and Grahn new genus Genus HERCOCHITINAJansonius, 1964 Type species.-By original designation: Fungochitina? jaglini Oulebsir and Paris, 1993. in Oulebsir and Paris, 1993, 1964. Ho- Holotype Type species.-Hercochitina crickmayi Jansonius, 274-279, pl. 4, fig. 1, Gd-1 bis borehole,north of Hassi Mes- in 9, Gamache p. lotype Jansonius, 1964, p. 908-909, pl. 1, fig. saoud, north-eastern Sahara, Hassi Touareg Formation, Azzel PrincetonLake 1 core from 210 m (690 ft.), borehole, sample Member, core sample at 4,033.50 m, late Abereiddian, IGR VaurealFormation, Ashgill, Anticosti Island, Canada;collec- collections of the Institut de de Canada. 60509 (R 41), Geologie tions of ImperialOil Limited,Calgary, Alberta, l'Universit6 de Rennes, France. Diagnosis.-Conochitinidae with a conical chamberand dis- with a conical to len- ornamenta- Diagnosis.-Lagenochitinidae glabrous tinct crests (vertical rows of spiny or membranous ticularchamber. tion). Occurrence.-Middle Ordovician-Silurian. Occurrence.-Middle Ordovician. Ordovician-Upper Discussion.-This genus differs from Fungochitina Taugour- deau, 1966 in being smooth and from PistillachitinaTaugour- SubfamilyPOGONOCHITININAE new subfamily deau, 1966, which displays a tubularchamber with an antiaper- Diagnosis.-Conochitinidae with a spiny ornamentationand turalwidening. a carina. Genus SPHAEROCHITINAEisenack, 1955a Genus POGONOCHITINATaugourdeau, 1961 emend. Paris, Grahn,Nestor, and Lakova Type species.-Pogonochitina simplex Taugourdeau, 1961. Type species.-Lagenochitina sphaerocephala Eisenack, Holotype in Taugourdeau,1961, p. 148, pl. 4, fig. 59, Avensac 1932. Lost holotype in Eisenack, 1932, p. 271, pl. 12, fig. 14, 101 borehole, 1,902 m, Avensac Formation,Llandeilian, south- erraticBeyrichia Limestone ("Beyrichia-kalk");neotype in Ei- western France,collections of the MuseumNational d'Histoire senack, 1955a, p. 162, pl. 1, fig. 6, erraticBeyrichia Limestone Naturelle de Paris, France. Representativespecimen in Paris, from the Baltic sea floor?, Pridoli, collections of the Sencken- 1996, pl. 1, fig. 9. berg Museum,Frankfurt am Main, Germany. Diagnosis.-Conochitinidae with a spiny conical chamber Diagnosis.-Lagenochitinidae with a glabrous spherical and with a laciniatedcarina on the margin. chamber. Occurrence.-Middle Ordovician-Silurian. Occurrence. -Upper Ordovician-Devonian.

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Discussion.-The spiny forms (spines exceeding two mi- Diagnosis.-Lagenochitinidae with an ovoid chamberenvel- crons) as well as those with a conical or a lenticularchamber oped by a membranoussleeve. are here excluded from the genus. Occurrence.-Lower-Middle Ordovician.

SubfamilyCYATHOCHITININAE Paris, 1981 Subfamily UROCHITININAEParis, 1981 Genus ANTHOCHITINAEisenack, 1971 Genus UROCHITINATaugourdeau and de Jekhowsky, 1960 Type species.-Anthochitina superba Eisenack, 1971. Holo- Type species.-Urochitina simplex Taugourdeau and de Jek- type in Eisenack, 1971, p. 452-454, fig. 1-1, erraticBeyrichia howsky, 1960. Holotype in Taugourdeauand de Jekhowsky, limestonesfrom the Baltic bottom?,Pridoli(?), collections of the 1960, p. 1233, pl. 11, fig. 160, boreholeGe-1, core samplefrom Geologisch-PalaontologischenInstitute der Universitat,Tubing- 1,586 m, Oued Saret Formation,latest Lochkovian,Sahara, Al- en, Germany.Representative specimens in Wrona,1980, pl. 26, geria, collections of the Museum National d'HistoireNaturelle figs. 1-9. de Paris, France. Diagnosis.-Lagenochitinidae with a conical to hemispherical Diagnosis.-Lagenochitinidae with an ovoid chamberbearing chamberwith a laciniated,spongy to cell-like carina. a long solid peduncle. Occurrence. -Upper Silurian-Lower Devonian. Occurrence.-Lower-Middle Devonian. Discussion.-Occasionally proximal perforations may exist. Subfamily ANCYROCHITININAEParis, 1981 Genus CYATHOCHITINA 1955b Eisenack, Genus ALPENACHITINADunn and Miller, 1964 emend. Paris, Grahn,Nestor, and Lakova Type species.-Alpenachitina eisenacki Dunn and Miller, Type species.-Conochitina Eisenack, campanulaeformis 1964. Holotype in Dunn and Miller, 1964, p. 725, pl. 119, fig. 1931. Lost holotype in Eisenack, 1931, p. 86-87, pl. 2, fig. 2, Dock Street erraticcalcareous to siltstoneof Ordovician 1, Clay Member,Alpena Limestone,Alpena, Mich- sandy age; neotype: USA, Middle Devonian, USNM collections 144880. in igan, Cyathochitina campanulaeformis Eisenack, 1962, p. 297-298, Diagnosis.-Lagenochitinidae with ovoid to pl. 14, fig. 5, from Harkunear Tallinn, likely lower Uhaku cylindrical Stage chamberand three distincthorizontal crowns of spiny ornamen- (Llandeilian),Estonia, collections of the SenckenbergMuseum, tation. Frankfurtam Main, Germany. Occurrence.-Middle Devonian. Diagnosis.-Lagenochitinidae with a conical to hemispherical glabrouschamber and with a membranouscarina on a complete Genus ANCYROCHITINAEisenack, 1955a sharpmargin. Discussion.