Taxonomy and Trunk-Ring Architecture of Pleurojulid Millipedes (Diplopoda: Chilognatha: Pleurojulida) from the Pennsylvanian of Europe and North America Author(s): Heather M. Wilson and Joseph T. Hannibal Source: Journal of Paleontology , Nov., 2005, Vol. 79, No. 6 (Nov., 2005), pp. 1105-1119 Published by: Paleontological Society Stable URL: https://www.jstor.org/stable/4094997 REFERENCES Linked references are available on JSTOR for this article: https://www.jstor.org/stable/4094997?seq=1&cid=pdfreference#references_tab_contents You may need to log in to JSTOR to access the linked references. JSTOR is a not-for-profit service that helps scholars, researchers, and students discover, use, and build upon a wide range of content in a trusted digital archive. We use information technology and tools to increase productivity and facilitate new forms of scholarship. For more information about JSTOR, please contact support@jstor.org. Your use of the JSTOR archive indicates your acceptance of the Terms & Conditions of Use, available at https://about.jstor.org/terms SEPM Society for Sedimentary Geology and are collaborating with JSTOR to digitize, preserve and extend access to Journal of Paleontology This content downloaded from 205.173.218.15 on Tue, 30 Mar 2021 15:35:15 UTC All use subject to https://about.jstor.org/terms J. Paleont., 79(6), 2005, pp. 1105-1119 Copyright C 2005, The Paleontological Society 0022-3360/05/0079-1105$03.00 TAXONOMY AND TRUNK-RING ARCHITECTURE OF PLEUROJULID MILLIPEDES (DIPLOPODA: CHILOGNATHA: PLEUROJULIDA) FROM THE PENNSYLVANIAN OF EUROPE AND NORTH AMERICA HEATHER M. WILSON',2 AND JOSEPH T. HANNIBAL3 'Department of Entomology, 4112 Plant Sciences Building, University of Maryland, College Park 20742; 2Current address: Department of Geology and Geophysics, Yale University, P.O. Box 208109, New Haven, Connecticut 06520-8109, <heather m wilson@yahoo.com> and 3The Cleveland Museum of Natural History, 1 Wade Oval Drive, Ohio 44106-1767, <jhanniba@cmnh.org> ABSTRACT-Pleurojulid millipedes, known since the turn of the last century to be relatively abundant in the Westphalian D (Carboniferous: Pennsylvanian) Gaskohl of Nyfany, Czech Republic, are here also identified as an important component of the Pennsylvanian (Westphalian D) Mazon Creek millipede fauna preserved in ironstone nodules. Pleurojulids reach lengths approaching 10 cm, have as many as 69 body segments, medium-sized heads, and large ocellaria with upwards of 40 ocelli. Pleurojulids have previously been interpreted as having either a juliform-like or a colobognathan-like trunk-ring architecture. In order to distinguish between these two hypotheses, almost all pleurojulid specimens in museum collections were surveyed to document the deformation pattern of exoskeletal elements to aid in reconstruction of the trunk-ring architecture. The Nyfany specimens are completely flattened while the Mazon Creek specimens retain a degree of three-dimensionality. In order to assess how trunk-ring architecture controls patterns of deformation, a variety of extant millipedes were experimentally compressed. The distribution of exoskeletal elements in pleurojulid fossils was most similar to that seen in compressed extant polyzoniid millipedes. Based on the available evidence, pleurojulid trunk-ring architecture is reconstructed as semicircular in cross section, consisting of arched diplotergites, free pleurites firmly articulated to the lateral margins of the tergites and held in a near horizontal position, and free sternites. Pleurojulida are hypothesized to be basal helminthomorph, the sister group to Colobognatha, though inclusion in Helminthomorpha is equivocal. The taxonomy of previously described pleurojulid