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J. Paleont., 80(2), 2006, pp. 264–271 Copyright 2006, The Paleontological Society 0022-3360/06/0080-264$03.

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SHELL HETEROSTROPHY IN EARLY ORDOVICIAN MACLURITELLA KIRK, 1927 AND ITS IMPLICATIONS FOR PHYLOGENY AND CLASSIFICATION OF MACLURITOIDEA (GASTROPODA)
ˇ´ ´ JIRI FRYDA
AND

DAVID M. ROHR

Czech Geological Survey, Klarov 3, 118 21 Praha 1, Czech Republic, fryda@cgu.cz and Department of Geology, Sul Ross State University, ´ Alpine, Texas 79832, drohr@sulross.edu

ABSTRACT—Study of the oldest macluritid gastropod, Macluritella stantoni Kirk, 1927 from the Lower Ordovician of Colorado, has revealed that its early whorls are openly and dextrally coiled, in contrast to those in later teleoconchs which are sinistrally coiled. This is the first documentation of heterostrophic coiling in members of the Macluritoidea, which have been considered to be dextrally hyperstrophic. Juvenile M. stantoni may be interpreted as dextrally orthostrophic and, thus, it had the same type of soft-body-shell arrangement as the vast majority of living and fossil gastropods. This intepretation also suggests that the Macluritoidea evolved from the dextrally orthostrophic gastropods, and their dextral hyperstrophy is derived and not a primary feature. In addition, occurrence of shell heterostrophy in M. stantoni brings additional evidence that the Macluritoidea and Onychochiloidea are not closely related taxa. Relationships between the Macluritoidea and Euomphaloidea are still uncertain. This study provides the oldest evidence (Early Ordovician) for shell heterostrophy in the class Gastropoda.

INTRODUCTION

majority of gastropod shells are asymmetrically (anisostrophically) coiled and most of them are right-handed (dextral). Although left-handed (sinistral) shells seem to be less common, they are known to occur in several unrelated gastropod groups during more than 500 My of gastropod evolution (Wenz, 1938; Knight et al., 1960; Pchelintsev and Korobkov, 1960; Vermeij, 1975). Symmetrical shells have been developed only in a few gastropod groups and they are either uncoiled (patelliform) or symmetrically coiled (isostrophic) like those in Paleozoic Bellerophontoidea (Amphigastropoda) and some Porcellioidea (Archaeogastropoda). Symmetrically coiled shells in the first group have been considered to represent a primitive stage (e.g., Knight, 1952). However, bilateral shell geometry in the Porcellioidea is a derived shell character (Fryda, 1997). Similarly, the development ´ of uncoiled patelliform shells in different post-Paleozoic gastropod groups is also a derived shell feature. This is most probably true also for the Patelligastropoda ( Docoglossa), which are considered to be the oldest gastropod offshoot (see discussion in Ponder and Lindberg, 1997). Regardless of the manner of shell coiling, the gastropod viscera are torted relative to the head-foot. This torsion of the soft body is one of the most typical (apomorphic) and thus also diagnostic characters for the class Gastropoda. The soft-body torsion is independent of shell coiling (Bandel, 1982). Dextrality and sinistrality of the body torsion is determined very early in the embryonic development, and once determined, is never reversed (Robertson, 1993). Thus shell-less gastropods or those with bilaterally symmetrical shells have had torted viscera. There are four possible combinations of soft-body organization and shell coiling: anatomically dextral as well as sinistral animals may have dextrally or sinistrally coiled shells. These four types of geometrical arrangement in shell-bearing gastropods (Fig. 1) were recognized more than 100 yr ago (Pelseneer, 1893) and may be described in terms discussed in detail by Knight (1952). Anatomically dextral animals with dextrally coiled shells are termed dextral orthostrophic (Fig. 1.1). The mirror image of that (i.e., an anatomically sinistral animal with a sinistrally coiled shell) is sinistral orthostrophic (Fig. 1.2). However, an anatomically dextral animal with a sinistrally coiled shell is called dextral hyperstrophic, and the mirror image of that is sinistral hyperstrophic (Fig. 1.3, 1.4). These terms are, of course, applicable only to the shell-bearing gastropods. The vast majority of gastropods