-Conical species without a conspicuous flexure Type species.-Conochitina ancyrea Eisenack, 1931. Lost ho- are transferredinto HyalochitinaParis and Grahnn. gen. lotype in Eisenack, 1931, p. 88-89, pl. 4, fig. 4, erraticlime- Occurrence.-Ordovician-lower Silurian(Llandovery). stones of the Baltic ("Ostseekalk");neotype: Ancyrochitina an- cyrea in Eisenack, 1955a, p. 163-164, pl. 2, fig. 7, erraticBey- Genus PARISOCHITINABoumendjel, 1985 richia limestones from the Baltic bottom?, Pridoli; collections of the Type species.-Parisochitina perforata Boumendjel, 1985. SenckenbergMuseum, Frankfurt am Main, Germany. Holotype in Boumendjel, 1985, p. 164-166, pl. 2, fig. 5, bore- Diagnosis.-Lagenochitinidae with a lenticular to conical hole Taouratine-3,1,352 m, Gazelle Formation,late Devonian, chamberbearing a crown of nonanastomosedhollow processes Illizi Basin, south-easternSahara, Algeria, IGR 60319 (M 39), on the margin. collections of the Institutde Geologie de l'Universitede Rennes, Occurrence.-Upper Ordovician-UpperDevonian. France. Diagnosis.-Lagenochitinidae with an ovoid chamber and Genus CLATHROCHITINAEisenack, 1959 with a reticulumextending beyond the base as a perforatedca- Type species.-Clathrochitina clathrata Eisenack, 1959. Ho- rina. lotype in Eisenack, 1959, p. 15 pl. 1, fig. 3, Dalhem canal, Pen- Occurrence.-Upper Devonian. tamerusgotlandicus beds from southeast Slite Marls,late Shein- woodian, Wenlock,Gotland, Sweden, collections of the Geolo- Genus SAGENACHITINAJenkins, 1970a gisch-PalaontologischenInstitute der Universitat, Tiibingen, Type species.-Clathrochitina oblonga Benoit and Taugour- Germany. deau, 1961. Holotypein Benoit and Taugourdeau,1961, p. 1406, Diagnosis.-Lagenochitinidae with a conical chamberand a pl. 1, fig. 1, Amg-1 borehole, 1,482 m, "Complexeargilo-gre- crown of anastomosedprocesses on the margin. seux superieur"Formation, late Arenig, Sahara,Algeria, collec- Occurrence.-Silurian. tions of the Museum National d'Histoire Naturelle de Paris, Discussion.-In order to separate Clathrochitina from An- France. thochitina,the emendationproposed by Laufeld(1974) focusing Diagnosis.-Lagenochitinidae with a glabrous conical to on a membranouscarina (cingulum) is not followed. hemisphericalchamber and with a perforatedor reticulatedca- rina extendingfrom the margin. Genus PLECTOCHITINACramer, 1964 Occurrence.-Lower-Middle Ordovician. Type species.-Plectochitina carminae Cramer, 1964. Lost holotype in Cramer,1964, p. 346-347, 20, La Vid Subfamily PELLICHITININAEAchab, Asselin and 1993 pl. fig. 21, Soufiane, de Gordon section, upper San Pedro Formation, north- Genus PELLICHITINAAchab, Asselin and Soufiane, 1993 Pridoli, western Spain; neotype: Plectochitina carminae in Priewalder, Type species.-Desmochitina pellucida Benoit and Taugour- 1997, p. 77, pl. 2, fig. 1, pl. 4, figs. 1, 7-8, slide 1997/1/1 (M. deau, 1961. Holotype in Benoit and Taugourdeau,1961, p. 35.3); collections of the Geological Survey of Austria,Vienna. 1408-1409, pl. 4, fig. 37, Amg-1 borehole, core sample from Diagnosis.-Lagenochitinidae with a conical to ovoid cham- 1,577 m, lower part of the HamraQuartzite Formation, middle ber providedwith a crown of cell-like processes. Arenig, CentralSahara, Algeria, collections of the MuseumNa- Occurrence.-Upper Ordovician-Silurian. tional d'HistoireNaturelle de Paris, France. Discussion.-The processes may be anastomosed.

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Genus SOMMEROCHITINACosta Cruz and Quadros, 1985 emendationof this genus. GotlandochitinaLaufeld, 1974 which Type species.-Sommerochitina langei Costa Cruz and Qua- has the same charactersis thereforea junior synonym. dros, 1985. Holotypein Costa Cruzand Quadros,1985, p. 289- 293, pl. 1, fig. 1-2, 2-PM-1-MA borehole, core sample from Order OPERCULATIFERAEisenack, 1931 1,258-1,262 m, CabefnasFormation (in Longa Formationac- Family DESMOCHITINIDAEEisenack, 1931, emend. Paris, 1981 cordingGrahn, 1992), early Famennian,Parnaiba Basin, Brazil, Subfamily DESMOCHITININAEParis, 1981 collections of departamentode Geologia e Paleontologia,Museu Genus BULBOCHITINAParis, 1981 Nacional, Rio de Janeiro,Brazil. Type species.-Bulbochitina bulbosa Paris, 1981. Holotype in Diagnosis.-Lagenochitinidae with a claviformchamber with Paris, 1981, p. 134-135, pl. 35, fig. 10, la Lezais section, Bois- sheathingprocesses. Roux Formation, Aubriais Member, late Pragian, western Occurrence.-Upper Devonian. France,IGR 51138 (P 36/3), collections of the Institutde Geo- Discussion.-When partlybroken, threadlike expansions sim- logie de l'Universitede Rennes, France. ulate a deeply laciniatedcarina located below the margin. Diagnosis.-Desmochitinidae with a hemisphericalglabrous chamber. ANGOCHITININAEParis, 1981 Subfamily Silurian-Lower Devonian. Genus ANGOCHITINAEisenack, 1931 Occurrence.