millipedes from N'fany is revised and newly recognized specimens from Mazon Creek specimens are described. Two genera are recognized within the new order Pleurojulida: Pleurojulus and Isojulus. Two species of Pleurojulus are recognized: P. biornatus and P. levis. Pleurojulus aculeatus and P. pinguis are synonymized with P. levis. Only one species of Isojulus, I. constans, is recognized with L setipes, I. marginatus synonymized with it along with Pleurojulus longipes and P. falcifer. INTRODUCTION THE UPPER Carboniferous strata at Nyfany in the Czech Re- public and Mazon Creek in Illinois comprise two of the most important Paleozoic Konservat-Lagerstaitten for millipedes in particular and terrestrial arthropods in general. Millipedes are generally rare in the fossil record due to their terrestrial habitus, but are superbly preserved and relatively abundant at these two localities. Many of the millipede taxa found at Ncirany and Mazon Creek were described around the turn of the last century by Fritsch (1899) and Scudder (e.g., 1889, 1890), respectively. Of these taxa, two groups in particular have captured the attention of those who have dealings with the Paleozoic terrestrial arthropod faunas and have frequently been figured in textbooks and treatises: the spiny euphoberiids, due to their fantastic armature, and the pleurojulids described by Fritsch (1899) from Nyfany, because of their ready acceptance as bona fide millipedes, due in large part to their superb preservation. Many of the Nyfany specimens are near complete with heads, legs, and tergal microsculpture preserved. Acceptance of pleurojulids as bona fide millipedes was a significant event in millipede taxonomy. Historically, all Paleozoic millipedes had been placed in Archipolypoda, a separate class from extant millipedes, due to mistaken morphological interpretations that led investigators to believe they were sufficiently anatomically different from extant millipedes as to preclude accommodation within the existing taxonomic framework (for a review of the history of the taxon Archipolypoda and a rediagnosis, see Wilson and Anderson, 2004). With all the attention that has been given to pleurojulids, it is surprising to note that none were ever previously described from Mazon Creek (Baird and Anderson, 1997) even though, as documented herein, they comprise a large proportion of the known millipede fossils from this locality. However, the fossils have not gone completely unnoticed as Langford (1963) mistakenly identified one or more specimens of Pleurojulus Fritsch, 1899 from Mazon Creek as a soft-bodied annelid. Superb preservation notwithstanding, the trunk-ring architecture of pleurojulid millipedes has been subject to various interpretations. Fritsch (1899) envisioned pleurojulids as having juliform-like trunk rings, roughly circular in cross section, but with visible sutures delimiting the portion of the ring formed by the pleurite from that formed by the tergite. Silvestri (1903, p. 103-105) interpreted these structures as ventral paratergites (in his terminology = true pleurites) and stated that the paratergites in Pleurojulidae were separate from the sternites as in Colobognatha. Sharov (1966, p. 70) re- garded the pleurites as paratergal lobes that were freely articulated with the tergites and which would have projected laterally. In con- trast, Hoffman (1969, p. R595) viewed the sutures separating the tergites and pleurites simply as cracks that occurred when pleurotergites were flattened. Kraus (1974) supported the contention of earlier authors that the pleurites were discrete structures. It is im- plied by Dzik (1981) that he also accepted the discrete nature of the pleurites as he suggested that