T

HE VAST

are dextrally orthostrophic, followed by sinistrally orthostrophic. As summarized by Robertson (1993), dextrally hyperstrophic as well as sinistrally hyperstrophic gastropods are known among modern gastropods. In fossil gastropods the manner of coiling of the soft body can be inferred only indirectly from the coiling of the operculum (if present) or, in some cases, from the apertural shape (Knight, 1952; Robertson, 1963, 1993; Peel and Horny, ´ 1996). The dextral orthostrophic gastropods have sinistrally (counterclockwise) coiled opercula or vice versa (Fig. 1.1, 1.2). Similarly, opercula in dextral hyperstrophic gastropods are coiled sinistrally (counterclockwise) and vice versa for sinistral hyperstrophic gastropods (Fig. 1.3, 1.4). However, in some gastropods the shell coiling may be even more complex. There are several gastropod groups in which the shell whorls coil in one direction during a part of their life and another direction for the other part. The term heterostrophy (Fig. 1.6) has been used for this condition (e.g., Knight, 1941, 1952). If all shell whorls are coiled in the same direction (i.e., dextral or sinistral) the shell is termed homeostrophic (Fig. 1.5). Some authors have used the term heterostrophy in a narrower sense, for a condition of the protoconch when its whorls appear to be coiled in the opposite direction to those of the teleoconch (Knight et al., 1960). The term larval hyperstrophy is also used for the latter condition (Robertson, 1993). A more detailed description and different usage of the term heterostrophy may be found in Dzik (1983) and Hadfield and Strathmann (1990). The Ordovician Macluritoidea have been interpreted as dextrally hyperstrophic on the basis of their sinistrally coiled teleoconchs and opercula (e.g., Knight, 1952; Knight et al., 1960). In this paper, we describe an occurrence of shell heterostrophy in the Early Ordovician macluritoidean gastropod Macluritella Kirk, 1927 and discuss its implications for the classification and phylogeny of the macluritoidean gastropods.
AGE AND GEOLOGICAL SETTING

Macluritella was established by Kirk (1927) based on 12 silicified specimens collected by T. W. Stanton from the Manitou Limestone in Williams Canyon, near Manitou Springs, Colorado. The original illustrations appear to have been retouched [compared to Knight’s (1941, pl. 66, fig. 2a–d) illustrations] in order to emphasize certain features of the shell. In addition, Yochelson and Stinchcomb (1987, p. 56) note that ‘‘the line drawing provided by Knight et al. (1960, fig. 105–2) does not accurately portray the features of the species.’’ For the present study, silicified gastropods were collected from Kirk’s (1927) type locality