-Upper Type species.-Angochitina echinata Eisenack, 1931. Lost Genus BURSACHITINATaugourdeau, 1966 holotype in Eisenack, 1931, p. 82, pl. 1, fig. 7, erraticBeyrichia restrict.Paris, 1981 limestones ("Beyrichia-kalk")from the Baltic bottom?; neo- type:Angochitina echinata in Eisenack, 1964, p. 139, pl. 29, fig. Type species.-Desmochitina bursa Taugourdeau and de Jek- 10, from the topmost part of the Hemse beds, Ludlow, early howsky, 1960. Lost holotype, in Taugourdeauand de Jekhows- Ludfordian,Gotland, Sweden, collections of the Geologisch-Pa- ky, 1960, p. 1225, pl. 7, fig. 89, Tb-I borehole, core sample laontologischenInstitute der Universitat,Ttibingen, Germany. from 1,268 m, EarlyDevonian, Sahara, Algeria; neotype: in Tau- Occurrence. -Upper Ordovician-Devonian. gourdeau,1967, p. 256, pl. 1, fig. 3, Nm-1, core sample from Diagnosis.-Lagenochitinidae with ovoid chamberand ran- 1,200 m, EarlyDevonian (likely lower Emsian),Sahara, Algeria, domly distributedsimple or complex spines. collections of the MuseumNational d'Histoire Naturelle de Par- is, France). Genus FUNGOCHITINATaugourdeau, 1966 Diagnosis.-Glabrous Desmochitinidaewith a conical cham- Type species.-Conochitina fungiformis Eisenack, 1931. Lost ber. holotype in Eisenack, 1931, p. 89, pl. 2, fig. 17, erraticgrayish Occurrence.-Ordovician(?)-Middle Devonian. limestones ("grauer Kalk") referredto the lower Silurianbut Discussion.-We have adoptedthe position of Paris (1981, p. likely from the RakvereStage (late Caradoc);proposed neotype: 137) who restrictedthe genus to glabrousspecies. Conochitina fungiformis subsp. spinifera. Eisenack, 1962, p. 310, pl. 14, fig. 15, Rakvere Stage (late Caradoc),Wesenberg, Genus CALPICHITINAWilson and Hedlund, 1964 Estonia, collections of the Geologisch-PalaontologischenInsti- Type species.-Calpichitina scabiosa Wilson and Hedlund, tute der Universitat,Tubingen, Germany. 1964. Holotype in Wilson and Hedlund,1964, p. 164, pl. 1, fig. Diagnosis.-Lagenochitinidae with conical to lenticular 1, lower Sylvan Shale, south of Davis, Murray County, chamberand randomlydistributed spines. Oklahoma, early Ashgill, OPC 235-113-1, collections of the Occurrence.-Upper Ordovician-UpperDevonian. Universityof Oklahoma,Norman. Discussion.-Eisenack's original holotype for the speciesfun- Diagnosis.-Desmochitinidae with a glabrous lenticular giformis was spiny. The subspecies spinifera is therefore chosen chamber. as the neotype for the genus and elevated to a specific rank. Occurrence.-Ordovician-LowerDevonian. the fact Genus MUSCOCHITINAParis, 1981 Discussion.-Calpichitina has been selected despite that Wilson and Dolly (1964) consideredit as a junior synonym Type species.-Muscochitina muscosa Paris, 1981. Holotype of Hoegisphaera.The latter,however, is here restrictedto spe- in Paris, 1981, p. 269-270, pl. 30, fig. 17, Saint-Germain-sur- cies with a subsphericalchamber (see discussion below). The Ay section, upper part of LaHaye-du-PuitsFormation, early definitionincludes the two subgeneraCalpichitina (Cal- collectionsof present Lochkovian,western France, IGR 51761 (M40/1), and Calpichitina (Densichitina), erected by Paris France. pichitina) the Institutde Geologie de l'Universitede Rennes, (1981, p.127 and 131), but no longer maintained. Diagnosis.-Lagenochitinidae with an ovoid chamberand a meshlike ornamentation. Genus DESMOCHITINAEisenack, 1931 Occurrence.-Lower Devonian. Type species.-Desmochitina nodosa Eisenack, 1931. Lost Genus RAMOCHITINASommer and van Boekel, 1964 holotypein Eisenack,1931, p. 92, pl. 3, fig. 1, erraticOrdovician emend. Paris, Grahn,Nestor, and Lakova herein calcareoussandy siltstone;proposed neotype: Desmochitina no- Type species.-Ramochitina ramosi Sommer and van Boekel, dosa in Laufeld, 1967, p. 330-332, fig. 26 from the Keila Stage 1964. Holotypein Sommerand van Boekel, 1964, p. 426, pl. 1, (Caradoc),Fjaka section, Dalarna, Sweden, collections of the fig. 3, Tocantina,middle Devonian,Parana Basin, Goias, Brazil. Instituteof Palaeontologyof the Universityof Lund, Sweden. Representativespecies of the genus in Laufeld, 1974, figs. 49- Diagnosis.-Desmochitinidae with an ovoid, glabrouscham- 50. ber. Diagnosis.-Lagenochitinidae with an ovoid chamber and Occurrence.-Lower Ordovician-LowerDevonian. distinct crests (verticalrows of spines or membranes). Discussion.-The presentdefinition includes the two subgen- Occurrence.-Silurian-Devonian. era Desmochitina (Desmochitina) and Desmochitina (Pseudo- Discussion.-The verticalrows of spines (crests)occurring on desmochitina),erected by Paris (1981, p. 116-117), but no lon- the holotype and confirmedby SEM observationon material ger maintainedas the presenceof a mucronin the Desmochitin- from the type locality (Grahn, unpublished),necessitate the idae is not regarded here as a generic character.