Pleurojulus represents the oldestknown colobognath millipede, calling attention to the similarity in tergal morphology to that of Polyzonium Brandt, 1837. However, in light of the unspecialized nature of the pleurojulid head, Dzik (1981) also suggested that a relationship to the Platydesmida should be considered. Most recently, Schneider and Werneburg (1998) echoed Sharov (1966) in hypothesizing that the pleurites in Pleurojulus were oriented horizontally to form a roof over the legs. The purpose of this paper is to describe newly identified pleurojulid millipedes from the Mazon Creek biota and also to document their presence in the Westphalian D of Glamorganshire, Wales. In addition, a thorough redescription of taxonomy and morphology of pleurojulids utilizing both the Nyfany and Mazon Creek material is undertaken. The goal was to understand better pleurojulid trunk-ring architecture and to document character states for use in formulating hypotheses about the phylogenetic position of pleurojulids within Diplopoda. GEOLOGICAL SETTING AND PRESERVATION Pleurojulid millipedes occur at three Westpha in Europe and North America: Nyfany in the C 1105 This content downloaded from 205.173.218.15 on Tue, 30 Mar 2021 15:35:15 UTC All use subject to https://about.jstor.org/terms 1106 JOURNAL OF PALEONTOLOGY, V. 79, NO. 6, 2005 SYSTEMATIC PALEONTOLOGY Material examined is housed in the following colle rodni Museum (NMP), Prague, Czech Republic; Brit ical Survey (BGS), Keyworth, United Kingdom; Na 300 seum of Scotland (NMS), Edinburgh, United Kin Natural History Museum (NHM), London, Unite United States National Museum of Natural Histo Washington, DC; Field Museum of Natural Histo Panthalassic Ocean Chicago; the Peabody Museum of Natural History 00 University, New Haven, Connecticut; Illinois St ny Pal Tethys Sea (ISM), Springfield; and the Museum fuir Naturkund boldt-Universitiit (MfNB), Berlin, Germany. Class DIPLOPODA Blainville in Gervais, 1844 Subclass CHILOGNATHA Latreille, 1802-1803 Infraclass? HELMINTHOMORPHA Pocock, 1887 Order PLEUROJULIDA new order (=Eurystema Verhoeff, 1926 in part) 300 Diagnosis.-As for family. Included families.-Pleurojulidae. Family PLEUROJULIDAE Schneider and Werneburg, 1998 FIGURE 1-Late Pennsylvanian paleogeographic reconstruction in Mollweide projection with land shaded and showing the location of the three Westphalian D localities which have produced fossils of pleurojulid millipedes. (After Scotese and McKerrow, 1990, fig. 19, and Emended diagnosis.-Large millipedes, approaching 10 cm in length, body composed of at least 69 segments. Head capsule juliform-like with short, robust antennae and two lateral ovoid to subcircular ocellaria with numerous ocelli. Collum small; as wide as, or slightly wider than, head. Tergites with distinct middorsal suture; prozonite short; metazonite smooth or with ornament. Cwm Clydach in Wales, and Mazon Creek in Illinois (Fig. 1). In Prominent paired lateral ozopores in midmetazonite position, the Westphalian, all three of these localities were located on the starting on fourth postcollum trunk segment and continuing to same paleocontinent in an equatorial position. The associated milpenultimate segment. Free ventral diplopleurites subrectangular lipede fauna found at Nyfany and Mazon Creek is similar, conand with marginal rim; anteromedial and posteromedial comers sisting mainly of euphoberiids, oniscomorphs, and juliform milrounded; same length as tergites. Sternites free and unfused. Legs lipedes such as xyloiulids (e.g., Scudder, 1889, 1890; Fritsch, with seven segments and terminal claw. 1899; Burke, 1979; Hannibal and Feldmann, 1981; Hannibal, Discussion.