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1861. In this region.. only a few poorly known species were placed into this genus. The gastropods here are larger than those that occur at Williams Canyon. 1941. 6). Omphalocirridae. 1861. placed Macluritella back in the Macluritoidea.? walcotti from the genus Macluritella because of its distinctly triangular shape and shallow sinus. 184. 1981. p. the coiling is termed heterostrophic (6). Gastropods are from locality R01–03D in the Ptarmigan Chert Member of the Manitou Formation (Myrow et al. 1958). 1861 Genus MACLURITELLA Kirk. According to these workers. 1992) from the Silurian (Llandoverian) of the Siberian Platform is poorly preserved.Macluritoidea are a well-known group of Ordovician gastropods (Knight. 1994.g. the entire Manitou is Early Ordovician in age (Rossodus manitouensis Zone) (Myrow et al. Knight et al. 2a–d. and the gastropods seem to be concentrated locally. Class GASTROPODA Cuvier. 1992. The depositional and stratigraphic setting of the Lower Ordovician Manitou Limestone has been reviewed by Rigby and Myrow (1999) and Myrow et al. Dzik. Yochelson. pl. 1–12. 1983. 2003). Yochelson and Stinchcomb (1987) interpreted the type lot of M. Wagner. 1861 were changed several times (Linsley and Kier. This corresponds to Rigby and Myrow’s (1999) sponge locality. figs. Rohr and Gubanov. Conversely. Wenz (1938) placed the family Macluritidae together with the Euomphalidae. Macluritella is generally considered to be the oldest representative of the family (Knight et al. Sando (1957) established Macluritella marylandicus from the Lower Ordovician Rockdale Run Formation [Rossodus manitouensis through Oepikodus communis conodont zones of Harris and Harris (1978)] in Maryland. Yochelson. 4304961N. Rohr. Rohr et al. 4) means that handedness of the shell and soft body is different. along the old road to the visitor center at Cave of the Winds.. 1999). 1984). 1861 Discussion. Orthostrophy (1. fig. Platyacridae. and they extended the range of Macluritella down from the Lower Ordovician into the Upper Cambrian with the addition of Macluritella? walcotti (Howell. 4302191N. stantoni to be rounded. who interpreted the Euomphaloidea to have been derived from the Macluritoidea during Early Ordovician time. but does not appear to belong to the genus because it lacks diagnostic features.´ FRYDA AND ROHR—SHELL HETEROSTROPHY IN EARLY ORDOVICIAN MACLURITELLA 265 FIGURE 1—Schematic diagrams showing four possibilities of the relationship between shell coiling and body asymmetry in shell-bearing gastropods (1–4) and the relationship between the coiling of larval (protoconch II) and postlarval (teleoconch) shells (5. The reddish cherty. opinions on the generic composition of the Macluritoidea Carpenter. Heterobranchia). Thus. 1797 Subclass UNCERTAIN Superfamily MACLURITOIDEA Carpenter. hyperstrophy (3. and this concept was followed by Knight et al. 1931. 2004). juvenile forms that during later ontogeny would evolve whorls with a triangular profile. 1999). The shell identified as Macluritella by Gubanov (in Sokolov. 2002). Rohr. Coiling of all shell whorls in the same direction (e. little is known ´ about their phylogenetic position. 1994. However. (1960) placed two subgenera under Macluritella: Macluritella and Euomphalopsis Ulrich and Bridge. 508877E. 2002). 1952. Family MACLURITIDAE Carpenter. in the suborder Macluritina of the Archaeogastropoda (Knight et al.Macluritella stantoni Kirk. 1960. a species with an acutely triangular profile and unknown early whorls. 288. 1927 Type species. The beds at this locality are within Trilobite Zone F (Loche in Rigby and Myrow. 2) means that anatomically dextral (or sinistral) animals occupy dextrally (or sinistrally) coiled shells. Wagner. The type locality corresponds to our locality R01–01 and is in the Middle Cherty Member of the Manitou Formation..Beside the type species.. together with the Euomphaloidea de Koninck. Yochelson and Stinchcomb (1987) restored Euomphalopsis to full generic status. Knight. in Williams Canyon as well as from the Ramparts Road section about 4 km north-northeast of the Williams Canyon section. Macluritella may also be represented by internal molds from the Lower Oslobreen Limestones (Lower Ordovician) in Spitsbergen (Hallam. 66. Macluritella stantoni. 1984. If handedness of some whorls is opposite (e. NAD 27. This concept was followed by Yochelson (1956). 1997. SYSTEMATIC PALEONTOLOGY not be linked together in a common taxon. and Poleumitidae within the superfamily Euomphaloidea ( Euomphalacea). Macluritella has no certain Cambrian record. Fryda and Rohr. p.. The second locality contains sponges. Similarly.. (2003). 1984. Discussion. Later Cox and Knight (1960) placed the Macluritoidea Carpenter. Linsley and Kier. 1881. Caenogastropoda and Neritimorpha) is termed homeostrophic (5). Wagner (2002). 1927. the Macluritoidea and Euomphaloidea should . 1960). dolomitic limestone is exposed on the Ramparts Forest Service road north of the Garden of the Gods at UTM zone 13. Later Yochelson and Stinchcomb (1987) moved the genus to the Euomphaloidea de Koninck.g. 1881. Oriostomatidae. UTM Zone 13. We exclude M.. Their concept of the Macluritina was later criticized by several authors (Morris and Cleevely. 1979. Included species. NAD 27. 506860E. Cirridae. based on a phylogenetic analysis of numerous teleoconch features. 1984.Wenz (1938) placed Macluritella in the family Macluritidae Carpenter. 1946). (1960).