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Genus HOEGISPHAERAStaplin, 1961 Diagnosis.-Desmochitinidae with an ovoid, glabrouscham- emend. Paris, Grahn,Nestor, and Lakova herein. ber, and short tubularcopula. Typespecies.-Hoegisphaera glabra Staplin, 1961. Holotype Occurrence.-Silurian-Lower Devonian. in Staplin, 1961, p. 419-420, pl. 50, fig. 5, Socony Vegreville Discussion.-Granules may be present on the wall. no. 1, Duvemay Member,Woodbend Formation, cuttings from m Late Devonian Al- Subfamily PTEROCHITININAEParis, 1981 1,067-1,097 (3,500-3,600 ft), (Frasnian), Genus ARMORICOCHITINA 1981 berta,Canada; collections of ImperialOil Limited,Calgary, Al- Paris, berta, Canada. Type species.-Linochitina? ceneratiensis Paris, 1976. Holo- Diagnosis.-Desmochitinidae with a spherical, glabrous type in Paris, 1976, p. 107, pl. 23, fig. 1, Saint-Ceneresection, chamber. upper part of the Saint-CenereFormation, Pragian, Western Occurrence.-Upper Devonian. France,IGR 50160 (N 48/4), collections of the Institutde Geo- Discussion.-The genus is restrictedhere to forms with a sub- logie de l'Universite de Rennes, France. sphericalchamber. Species with a lenticularchamber are trans- Diagnosis.-Desmochitinidae with an ovoid chamberand a ferred into Calpichitina.The emendationsproposed by Urban membranouscarina extending below the margin. (1972) and by Legault (1973b), based on the occurrenceof a Occurrence.-Middle Ordovician-Lower Devonian. membranousmatlike structure,are not adoptedas the primary Discussion.-Granules may be present on the wall. or secondaryorigin of such a structureis not yet demonstrated. Genus CINGULOCHITINAParis, 1981 Genus OLLACHITINAPoumot, 1968 Type species.-Desmochitina cingulata Eisenack, 1937. Lost Typespecies.-Ollachitina ingens Poumot, 1968. Holotypein holotype in Eisenack, 1937, p. 220, pl. 15, fig. 6 from erratic Poumot, 1968, p. 47-48, pl. 1, fig. 1, Gs-2 borehole,core sample graptoliticshale ("Graptolithengestein");neotype: Linochitina from 3,451.50 m, El Gassi, northernSahara, Algeria, unknown cingulata, in Eisenack, 1968, p. 170-171, pl. 29, fig. 29, from formation,Tremadoc, collections of the Service Palynoplancto- erraticgraptolitic shale, Ludlow; collections of the Geologisch- logique de la SNPA-Elf,France. PalaontologischenInstitute der Universitat,Tiibingen, Germany. Diagnosis.-Desmochitinidae with a glabrous, cylindrical Representativespecimens from outcropsin Laufeld, 1974, fig. chamber. 37 B and D, Valbyte 1, Slite Marls, Sheinwoodian,Wenlock, Occurrence.-Lower Ordovician. Gotland,Sweden. with a thin-walled, conical to MARGACHITININAEParis, 1981 Diagnosis.-Desmochitinidae Subfamily ovoid glabrouschamber bearing a complete membranouscarina Genus LINOCHITINAEisenack, 1968 a short tubular restrict. 1981 on the margin; copula is present. Paris, Occurrence.-Silurian-Lower Devonian. Type species.-Desmochitina erratica Eisenack, 1931. Lost holotype in Eisenack, 1931, p. 92, pl. 3, fig. 6, erraticgraptolitic Genus PSEUDOCLATHROCHITINACramer, 1966 rock of early Ludlow age ("Graptolithengestein");neotype in emend. Priewalder,1997 Eisenack, 1962, p. 307, pl. 17, fig. 11, erraticgraptolitic rock, carmenchui 1964. Ho- Wenlockto collections of the Type species.-Clathrochitina Cramer, early Ludlow, Geologisch-Palaon- in Cramer,1964, 346, 24, 18, La Vid de Gordon Instituteder lotype p. pl. fig. tologischen Universitat,Ttibingen, Germany. section, upper San Pedro Formation, Pridoli, north-western Diagnosis.-Desmochitinidae with a claviformto cylindrical Pseudoclathrochitina carmenchui in chamberand a hollow tubular Spain; neotype: Priewalder, copula. 1997, 78-81, 1, 2, 6, 5, 8, slide 1997/1/2 Occurrence.-Middle Ordovician-MiddleDevonian. p. pl. figs. pl. fig. (L. 39.3); collections of the Geological Survey of Austria,Vienna. Discussion.-As expressed by Paris (1981, p. 148), forms Desmochitinidaewith a reticulum ex- a carinaare excluded from this Diagnosis.-Conical bearing genus. tending in a perforatedcarina. Genus MARGACHITINAEisenack, 1968 Occurrence.-Silurian. Discussion.-The emendation Priewalder Eisenack, 1937. proposedby (1997, Type species.-Desmochitina margaritana is However, we consider the of the Lost holotype in Eisenack, 1937, p. 221, pl. 15, fig. 9, erratic p. 78) accepted. folding limestones of Silurian in meshworkaround the marginas a structureequivalent to a ca- bluish-graymarly early age; neotype rina. Eisenack, 1962, p. 306, pl. 17, fig. 13, erraticSilurian graptolitic rocks, collections of the Geologisch-PalaontologischenInstitute Genus PTEROCHITINAEisenack, 1955a der Universitat,Tiibingen, Germany. Representative specimens in Laufeld, 1974, fig. 62. Type species.-Bion perivelatum Eisenack, 1937. Lost holo- Diagnosis.-Desmochitinidae with a lenticularto spherical, type in Eisenack, 1937, p. 229-230, pl. 16, fig. 4, erraticBey- glabrouschamber and a solid peduncle linking the apex to the richia limestones ("Beyrichia-kalk"); neotype in Eisenack, operculumof the precedingvesicle. 1955a, p. 177, pl. 3, fig. 10, erraticBeyrichia limestones from Occurrence.-Silurian-Lower Devonian. the Baltic bottom?,Pridoli; collections of the SenckenbergMu- Discussion.-Collarette is absent. seum, Frankfurtam Main, Germany. Diagnosis.