-Fritsch (1899, p. 27) placed Isojulus Fritsch, 1899 1997). At Nyfany the specimens are preserved as impression fos- and Pleurojulus in the Projuloidae within the Chilognatha, tosils in Gaskohle from the Nyfany Member of the Kladno For-gether with Anthracojulus Fritsch, 1899 and Pylojulus Fritsch, mation (Holub, 1988). The Mazon Creek specimens are preserved1899 (=Xyloiulus Cook, 1895). Projuloidae was not based on a www.scotese.com/late.html) as external and internal molds in siderite nodules found within the Francis Creek Shale Member of the Carbondale Formation. genus and was a wastebasket taxon for fossil millipedes that were considered at the time to be closely allied to the Juliformia. VerMany of these specimens come from the Braidwood assemblage, hoeff (1926, p. 355) followed Fritsch's classification although he collected from strip-mine spoil dumps in Grundy, Will, and replaced KanProjuloidae with Eurysterna, a taxon erroneously defined kakee counties. The sediments of this portion of the Mazon Creek by the occurrence of broad sternites with widely separated coxal complex have been interpreted as being deposited in a brackish sockets. Laurentiaux (1953, p. 391) utitlized Verhoeff's taxono- to freshwater environment as part of a prograding delta complex my. Sharov (1962, 1991) further contributed to the mdlange by (Baird et al., 1985; Baird, 1997a, 1997b). The single specimen synonymizing Projulidae with Archijulidae Scudder, 1873 with from Wales is preserved in shale with a thin layer of coalified the addition of Tomiulus Martynov, 1936 and Purkynia Fritsch, cuticle. 1899. Hoffman (1969, p. R595) did not attempt to place the pleuMATERIALS AND METHODS rojulids within a family, but suggested that they were referable to the juliform millipedes. Schneider and Werneburg (1998) erected Select Mazon Creek specimens, or, for the sake conservaa newof family Pleurojulidae to accommodate the genus Pleurojution, only partial specimens, were prepared and lus, castincluding in latex using a new species, P. steuri. However, the specimens methods outlined in Hannibal (2001). Millipedesdescribed representing the as P. steuri have lateral paranota rather than ventrolatextant orders Polyzoniida (Siphonotus sp.), Callipodida [Abacion eral pleurites and are not pleurojulid millipedes. Schneider and magnum (Loomis, 1943)], and Julida [Cylindroiulus punctatus Wemrneberg (1998) did not include Isojulus in the Pleurojulidae, (Leach, 1815)] were experimentally compressedsuggesting in orderthat to it get a is allied with the Plagiascetidae. However, as rough idea of the relationship between trunk-ring architecture and has the same trunk-ring architecture as discussed below, Isojulus deformation pattern among extant orders. Millipedes preserved inof a diplotergite, pair of free diplopleuriPleurojulus, consisting 70%-100% ethanol were treated overnight with protease (one tes, and two freetabsternites. An emended diagnosis of Pleurojuli- let of Rite Aid brand enzymatic cleaner containing subtilisin A dae, including both Pleurojulus and Isojulus, is given above. dissolved in 25 ml of distilled water) to mimic limited decay. The treated millipedes were then mounted between two glassGenus microPLEUROJULUS Fritsch, 1899 scope slides using Euparal mounting medium. The slides were compressed using a Comten Industries model SSB1000 materials [=Pleuroiulus VERHOEFF 1910-1914, p. 33, pl. 2, fig. 28; VERHOEFF, 1926-1928, p. 342; LAURENTIAUX, 1953, p. 391, and others]. testing device with a maximum force of -400 N. This content downloaded from 205.173.218.15 on Tue, 30 Mar 2021 15:35:15 UTC All use subject to https://about.jstor.org/terms WILSON AND HANNIBAL-CARBONIFEROUS PLEUROJULID MILLIPEDES 1107 Type species.-Pleurojulus biornatus Fritsch, Types.