Shell heterostrophy has been considered by many neontologists to be limited to the gastropod subclass Heterobranchia. This represents the first evidence of heterostrophic coiling in the Macluritoidea. p. Museum of Natural History. As noted by Knight (1952).Discoidal shells with rounded. The present study of Macluritella stantoni has yielded evidence for heterostrophic coiling of its shell. 57. 2. 9. This interpretation of the oldest macluritid genus suggests that the Macluritoidea evolved from dextrally orthostrophic gastropods (Haeckel’s biogenetic law). 289. and their dextral hyperstrophy is a derived rather than a primary shell feature. 2. Hadfield and Strathmann (1990). EVIDENCE OF HYPERSTROPHY IN MACLURITOIDEA The Ordovician genus Maclurites LeSueur. Knight et al. 8. 2. At least the first two shell whorls in Macluritella stantoni are distinctly dextrally coiled (Fig. 1990.Macluritella stantoni was described from a horizon about 125 ft below the top of the Manitou Limestone (‘‘Tremadocian. 2.8–2. KNIGHT. 184. In heterobranch gastropods with a lecitotrophic or direct development (some marine. 2. diameter of first preserved whorl about 0. width of teleoconch about double its height.7).8–2. 25. the Macluritoidea had the same type of soft-body-shell arrangement (anatomically dextral body in a dextrally coiled shell) as the vast majority of living and fossil gastropods. This type of operculum had a high probability of being fossilized. However. In these gastropods the timing of the change of the shell coiling (from sinistral to dextral or vice versa) is always connected with an ontogenetic stage when a true larval shell (protoconch II) is completed and the teleoconch starts to form. apertural.4. 2. YOCHELSON AND STINCHCOMB.7). 1952.S. oblique apical. diameter of subcircular aperture about one-third of maximum shell diameter (Fig.. 1990. 1996). internally tapering aperture on some specimens suggests an operculum may have been retracted a short distance inside shell. The remaining five syntypes represent paratypes. lateral. ´ The Macluritoidea are one of the oldest gastropod groups and. Specimen numbers are those of the U. NO. 12. Emended description. 2. According to Yochelson. near Manitou Springs. Colorado. whorl profile roughly subcircular but somewhat flattened at upper whorl surface (Fig. shell wall moderately thick. 1990. apical. and oblique lateral views of the same specimen showing distinct dextral coiling in early whorls. Colorado. apical. The latter type of shell heterostrophy is considered to be an apomorphy of the Heterobranchia and it occurs in all heterobranchs where planktotrophic larvae have been developed.4. they have been interpreted as dextrally hyperstrophic.1–2. apical. nearly flat upper shell surface bearing central depression formed by umbilicus of dextrally coiled initial part. 1987. Washington. umbilical and lateral whorl surfaces separated by rounded keel (Fig. Occurrence. sinistrally coiled whorls in adult shells. DC. 2. The majority of marine. WAGNER.15). Dzik (1983). all 5. pl. 1960. Detailed discussion on dextral hyperstrophy in the Macluritoidea may be found in several papers (Knight.Knight (1941) designated the specimen illustrated by Kirk. 18. 14. this shell feature was later also documented in some archaeogastropods (Hadfield and Strathmann.266 JOURNAL OF PALEONTOLOGY. oblique apical. irregular transverse lirae (Fig. 35. Nevertheless. 14. thick. 2.’’ Lower Ordovician). The spiral operculum of Maclurites is sinistrally (counterclockwise) coiled. 2). Linsley and Kier. This interpretation Heterostrophy is defined as the condition where the protoconch and/or early whorls of the teleoconch appear to be coiled in an opposite direction to the later teleoconch whorls. oblique. 