-Desmochitinidae with a lenticularto spherical, Genus URNOCHITINAParis, 1981 glabrouschamber with a membranouscarina. Type species.-Desmochitina urna Eisenack, 1934. Lost ho- Occurrence.-Middle Ordovician-Lower Devonian. lotype, in Eisenack, 1934, p. 69-70, pl. 5, fig. 7, late Silurian (Pridoli) limestones from Karlstejn,Bohemia, Czech Republic; Subfamily CUTICHITININAEAchab, Asselin and Soufiane, 1993 proposed neotype: Desmochitina urna in Paris, Laufeld and Genus CUTICHITINAAchab, Asselin and Soufiane, 1993 Chlupac, 1981, p. 15-16, pl. 1, fig. 1, bed 29, late Pridolilime- Type species.-Cutichitina legrandi Achab, Asselin and Sou- stones from Karlstejn,Bohemia, Czech Republic,IGR 51365(0 fiane, 1993. Holotype in Achab et al., 1993, p. 7, pl. 1, fig. 9, 35), collections of the Institut de Geologie de l'Universite de QD-1 borehole, core sample from 3,361 m, upper part of the Rennes, France. OuarglaSandstone Formation, Arenig, CentralSahara, Algeria,

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GSC 103 969 (J.55.3), collections of the Geological Survey of Genus ORBICHITINAAchab, Asselin and Soufiane, 1993 Canada,Ottawa, Ontario, Canada. Type species.-Orbichitina vulpina Achab, Asselin and Sou- Diagnosis.-Desmochitinidae with an ovoid chamberenvel- fiane, 1993. Holotype in Achab et al., 1993, p. 7, pl. 1, fig. 5, oped by a membranoussleeve. Riviere-au-Renard,Gaspe Peninsula, Quebec, Canada, lower Occurrence.-Ordovician. part of the Rosebush Cove Member of the Indian Point For- collections of Geo- EISENACKITININAE 1981 mation, Pridoli, GSC 103967 (E 44.3), the Subfamily Paris, of Genus EISENACKITINA 1964 logical Survey Canada,Ottawa, Ontario, Canada. Jansonius, with a conical chamberand a restrict.Paris, 1981 Diagnosis.-Desmochitinidae crown of processes. Type species.-Eisenackitina castor Jansonius, 1964. Holo- Occurrence.-Silurian (Pridoli). type in Jansonius, 1964, p. 912-913, pl. 2, fig. 16, Imperial Cartridgeborehole, core sample from 253 m (758 ft.), Hume Genus SALOPOCHITINASwire, 1990 Formation,early Givetian, northernCanada, collections of Im- emend Paris, Grahn,Nestor, and Lakova perial Oil Limited,Calgary, Alberta, Canada. Type species.-Conochitina? monterrosae Cramer, 1969. Ho- Emendeddiagnosis.-Desmochitinidae with an ovoid cham- lotype: Salopochitina bella Swire, 1990 (=S. monterrosae), p. ber and a randomlydistributed, spiny ornamentation. 109-110, pl. 2, fig.3, Lover Hill Farmborehole, Buildwas For- Occurrence.-Middle Ordovician-MiddleDevonian. mation,Sheinwoodian, Shropshire, England, MPK 5913, collec- Discussion.-In agreementwith Paris (1981, p. 155), smooth, tions of the British Geological Survey, Keyworth,England. ovoid forms are transferredinto Desmochitinaand smooth,con- Diagnosis.-Desmochitinidae with a conical to ovoid cham- ical ones into Bursachitina. ber, and sheathingprocesses. Occurrence.-Silurian (Wenlock-Ludlow). Genus KALOCHITINAJansonius, 1964 Discussion.-The chamber surface may be granulous and, Type species.-Kalochitina multispinata Jansonius, 1964. Ho- when damaged, the remainingprocesses may simulate a laci- lotype in Jansonius, 1964, p. 909-910, pl. 2, fig. 21, Imperial- niated carina.The originaldiagnosis of the genus includes mis- Calvan Anderson 9-6 borehole, cuttings from 736-737 m interpretations(e.g., centralprocess); in additionwe considerS. (2,420-2,425 ft), AndersonTownship, Essex County, Ontario, bella Swire, 1990 to be a junior synonym of S. monterrosae CanadaMaeford-Dundas Formation, Cincinnatian (late Caradoc- (Cramer,1969). Ashgill), collections of ImperialOil Limited, Calgary,Alberta, Canada. ACKNOWLEDGMENTS Diagnosis.-Desmochitinidae with a conical chamber and We are particularlygrateful to T. Winchester-Seeto,and to the randomspines. reviewers, M. A. Miller and S. J. E. Sutherland,for valuable Occurrence.-Upper Ordovician. suggestions and for a careful readingof the manuscript.Many colleagues from the ChitinozoanSubcommission of the CIMP Genus ORDOCHITINAAchab, Asselin and Soufiane, 1993 (CommissionInternationale de Microfloredu Paleozoique)made Type species.-Ordochitina tadlaiensis Achab, Asselin and helpful comments that improved the final version the manu- Soufiane, 1993. Holotype in Achab et al., 1993, p. 6, pl. 1, fig. script. We also thankthe differentmuseums that answeredour 1, Bj-103 borehole,cuttings sample from 849 m, unnamedfor- requestsconcerning the repositoryplaces of type material. mation,early Caradoc,Tadla Basin, Morocco, GSC 103 965 (P This is a contributionto the ChitinozoaClade Groupof IGCP 27.2), collections of the Geological Survey of Canada,Ottawa, Projectnumber 410. Ontario,Canada. REFERENCES Diagnosis.-Desmochitinidae with a conical chambercovered by crests (verticalrows of spiny or membranousornamentation). ACHAB,A. 1980. 