-Syntypes 1899, designated NMP 54125, 510, 512. by Hoffman, 1969. Other material examined.--NMS 1903.95.5; NMP St504; BGS 48790-1. Other species.-P. levis Fritsch, 1899. Diagnosis.---Ocellaria as wide as long. CollumOccurrence.-Pennsylvanian, semicircular in Westphalian D, Nyffany Horizo shape, as wide as head. Pleurites with broadlyofrounded posterothe Kladno Formation, Nyfany, Czech Republic; Westphalia medial corners. D, Tenuis Zone, Clydach Merthyr Colliery, Cwm Clydach, G Discussion.-Pleurojulus can be distinguished from contemmorganshire, Wales. poraneous Isojulus based on the width of the collum segmentDiscussion.-Fritsch (1899, p. 20-29) described two additiona (wider than the head in Isojulus and narrower, approximatelyspecies as of Pleurojulus, P. aculeatus and P. pinguis, each based wide as the head in Pleurojulus), the shape of the ocellaria on (as a single specimen. Fritsch diagnosed P. aculeatus based on wide as long in Pleurojulus and wider than long in Isojulus), dimpling and of the tergal cuticle, though he suggested that the h the lack of lateral wrinkling of the tergites, which is only seen in lotype might actually represent a specimen of P. levis. Upon r Isojulus). examination, this dimpling appears to be taphonomic as it is n ther regular nor continuous across the specimen. As there are PLEUROJULUS BIORNATUS Fritsch, 1899 other features to distinguish it from P. levis, the two species a Figure 2.5, 2.6 here synonymized. The features that Fritsch used to distingui Pleurojulus biornatus FRITSCH, 1899, p. 27-28, pl. 139, figs. 1-9;P.pl.pinguis from P. levis also appear to be taphonomic. The ho 143, fig. 8; VERHOEFF, 1926-1932, figs. 77-82. lotype of this species is preserved in a tight 'C' with a great d of segmental overlap in comparison with many of the Pleuroju Diagnosis.-Prozonites covered with very fine striations. Despecimens where the trunk is telescoped to a varying degree pressed anterior band of metazonite covered with pits or irregular Thus, features that Fritsch described as distinctive are actuall longitudinal striations. Remainder of metazonite covered with structures of one segment impressed through the overlying cutic fine, slightly irregular, longitudinal striae. Pleurites generally of another segment. For example, Fritsch described the pleurit smooth, though occasionally with faint punctae. of P. pinguis as unique because they have a ridge that truncat Description.-Large millipede, reconstructed length approachthe anterior of the pleurite. However, it appears that this rid ing 8 cm, elongate, tapering front and back, up to about 66 body represents the posterior margin of the preceding pleurite. We he segments. Head poorly preserved in NMP IFG Me7, IFG synonymize 606 P. pinguis with P. levis. This leaves only two va Me17 (Fig. 2.5). Antennae with robust, distally flaring antennomNrfany species of Pleurojulus (P. biornatus and P. levis) as P. eres. Ocellaria poorly preserved. Ornament of collum unknown. longipes Fritsch, 1899 and P. falcifer Fritsch, 1899 are synony Prozonite covered with very fine striations, terminating with mized rim with Isojulus constans Fritsch, 1899 as discussed below (Fig. 2.5, 2.6). Metazonite with depressed anterior band of pits or PLEUROJULUS cf. BIORNATUS Fritsch, 1899 irregular longitudinal striae, remainder ornamented by fine, slightFigures 3-6, 9.1 ly irregular, longitudinal striae; posterior with pronounced raised rim (Fig. 2.5). Ozopores located at midheight of tergites in subsoft bodied annelid LANGFORD, 1963, fig. 19a-d. circular pits midway between middorsal suture and lateral tergal margin (Fig. 2.5). Pleurites generally smooth, with raised rim on Diagnosis.