15. shell-bearing Heterobranchia has developed heterostrophically coiled shells. and Robertson (1993). 2006 MACLURITELLA STANTONI Kirk. Yochelson. this feature was interpreted by Woodward (1854) as evidence for dextral hyperstrophic coiling. freshwater. in Williams Canyon. 66.7). detail of figure 8. 15. Fryda and Blodgett. and oblique basal views of specimen USNM 528374 from locality R01-01. The distribution of heterostrophy within the class Gastropoda is discussed in the following paragraphs. 13. V. and terrestrial Heterobranchia). National Museum. the documentation of sinistral heterostrophy in the oldest macluritid genus Macluritella (see below) has significant implications for interpretation of shell ontogeny in the Macluritoidea. 2. and in some specimens it has been found in situ within the aperture. 1984.1. figs. near Manitou Springs. dextrally coiled juvenile specimen USNM 528375 from locality R01-01. Macluritella stantoni has been reported from the upper part of the Roubidoux Formation in Missouri (Heller. 1984. p. fig.15). p. initial part of shell not well preserved. 10–12. 15. but no opercula were found with shells. 80. 1927 as figures 10–12 on page 289 as the holotype (one of six original syntypes under No. counterclockwise curvature of the exterior surface of the operculum in Maclurites supports its interpretation as dextrally hyperstrophic (Yochelson. → FIGURE 2—Macluritella stantoni Kirk.8–2. all whorls openly coiled. first preserved (probably juvenile teleoconch) whorl openly coiled. protoconch/ teleoconch transition not preserved due to silicification of shell. 8. 2.10–12.3). Peel and Horny. Williams Canyon area. at this ontogenetic stage. ornamentation consisting of rather widely spaced. 2. 15. 1996). Macluritella can be interpreted as dextrally orthostrophic during its early ontogeny when its shell was dextrally coiled (Fig. calcified operculum. Oblique lateral. 11. gap separating first and second whorls much wider than that in other whorls (Fig. 1927 from the Manitou Limestone.6. 2. Aperture tangential with no reentrants. (1960). 1927 Figure 2 is based on the polygonal aperture of Maclurites. but at least two first whorls distinctly dextrally coiled (Fig. 10. Washington. 1954). 1998. juvenile whorls distinctly dextrally coiled (Fig. closely spaced growth lamella developed near aperture of larger specimens indicates adult stage. 2. in contrast to vast majority of fossil and living gastropods. Thus.4. ´ 2001). 2a–d. National Museum.15). 2. During later ontogeny the shell whorls become planispirally and subsequently sinistrally coiled (Fig. oblique apertural. slightly concave.15). . 2. p.8–2. detail of dextrally coiled early whorls in specimen USNM 528376 from locality R01-03. 2002. and thus became dextrally hyperstrophic. 1817 belongs to a limited group of Paleozoic gastropods which developed a thick.5 mm. its diameter wider than that of aperture (Fig. 20. This fact has significant implications for evaluation of their phylogeny and relationships to other gastropod groups (see below). 1–7. 25. 1927. DC).S. HETEROSTROPHIC COILING IN MACLURITELLA AND WITHIN THE CLASS GASTROPODA Macluritella stantoni KIRK. 2. Peel and Horny.4. A more detailed description and different usage of the term heterostrophy may be found in Knight et al.6). In addition to the type area. 12.1–2. Material examined. umbilicus of sinistrally coiled teleoconch wide. 71710 in the U. 1941. During later ontogeny the shell of Macluritella changed the coiling of its shell to sinistral. 14. Thus.6). the macluritid opercula grew by accretion around the entire margin and thus not just at a narrow growth zone as in rounded spiral opercula. Yochelson (1990) ´ suggested that the operculum in Maclurites grew differently than spiral opercula in more advanced gastropods.