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Amphorachitina, Ollachitina, Velatachitina, trois Geological Notes, 24(10):224-232. nouveaux genres de Chitinozoairesde 1'Ergoriental (Alg6rie-Tuni- -, AND R. W. HEDLUND.1964. Calpichitina scabiosa, a new Chiti- sie). Bulletin du Centrede Recherchede Pau, 2:45-55. nozoan from the Sylvan Shale (Ordovician) of Oklahoma. Oklahoma PRIEWALDER,H. 1997. SEM-revisionof a chitinozoanassemblage from Geological Notes, 24(7):161-164. the uppermostSan Pedro Formation(Pridoli), Cantabrian Mountains WOOD,G. D. 1994. Togachitina,a new bilayered chitinozoangenus (Spain). Jahrbuchder GeologischenBundesandstalt, 140:73-93. from the Devonian of the Sierra Subandinas Region, Bolivia. Paly- REID,P. C., ANDA. W. G. JOHN.1981. A possible relationshipbetween nology, 18:195-204. Chitinozoaand Tintinnids.Review of Palaeobotanyand Palynology, -, AND M. A. MILLER.1991. Distinctive Silurianchitinozoans from 34:251-262. the ItacurubaGroup (Vargas Pefia Shale), Chaco Basin, Paraguay. SCHALLREUTER,R. 1963. Neue Chitinozoen aus ordovizischen Geschie- Palynology, 15:181-192. ben und Bemerkungenzur GattungIllichitina. Palaeontologische Ab- WRONA, R. 1980. Upper Silurian-Lower Devonian Chitinozoa from the handlungen,1:392-405. subsurface of southeastern Poland. Palaeontologia Polonica, 41:103- . 1981. Chitinozoenaus dem Sularpschieffer(Mittelordoviz) von 165. Schonen (Schweden).Palaeontographica B, 178:89-142. ZASLAVSKAYA, N. M. 1980. SilurianChitinozoa from the SiberianPlat- . 1983. Sularpschiefer(Mittelordoviz) als Geschiebe in Nord- form. AkademiaNauk SSR, 435:52-80. (In Russian) deutschland. aus Dem In- Mitteilungen Geologisch-Palaontologischen ACCEPTED18 FEBRUARY1999 stitut UniversitatHambourg, 54:55-64. SCHWEINEBERG,J. 1987. Silurische Chitinozoen aus der Provinz Palen- cia (KantabrischesGebirge, N-Spanien). Gottinger Arbeiten zur Geo- APPENDIX: LIST OF THE REJECTEDGENERA logie und Palaontologie,33:1-122. SOMMER,F. W., AND N. M. VAN BOEKEL.1964. Quitinozoarios do De- Acmochitina Tsegelnyuk, 1982. Type species: Acmochitina corollata voniano de Goias. Anais Da Academia BrasileiraDe Ciencias, 36: Tsegelnyuk, 1982. Junior synonym of Ancyrochitina Eisenack, 423-431. 1955a. STAPLIN, F. L. 1961. Reef-controlleddistribution of Devonian micro- Agathochitina Tsegelnyuk, 1982. Type species: Ancyrochitina primitiva planktonin Alberta.Palaeontology, 4:392-424. Eisenack, 1955a. Junior synonym of Ancyrochitina Eisenack, 1955a. SUTHERLAND,S. J. E. 1994. Ludlow chitinozoansfrom the type area Aleurochitina Locquin, 1976. Type-species: Desmochitina erinacea Ei- and adjacentregions. PalaeontologicalSociety Monographs,London, senack, 1931. Nomen nudum. 148, 104 p. Amphorachitina Poumot, 1968. Type species: Amphorachitina conifun- SWIRE,P. H. 1990. New Chitinozoantaxa from the Lower Wenlock dus Poumot, 1968. Junior synonym of Lagenochitina Eisenack, 1931.

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AqualichitinaLocquin, 1976. Type species: Lagenochitinaovoidea Tau- FlascachitinaJansonius, 1967. Type species:Angochitina fiasca Collin- gourdeauand de Jekhowsky,1960. Nomen nudum. son and Schwalb, 1955. Nomen nudum(see Jansonius,1967, p. 351). AmpullachitinaCollinson and Schwalb, 1955. Type species: Ampul- FustichitinaAchab, 1980. Type species: Fustichitinaventriosa Achab, lachitina lagunculaCollinson and Schwalb, 1955. Remarks:Ampul- 1980. Junior synonym of ConochitinaEisenack, 1931, restrict.Ei- lachitinalaguncula Collinson and Schwalbwas publishedon 16 May senack, 1955b, 1965 and restrict.Paris, 1981. 1955. It is thereforea senior synonym of AncyrochitinaEisenack, GiraffachitinaLocquin, 1985. Type species: Giraffachitinapoignantii 1955a publishedon 5 June 1955. However,despite the fact that it is Locquin, 1985. We exclude this taxon from the chitinozoans. not a nomen oblitum, Ampulachitinahas not been used since 1955, GotlandochitinaLaufeld, 1974. Type species: Gotlandochitinamartins- thus for stability of nomenclature,it is suggested to maintainAncy- soni Laufeld, 1974. Juniorsynonym of RamochitinaSommer and van rochitinabecause Collinson himself adoptedthis genus (see Collin- Boekel, 1964 emend. son and Scott, 1958). GuizhouchitinaLai, 1982. Type species: Guizhouchitinalagena Lai, Ascochitina Tsegelnyuk, 1982. Type species: Ascochitina seccata Tse- 1982. Juniorsynonym of LagenochitinaEisenack, 1931 if the vesicle gelnyuk, 1982. Junior synonym of Angochitina Eisenack, 1931, is glabrous.Remarks: the "reticulate"pattern of the holotype is an emend. Eisenack, 1968. artifactrelated to pyrite crystal casts. Baculochitina Locquin, 1976. Type species: Cyathochitina cylindrica HalochitinaEisenack, 1968. Type species: Pterochitinaretracta Eisen- Taugourdeauand de Jekhowsky,1960. Nomen nudum. ack, 1955b. Juniorsynonym of PterochitinaEisenack, 1955a. Biconigutta Schallreuter, 1981. Type species: Biconigutta catinus HaplochitinaGrignani and Mantovani,1964. Type species: Haplochi- Schallreuter,1981. Consideredhere as a juniorsynonym of Armigutta tina omdoulensisGrignani and Mantovani,1964. Juniorsynonym of Schallreuter,1981. Remarks:this genus is based on a single specimen SphaerochitinaEisenack, 1955a. from an erraticrock sample. HelicochitinaLocquin, 1976. Type species: Helicochitinaviticula Loc- Calpichitina (Densichitina) Paris, 1981. Type species: Desmochitina quin, 1976. Nomen nudum. densa Eisenack, 1962. This subgenus, sometimes used at a generic IberichitinaJansonius, 1967. Type species: Clathrochitinacarmenchui rank (e.g., Grahnand Paris, 1992), is here includedwithin Calpichi- Cramer,1964. Nomen nudum. tina emend. Paris 1981. IdiochitinaTsegelnyuk, 1982. Type species: Idiochitinaplatycera