-Large millipedes with free, thick-rimmed, diplo all margins except lateral. pleurites, medium-sized heads, large ocellaria, and up to about Types.-Syntypes NMP IFG 606 Mel7, IFG 589 Me7, IFGbody segments. Prozonite fairly smooth, metazonite covered wi 588. irregularly transverse striations. Diplopleurites smooth or wit Other material examined.-NMP 509, IFG 590 Me8, 35125, vague ornamentation. 491/35425, 512. Description.--Large millipede, reconstructed length approach Occurrence.--Pennsylvanian, Westphalian D, N'fany Horizon ing 10 cm, elongate, tapering front and back, up to about 66 bod of the Kladno Formation, Czech Republic. segments. Height (flattened) at midpoint up to about 5.7 mm. Discussion.-As the head is poorly preserved in the PleuroHead subrounded, with medial longitudinal depression (Fig julus biornatus specimens, the only character that can be reliably3.4). Ocellarium large and (?)reniform, composed of about 40 used to distinguish P. biornatus from P. levis is the tergal orna- ocelli (Fig. 3.5). Mandibles stout (Figs. 3.1, 4). Antennae wi ment, with the tergites of P. levis being relatively unornamented. cone-shaped segments (Fig. 3.4). Collum small, slightly narrow than head, (?)smooth (Figs. 3.4, 5.3). Tergal prozonites short, metazonites long; prozonites raise Figure 2.1-2.4 posteriorly, separated from metazonites by groove which begi Pleurojulus levis FRITSCH, 1899, p. 28, pl. 141, figs. 1-11; KUHN, 1949, as sharp-edged constriction at distal margin of prozonite. Bot fig. 155, la, b; MOLLER, 1963, fig. 198; 1992, figs. 177, 178; SHAROV, prozonites and metazonites marked by fine horizontal striatio 1966, fig. 29; KRAUS, 1974, fig. 4; SHAROV, 1991, fig. 7. (15-20 per mm), striations more pronounced and grade into sm Pleurojulus FRITSCH, 1899; MULLER AND ZIMMERMANN, 1962, p. 199, pits anteriorly on metazonites. Ozopores located at about mid fig. 152; HOFFMAN, 1969, p. 595, fig. 379. height on metazonites in suboval depressions (Figs. 3.1, 3.4, 5.1 Pleurojulus aculeatus FRITSCH, 1899, p. 28, pl. 141, figs. 12-14. PLEUROJULUS LEVIS Fritsch, 1899 Pleurojulus pinguis FRITSCH, 1899, p. 29, pl. 140, figs. 8, 9; VERHOEFF, 5.3) on fifth to penultimate tergites. Diplopleurites with raised rims on anterior, posterior, and ven 1926-1932, fig. 83. tral borders, thickest on anterior border; anteromedial border Diagnosis.-Diplotergite generally smooth, diplopleurites usu-rounded, posteroventral border strongly rounded; surface marked ally have ornamentation of fine horizontal striations. by faint ridges generally radiating towards borders (Fig. 6.4). Description.-Large millipede with reconstructed length over Sternites narrow with acutely rounded lateral margins (Fig. 6.2, 4 cm, up to 69 trunk segments. Head slightly wider than long6.3), anteromedial area forming raised triangle (Fig. 6.4), narrow with longitudinal groove at position of presumed internal median elliptical grooves adjacent to coxal sockets housing spiracles (Fig. septum (Fig. 2.2). Antennomeres short, robust, and flared distally. 6.2, 6.3). Both sternites of same trunk segment with similar shape. Ocellaria large, with at least 80 ocelli. Collum same width as head, without middorsal suture. Legs with seven segments and terminal claw, when fully extended as long as half width of tergite (Figs. 3.1, 5.1, 5.3). Coxae This content downloaded from 205.173.218.15 on Tue, 30 Mar 2021 15:35:15 UTC All use subject to https://about.jstor.org/terms 1108 JOURNAL OF PALEONTOLOGY, V. 79, NO. 6, 2005 Is Itt ilit 4 1 I; V,2 Au Is 04. Is,- azirr ?c 'Pol :ss;R' IfI .oil I Is Af i 1IC -4LP ?I1~a J4.1- Is, 4k. VF~ ORC This content downloaded from 205.173.218.15 on Tue, 30 Mar 2021 15:35:15 UTC All use subject to https://about.jstor.org/terms WILSON AND HANNIBAL-CARBONIFEROUS