´ FRYDA AND ROHR—SHELL HETEROSTROPHY IN EARLY ORDOVICIAN MACLURITELLA 267 .

Nutzel ¨ (2002) confirmed earlier observations (Yoo. McLean (1981) united members of the modern Neomphaloidea McLean. the fossil record of shell-bearing heterobranchs is relatively rich in Tertiary and Mesozoic strata. and Heterobranchia). Fryda et al. 1956 (Porcelliidae Koken in Zittel. They interpreted their findings as evidence for heterostrophic shell coiling. 1925. 2002. in contrast to Wenz (1938).. The Givetian (late Middle Devonian) genus Heteroloxonema Fryda. Bandel and Fryda (1998) consid´ ered euomphaloideans to represent an independent. the vast majority of which belong to the superfamily Streptacidoidea Knight. 1941. 1984). Bandel and Heidelberger. Knight et al. 1984). In addition. The phylogenetic relationships of the Devonian taxa within the Heterobranchia are still uncertain. Later Bandel and Fryda ´ (1998) showed that an unusual protoconch morphology of Devonian and Carboniferous euomphaloidean genera distinguishes them from members of extant gastropod groups (Patellogastropoda. 2002). 1966. 1999. 2) both groups are not closely related and belong to the Archaeogastropoda (Yochelson. 2004) discussed the origin of ´ shell heterostrophy in the Agnesiinae and concluded that the development of shell heterostrophy in the Archaeogastropoda and Heterobranchia is not homologous. Modern Heterobranchia represent a very successful gastropod group which unites many thousands of extant species. 1925 to be closely related to macluritid gastropods. analysis of the shell morphologies in later-discovered clisospirid and onychochilid taxa suggests that both the families Clisospiridae and Onychochilidae are closely related (Horny. Peel. Allogastropoda Haszprunar. Bandel. 1984. 2001. Fryda. Cocculiniformia. Linsley and Kier (1984) proposed to unite the Onychochiloidea . although an integration of ´ the Macluritoidea and Euomphaloidea into one higher taxon (e. Knight et al. Batten. Taken together. several additional Paleozoic groups of Heterobranchia with heterostrophic shells have been documented (Fryda and ´ Blodgett. recent ¨ analysis of the biodiversity of Ordovician gastropods (Fryda and ´ Rohr. McLean (1981) ¨ suggested that the members of the superfamilies Macluritoidea and Onychochiloidea do not belong to his suborder Euomphalina but represent a separate lineage. 1994. 1994. 1895) and the Mesozoic Cirridae Cossmann. 2. separated both the Clisospiridae and the Onychochilidae from the suborder Macluritina and established a new suborder Mimospirina for them. 2004. or if this character originated independently in these two archaeogastropod groups. Yoo. however. 2002. Nutzel. Discovery of shell heterostrophy in Macluritella stantoni represents the first evidence for the occurrence of this remarkable shell feature in the Macluritoidea. 2004) has shown a similarity in the diversity and turnover patterns of the Ordovician Euomphaloidea and Macluritoidea. in Paleozoic time there ¨ were at least three distinct groups belonging to the subclass Heterobranchia Gray. but distinct sinistral coiling in the first whorl of three members of extant Trochoidea (Vetigastropoda. This concept was subsequently followed by Knight et al. 1931 (?Middle Devonian. Fryda and Blodgett (1998) showed that ´ both types of shell heterostrophy (anastrophic as well as inclined heterostrophic coiling) were present in the Archaeogastropoda. (1960) and many other paleontologists. Golikov and Starobogatov. Hei´ delberger and Bandel. Unfortunately. V. 1984). 1985). Batten. 1986). On the other hand. Neritimorpha. 2006 Lesueurilla-like forms of the pleurotomarioidean stock. Consequently. 1996. Recently. 2002). 1952. the morphology of the openly coiled early whorls in Macluritella stantoni (Fig. 1840: the Streptacidoidea Knight. 2002). 1998. 1931. Nutzel et al. Hadfield and Strathmann (1990) described a slight. 2000 represents probably ´ the oldest-known streptacidoidean genus. 