Tse- Calychitina Tsegelnyuk, 1982. Type species: Desmochitina? urna Ei- gelnyuk, 1982. Juniorsynonym of AncyrochitinaEisenack, 1955a. senack, 1934. Juniorsynonym of UrnochitinaParis, 1981. Illichitina Collinson and Schwalb, 1955. Type species: Illichitina cro- Catenachitina Jansonius, 1967. Type species: Desmochitina erratica Ei- talum Collinson and Schwalb, 1955. Remarks:the authors of this senack, 1931. Nomen nudum,see Jansonius,1967, p. 351. genus laterconsidered it to be a synonymof CyathochitinaEisenack, Catenochitina Locquin, 1976. Type species: Desmochitina elegans Tau- 1955b (see Taugourdeauet al., 1967, p. 62). NormallyIllichitina has gourdeauand de Jekhowsky,1960. Nomen nudum. the anteriority(16 May 1955, instead of 12 December 1955 for Ceratochitina Tsegelnyuk, 1982. Type species: Ceratochitina cornuta Cyathochitina)but for stability of the taxonomy and because the Tsegelnyuk,1982. Juniorsynonym of AncyrochitinaEisenack, 1955a definitionwas too broad,we follow the decision of CIMP (see Tau- Cingulochitina Locquin, 1976. Type species: Cyathochitina dispar Tau- gourdeauet al., 1967, p. 62) and do not retainIllichitina. gourdeauand de Jekhowsky,1960. Nomen nudum. JansoniuchitinaLocquin, 1976. Type species: Kalochitina cf. inflata Cladochitina Lange, 1967. Type species: Conochitina biconstricta (Taugourdeau,1961) in Jansonius, 1964, pl. 2, fig. 23. Nomen nu- Lange, 1949. Juniorsynonym of SpinachitinaSchallreuter, 1963. dum. ClathrochitinellaCramer, 1967. Type species: Clathrochitinaoblonga JenkinochitinaParis, 1981. Type species: Conochitinaoelandica Eisen- Benoit and Taugourdeau,1961. Invalid taxon accordingto the code ack, 1955b. Juniorsynonym of EuconochitinaTaugourdeau, 1966. of Zoological Nomenclature(no definitiongiven). LabrochitinaLocquin, 1976. Type species: Conochitinaelegans Eisen- CombachitinaLocquin, 1976. Type species: Lagenochitinadubia Tau- ack, 1931. Nomen nudum. gourdeauand de Jekhowsky,1960. Nomen nudum. LachkarichitinaLocquin, 1976. Type species: Cyathochitinaelongata Coronochitina Eisenack, 1965. Type species: Coronochitina coronata Bouche, 1965. Nomen nudum. Eisenack, 1965. Juniorsynonym of SpinachitinaSchallreuter, 1963. LambdachitinaLakova, 1986. Type species: Lambdachitinacoronata CramerochitinaLocquin, 1976. Type species: Halochitinaretracta Ei- Lakova, 1986. Junior synonym of AngochitinaEisenack, 1931 re- senack, 1968. Nomen nudum. strict. Eisenack, 1968. Cupulachitina Jansonius, 1967. Type species: Desmochitina elegans LeiochitinaLocquin, 1976. Type species: Cyathochitinaelenitae Cra- Taugourdeauand de Jekhowsky,1960. Nomen nudum. mer, 1964. Nomen nudum. Cylindrochitina Schallreuter, 1963. Type species: Cylindrochitina gran- Leiochitina Tsegelnyuk, 1982. Type species: Lagenochitina elegans ata Schallreuter,1963. Junior synonym of Angochitina Eisenack, Beju and Danet, 1962. Juniorsynonym of LagenochitinaEisenack, 1931, restrict.,1968. 1931. Dasychitina Tsegelnyuk, 1982. Type species: Dasychitina implexa Tse- LigulachitinaLocquin, 1976. Type species: Rhabdochitinavirgata Tau- gelnyuk, 1982. Juniorsynonym of AncyrochitinaEisenack, 1955a. gourdeau,1961. Nomen nudum. Deflandrochitina Locquin, 1976. Type species: Angochitina milanensis LissochitinaTsegelnyuk, 1982. Type species: Sphaerochitinaresupina Collinson and Scott, 1958. Nomen nudum. (Dicevichius, 1971). Junior synonym of SphaerochitinaEisenack, Desmochitina (Pseudodesmochitina) Paris, 1981. Type species: Des- 1955a mochitina?cocca Eisenack, 1931. This subgenus,sometimes used at MacrostomachitinaTaugourdeau, 1965. Type species: Lagenochitina generic rank (e.g., Schallreuter,1983), is here included within Des- macrostomaTaugourdeau and de Jekhowsky, 1960. Nomen nudum; mochitinaEisenack, 1931, emend. Paris, 1981. see Jansonius(1967, p. 352). DiabolochitinaLocquin, 1976. Type species: Conochitinasymmetrica MirachitinaEisenack, 1931. Type species:Mirachitina quadrupedis Ei- Taugourdeauand de Jekhowsky,1960. Nomen nudum. senack, 1931. No longer includedin the chitinozoans. Discochitina Tsegelnyuk, 1982. Type species: Discochitina discoides MonicachitinaLocquin, 1976. Type species: Cyathochitinastriata Ei- Tsegelnyuk, 1982. Junior synonym of Ancyrochitina Eisenack, senack, 1937. Nomen nudum. 1955a. Monilichitina Locquin, 1976. Type species: Desmochitina sommeri EarlachitinaCollinson and Scott, 1958. Type species: Earlachitinala- Lange, 1952. Nomen nudum. tipes Collinson and Scott, 1958. Junior synonym of Ancyrochitina Mucrochitina Locquin, 1976. Type species: Lagenochitina esthonica Eisenack, 1955a. Eisenack, 1955b. Nomen nudum. Edouardochitina Locquin, 1976. Type species: Angochitina communis Mycetochitina Locquin, 1976. Type species: Mycetochitina hypha Loc- Jenkins, 1967. Nomen nudum. quin, 1976. Nomen nudum. ErinaceochitinaLocquin, 1976. Type species: Sphaerochitinacollinsoni NanochitinaNestor, 1994. Type species Nanochitinanana Nestor,1994. Dunn, 1959. Nomen nudum. Regardedas a junior synonym of BursachitinaTaugourdeau, 1966. EurychitinaTsegelnyuk, 1982. Type species: Bursachitinaoviformis Ei- Nematochitina Locquin, 1976. Type species: Ancyrochitina longicornis senack, 1972. Junior synonym of Bursachitina Taugourdeau, 1966. Taugourdeauand Jekhowsky,1960. Nomen nudum.