PLEUROJULID MILLIPEDES 1109 ISOJULUS CONSTANS of adjacent legs in close proximity, no evidence of eversible ves-(Fritsch, 1879) Figures icles (Fig. 6.4), approximately 1.3 times wider than long (Fig.7, 8 5.1). Trochanter very reduced, five times wider than long, wedgeJulus constans FRITSCH, shaped (Figs. 5.4, 6.1). Prefemur and femur similar size,1879. nearly Archijulus constans FRITSCH, 1895, p. 2. square with recurved anterodorsal margins (Fig. 6.1). Postfemur Isojulus constans FRITSCH, 1899, p. 25-26, pl. 142, figs. 1-3, text-fig. approximately 1.5 times longer than wide 336. (Figs. 3.2, 6.1). Tibia elongate, approximately four times longer than (Fig. Isojulus setipeswide FRITSCH, 1899, 3.2). p. 26, pl. 142, figs. 4-8. Tarsus shorter than tibia, approximatelyIsojulus 3.7 times than marginatuslonger FRITSCH, 1899, p. 26-27, pl. 140, figs. 1, 2; pl. 142, wide (Fig. 3.2). Terminal claw short (Fig. figs. 3.2). 9, 16.Leg 10 possibly modified as gonopod in males (Fig. 4). Pleurojulus longipes FRITSCH, 1899, p. 28-29, pl. 140, figs. 3-9. Pleurojulus falcifer FRITSCH, 1899, p.6.4). 29, pl. 140, fig. 10. Anal opening located ventrally, oval, relatively small (Fig. Shape of valves as in Rhinotus purpureus (Pocock, 1894) (Polyzoniida: Siphonotidae) (see Hoffman, 1990, fig. 26.9). Diagnosis.-Large pleurojulidan millipede with reconstructed Material examined.-FMNH PE 42, PE 3272, PE 11296, PE length approaching 7 cm, at least 55 trunk segments. Collum 26921, PE 29360, PE 29358, PE 29445, PE 30612, PE 32259, short, slightly wider than head; ocellaria wider than long; tergites PE 32260, PE 32261, PE 32262, PE 32266, PE 32267, PE 32269, finely punctate with prominent longitudinal striations near lateral PE 32274, PE 32279, PE 32282, PE 32288, PE 32293, PE 32296, margins; legs with numerous setal sockets. PE 32300, PE 32301, PE 32310, PE 32312, PE 32313, PE 32315, Description.-Head and collum preserved in NMS PE 32316, PE 32323, PE 32324, PE 32325, PE 32329, PE 32360, 1898.105.21 (Fig. 7.1), NMP IFG 593 MelO, and NMP K1040 PE 39258, PE 39361, PE 51789, PE 45721 (in coprolite); ISM (Fig. 7.2). Labrum nowhere completely preserved. Ocellaria wid- 14855; YPM 42861; USNM 379991. Preserved in siderite coner than long, subovoid in shape (Fig. 7.2). Collum short, slightly cretions from the Francis Creek Shale Member (Upper Pennsylwider than head, with raised marginal rim (Fig. 7.1, 7.2). Collum vanian, Westphalian D) of the Carbondale Formation, part ofin the NMP K1040 (Fig. 7.2) possibly with middorsal suture as in Mazon Creek fauna. Collected from old coal strip-mine dumps in postcollum tergites. Postcollum tergites with longitudinal midWill, Grundy, or Kankakee counties, Illinois (see Hannibal and dorsal suture (Fig. 7.1-7.3), raised rim on posterior margin, finely Feldmann, 1981, text-fig. 2; Baird et al., 1985, fig. 3). Many specpunctate surface texture, prominent longitudinal striations near imens were collected in Pits 1 or 6, which straddle the Grundy lateral margins (Fig. 7.2). Paired, slanting ozopores located at and Will county lines (see Wilmington and Coal City, Illinois, midheight of tergite midway between middorsal suture and lateral U.S. Geological Survey 7.5-minute-quadrangle topographic margin of tergite. Ozopores present in continuous series beginning maps). These localities are within the Braidwood (freshwater)with as- fifth tergite (Fig. 7.1), ending with posteriormost leg-bearing semblage. Additional information on the Mazon Creek localities segment (last two trunk rings anterior to telson appear apodous and their biota can be found in the articles in Shabica and Hay without ozopores). Pleurites, clearly visible at posterior of coun(1997). terpart of IFG 600 Me15 (Fig. 8.2), with straight lateral margin, Discussion.