1998) on the nature of the protoconch and the shape of ´ the boundary between the protoconch and teleoconch. both groups represent good examples of the occurrence of sinistral heterostrophy in fossil archaeogastropods. Fryda and Blodgett. multiwhorled shells bearing a sinistral protoconch were relatively common in Carboniferous faunal communities (Donald. Archaeogastropoda) with dextrally coiled teleoconchs. 1979.10–2. Stuoraxidae Bandel. In summary. 2001 (Devonian). Thus. In addition. recent models of gastropod evolution have offered the following possibilities for the relationships of the Macluritoidea and Euomphaloidea: 1) both groups are closely related and belong to the Archaeogastropoda (Knight. Linsley and Kier.. At present it is difficult to solve the question of whether sinistral heterostrophy is a character which is apomorphic for the Porcellioidea (thus. Fryda ´ ´ ´ ´ and Farrell. (1960). Caenogastropoda. 1916. 1966. RELATIONSHIPS OF MACLURITOIDEA TO ONYCHOCHILOIDEA this shell feature occurs rarely because the morphology of the early shell is simplified. our knowledge of early members of the Heterobranchia. absence of knowledge on the protoconch morphology in the Macluritoidea prevents more detailed analysis of relationships between the Macluritoidea and Euomphaloidea. He also noted that the protoconch of some euomphalids is not fundamentally different from that of Docoglossa. (1960) considered members of the Onychochilidae Koken. or 3) both groups are closely related but they are not gastropods (Linsley and Kier. Significantly... ¨ 1979. 1960. 1850). 2001. 1997. However. Bandel and Fryda. This approach was followed by several authors (Wangberg-Eriksson. In contrast to the Heterobranchia. 1990. is still very poor.. 1985. 1898. Anderson et al. 1960. These gastropods which have slender. it also represents the oldest evidence (Early Ordovician) for shell heterostrophy in the class Gastropoda. 2. NO. Yochelson. uniting the Mesozoic Cirridae and the Paleozoic Porcelliidae into one natural group). Knight et al. 1931. Wagner. Heterobranchia have been well documented from late Paleozoic strata. These taxa belong to the Paleozoic Agnesiinae Knight. Later Fryda and Blodgett (2001. Nevertheless. 1981. 2005). which undoubtedly evolved in the Paleozoic. The latter taxon was interpreted as belonging to the Archaeogastropoda. Dzik (1983).268 JOURNAL OF PALEONTOLOGY. 1964. 2002. The protoconch morphology of the Macluritoidea is still unknown. 1981 with the Paleozoic Euomphaloidea and placed them in the Archaeogastropoda. Knight. However. This fact may suggest similarities in some of their life strategies (Fryda and Rohr. Also. Bandel and Fryda. 80. Archaeogastropoda.. Clisospirids were interpreted as sinistral orthostrophic gastropods with an uncertain higher taxonomic position. exclusively Paleozoic (Cambrian–Permian) gastropod group. Macluritina) was criticized by several authors (Morris and Cleevely. 1996 (Late Permian. considered onychochylids to be dextral hyperstrophic gastropods like the macluritids. 1998. Bandel. Heterostrophy also has been found in some fossil archaeogastropods (Knight. highspired. the development of shell heterostrophy in members of the Archaeogastropoda is very rare and is limited to several small groups. Mississippian–Late Permian. In addition to the Streptacidoidea. 1993. Nevertheless. ´ 1975. Yochelson (1984) concluded that the concept of the Macluritina uniting the superfamilies Macluritoidea and Euomphaloidea should be abandoned. 2004). He interpreted the Euomphaloidea as an independent superfamily derived from Knight et al. Architectonicoidea Gray. 2002). on the basis of his study of the early shell ontogeny of the onychochiloidean genus Mimospira Koken. RELATIONSHIPS OF MACLURITOIDEA TO EUOMPHALOIDEA Yochelson (1956) considered the Early Ordovician Macluritoidea to be the ancestral group of the Euomphaloidea. Bandel. Wangberg-Eriksson. and Kuskokwimiidae Fryda and ´ Blodgett. 1994. and Neomphalidae.g.12) resembles that in the Euomphaloidea (Yoo. Linsley and Kier. the similarity in early shell morphology suggests a dextrally coiled common ancestor and seems to support the old concept uniting both of these superfamilies in a common higher taxon. 1984).