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Oochitina Tsegelnyuk, 1982. Type species: Oochitinafugax Tsegel- StriatolagenochitinaSchallreuter, 1981. Type species: Striatolageno- nyuk, 1982. Juniorsynonym of AncyrochitinaEisenack, 1955a. chitina clava Schallreuter,1981. Junior synonym of Lagenochitina OrochitinaTsegelnyuk, 1982. Type species: Orochitinainflata Tsegel- Eisenack, 1931. nyuk, 1982. Juniorsynonym of AngochitinaEisenack, 1931 restrict SubulochitinaLocquin, 1976. Type species: Acanthochitinasecunda 1968. Schallreuter,1963. Nomen nudum. Pallachitina Costa, 1970. Type species: Pallachitina wilhelmi Costa, TaugourdochitinaLocquin, 1976. Type species: Conochitinamicracan- 1970. Juniorsynonym of BursachitinaTaugourdeau, 1966. tha Eisenack, 1931. Nomen nudum. Palenchitina Schweineberg, 1987. Type species: Palenchitinapisuer- Togachitina Wood, 1994. Type species: Togachitina eamesi Wood, gensis Schweineberg,1987. Juniorsynonym of AncyrochitinaEisen- 1994. Juniorsynonym of Pellichitina Achab, Asselin, and Soufiane, ack, 1955a. 1993 ParachitinaEisenack, 1937. Type species: Parachitinacurvata Eisen- TrochochitinaMiller, 1976. Type species Trochochitinaradiata Miller ack, 1937. Incertaesedis (see Jansonius,1967, 1970). No longer in- 1976. Nomen nudum. cluded in the chitinozoans. UrnichitinaLocquin, 1976. Type species:Desmochitina rhenana Eisen- ParvichitinaLocquin, 1976. Type species: Conochitinavasculiformis ack, 1939. Nomen nudum. Bouch6, 1965. Nomen nudum. VirgilochitinaTasch and Hutter,1978. Type species: Virgilochitinakan- PhiolachitinaMontenari and Maass, 1996. Type species: Phiolachitina sasensis Tasch and Hutter,1978. Juniorsynonym of Lagenochitina silvanegraticaMontenari and Maass, 1996. Not includedin the chi- Eisenack, 1931. tinozoans. VitreachitinaNestor, 1994. Type species: Sphaerochitinavitrea Tau- PoteriochitinaLegault, 1973a. Type species: Poteriochitinabriarca Le- gourdeau, 1962. Regarded as a junior synonym of Euconochitina gault, 1973a. Juniorsynonym of EisenackitinaJansonius, 1964. Taugourdeau,1966 because the wall thicknessis not used as a dis- PoumochitinaLocquin, 1976. Type species:Ancyrochitina multiradiata criminatecharacter in the presentclassification. Eisenack, 1959. Nomen nudum. RetrochitinaLocquin, 1976. Type species: Halochitinaretracta Eisen- Discussion.-Locquin's document(1976), includingnew chitinozoan ack, 1955b. Nomen nudum. genera and distributedto some chitinozoanworkers by this author,is RhizochitinaTsegelnyuk, 1982. Type species:Rhizochitina laufeldi Tse- not registeredin the official recordfor Frenchpublications in the Bib- gelnyuk, 1982. Juniorsynonym of AncyrochitinaEisenack, 1955a. liotheque Nationale de France. Concerningvalid publications,the In- SclerochitinaTsegelnyuk, 1982. Type species: Conochitinaintermedia ternationalCode of Zoological Nomenclature(third edition, 1985) stip- Eisenack, 1955a. Juniorsynonym of BelonechitinaJansonius, 1964. ulates: Article 8a(i) "it must be issued for the purposeof providinga SolenochitinaTsegelnyuk, 1982. Type species: Solenochitinafistulata permanentscientific record." We consider that this requirementand Tsegelnyuk, 1982. Juniorsynonym of ConochitinaEisenack, 1931, point (i) of Article 13a of the code are not fulfilled in Locquin'sdoc- emend. Paris, 1981. ument (1976). Therefore,we consider the taxa describedin this docu- SpathachitinaCosta, 1970. Type species: Spathachitinacruzi Costa, ment as nominanuda, like the taxa describedin unpublishedtheses. 1970. Juniorsynonym of ConochitinaEisenack, 1931, emend.Paris, Four new genera, Grahnichitina,Nanochitina, Retiachitina, and Ser- 1981 icochitina, have been erected recently by Geng et al. (1997). Because SteneyochitinaZaslavskaya, 1980. Type species: Steneyochitinaovatoe- they were publishedafter the submissionof our manuscript,it was not longata Zaslavkaya, 1980. Junior synonym of SpinachitinaSchall- possible to include them in our multivariateanalysis and therefore,they reuter,1963. are not discussed herein. It should be noted that NanochitinaGeng et StephanochitinaGrignani and Mantovani,1964. Type species: Stephan- al., 1997 is an homonymof NanochitinaNestor, 1994. However,if the ochitina africana Grignaniand Mantovani,1964. Probablya junior synonymyof NanochitinaNestor, 1994 with BursachitinaTaugourdeau, synonym of AngochitinaEisenack, 1931, but the diagnoses and il- 1966 is accepted, then the name Nanochitinacan be used again as a lustrationof the type materialdo not allow a definite statement. generic name.