¨ ¨ BATTEN. and D. ANDERSON. K. and possibly the Euomphaloidea in the new order Hyperstrophina of the new class Paragastropoda. Somerset. 2004). P. ` Troisieme partie. However. Thus the dextral hyperstrophy of the Macluritoidea is a derived and not a primary shell feature.. 59: 1011–1027. 70(3–4):325–365. Monatshefte. .. 1985. 1966. 104. Myrow. Mitteilungen aus dem Geologisch-Palaonto¨ logischen Institut der Universitat Hamburg. Vigo. M. 2001. ´ 1999. the discovery of dextral coiling of the early shell in Macluritella stantoni (Fig. Journal of Paleontology. COSSMANN. was dextral orthostrophic at this ontogenetic stage and so it had the same type of soft-body-shell arrangement (anatomically dextral body in dextrally coiled shell) as in the vast majority of gastropods. L. Colorado College. 2001. The Pennsylvanian gastropod genera Orthonema Meek and Worthen and Streptacis Meek from the Appalachian Basin. AND M. ´ ´ DONALD. BANDEL. Interpretation of juvenile Macluritella. K. 2001. Dissimilarity in protoconch ´ morphologies of the Onychochiloidea and Euomphaloidea was interpreted as a basis for suggesting that the class Paragastropoda is an artificial group (Fryda. R. FRYDA. R. Paris.. STURGEON. The adult teleoconch of this species is sinistrally coiled as are those in all macluritoidean gastropods. AND D. sinistrally coiled) shells. STURGEON. 305–306. Gastropoda) from the Late Ordovician of Bohemia. 1993. M. 2. J. 1916. Our discovery of sinistral heterostrophy in the oldest macluritid genus. 9:503–550. D. 78.1–2:103–131. The Paragastropoda has been considered to unite untorted mollusks. C450:41–81. 1999. KNIGHT. Twelfth International Malacological Congress. D. HOARE. ¨ ´ BANDEL. However. 1797. provided invaluable assistance with his knowledge of the Manitou Formation. ´ 2001) and Fryda and Rohr (1999. Vestnık Ustrednıho ustavu geoˇ ´ ´ ˇ ´ ´ logickeho. Observations on the genus Aclisina de Koninck. 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The sinistrally coiled teleoconch of the Macluritoidea has been interpreted as dextrally hyperstrophic. ´ ´ Moreover. Abstract. 1998. T. 2) as well as its shell heterostrophy provides evidence that the Macluritoidea and Onychochiloidea are not closely related taxa. G. The Pennsylvanian gastropod Donaldina Knight in the Appalachian Basin. 1999).. R. 64:557–562. Some heterostrophic gastropods from Triassic St. Nutzel (Germany). AND M. Annales du Musee royal d’Histoire na` ´ ´ turelle de Belgique. with descriptions of British species and of some other Carboniferous Gastropoda. R. Thus juvenile Macluritella stantoni. Proceedings of the Malacological Society of London. and the Ordovician ‘‘sinistral hyperstrophic’’ Mimospira is mentioned as their possible example. 66:200–205. 2002. Ponder and Lindberg (1997. J. 33:262–264. 3. 1983. Palaeontologische Zeitschrift. ACKNOWLEDGMENTS P. As noted above. 1996. Bandel and Fryda. Macluritella. Mode of life of a new onychochilid mollusc from the Lower Devonian of Bohemia. J. which is the oldest macluritid gastropod. B. 1960. as dextrally orthostrophic suggests that the Macluritoidea evolved from the dextrally orthostrophic gastropods (Haeckels biogenetic law). Faune du calcaire carbonifere de la Belgique. there is no evidence that Mimospira and related genera belonging to the superfamily Onychochiloidea were hyperstrophic. 2004). AND J. 119–120:1–109. DZIK. Journal of Paleontology. Fryda. curved. 1992. 1898. Freiberger Forschungshefte. Tableau Elementaire de l’Histoire Naturelle des Animaux. About the Heterostropha (Gastropoda) from the Carboniferous and Permian. Fryda ´ ´ and Rohr. Vetigastropoda). sinistrally coiled shells and most probably were not closely related to the Macluritoidea (Dzik. 2001. They suggested that such an ancestor might be found among the Macluritoidea. 2004). 1994. Cassian Formation with a discussion of the classification of the Allogastropoda. J. Wagner. 1881. serie paleontologique. 6. A. ¨ BANDEL. Evolutionary history of sinistral archaeogastropods with and without slit (Cirroidea. The Quarterly Journal of the Geological Society of London. p. sinistrally coiled protoconch consisting of several whorls (Dzik. 2004) pointed out that members ´ of the Onychochiloidea and Euomphaloidea have quite different protoconch morphologies. 1992. The Lower Carboniferous gastropod fauna from the Hotwells limestone of Compton Martin. 2004). the Onychochiloidea have homeostrophic. CONCLUSIONS 269 Macluritoidea and Onychochiloidea are not closely related taxa.A. A Devonian member of the subclass Heterostropha (Gastropoda) with valvatoid shell shape. ´ ´ FRYDA. 1995. AND J. eastern U. Early whorls in Macluritella stantoni are openly and dextrally coiled. This protoconch type is ´ unlike that of the Archaeogastropoda and it has been interpreted as a true larval shell (protoconch II.. COX. 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