-The Mazon Creek species of Pleurojulus resembroadly rounded anteromedial and posteromedial comers. Legs bles Pleurojulus biornatus Fritsch from Nyfany so closely that relatively stout, composed of seven segments (Fig. 7.3), covered we were unable to identify any reliable characters with which to numerous fine pits interpreted as setal sockets. Sternites not with differentiate consistently between specimens from these twoclearly lopreserved. calities. This is due in large part to variation in ornamentationType.-Holotype NMP K1040. present within each fossil population. This is reflected in Fritsch's Other material examined.-NMS 1898.105.21; NHM In. original description of P. biornatus, which embraced millipedes 59546; NMP IFG 593 MelO, IFG599 M1040, IFG 600 Mel5 IFG 594 Me lt, IFG 596 M1008. with variable ornamentation of the tergites. The ornament seems to be more pronounced in many of the Nyi~any specimens (comOccurrence.-Pennsylvanian, Westphalian D, Nyfany Member pare Fig. 2.5 with 3.3), however there is no obvious way toof adthe Kladno Formation, Czech Republic. dress whether this difference is real or due to taphonomy. ProDiscussion.-Fritsch (1899, p. 25) described the genus Isojulus portions of the skeletal elements appear to be similar between as lacking separate pleurites. However, several specimens of IsoNyfany and Mazon Creek specimens. julus preserve unarguable evidence of distinct pleurites. In fact, Genus ISOJULUS Fritsch, 1899 the specimen that exhibits the most completely preserved pleurites is the counterpart of the specimen designated as the holotype of I. setipes by Fritsch (1899) (Fig. 8.2). Thus the ring structure of [=Isoiulus VERHOEFF 1910-1914, p. 33; 1926-1928, p. 342; LAURENIsojulus appears to have been very similar to that of Pleurojulus TIAUX, 1953, p. 391-392, and others]. and we place them together in the family Pleurojulidae. Isojulus can be distinguished from Pleurojulus by the presence of longiType species-Isojulus constans Fritsch, 1899, designated by tudinal striations on the lateral ends of the tergites, the lack of Hoffman, 1969. other tergal ornament (other than fine punctae), by ocellaria that Diagnosis.-As for genus. are wider than long [in Pleurojulus the ocellaria have nearly equal FIGURE 2-1-4, Pleurojulus levis Fritsch, 1899, Westphalian D, Nrfany Member, Kladno Formation, Czech Republic. 1, NMP 54125, entire specimen preserved in lateral aspect; 2, enlarged view of the anterior of NMP 54125, showing details of the collum and head, including the eyes and antennae; 3, NMS 1903.59.5, posterior of specimen in lateral aspect; 4, NMP 35125, preserved in lateral aspect showing pleurites partially detached from lateral margin of tergites. 5, 6, Pleurojulus biornatus Fritsch, 1899, locality and horizon as above. 5, NMP IFG 606 Mel7, preserved in dorsal aspect showing head, with left antenna preserved, and anterior tergites with characteristic ornament; 6, NMP 509, preserved in dorsal aspect and with characteristic ornament. Scale bars = 2 mm. an, antenna; c, collum; h, head; ms, middorsal suture; oc, ocellarium; oz, ozopore; pl, pleurite; st, sternite; ta, trunk appendages; tg, tergite. This content downloaded from 205.173.218.15 on Tue, 30 Mar 2021 15:35:15 UTC All use subject to https://about.jstor.org/terms 1110 JOURNAL OF PALEONTOLOGY, V. 79, NO. 6, 2005 ttw ~m 1? Ee A.~ II )TAL 2,tg 4 3 6P I-5 pre This content downloaded from 205.173.218.15 on Tue, 30 Mar 2021 15:35:15 UTC All use subject to https://about.jstor.org/terms WILSON AND HANNIBAL-CARBONIFEROUS PLEUROJULID MILLIPEDES 1111 lengths and widths (Fig. 2.2)] and by a collum that is wider than the head [in Pleurojulus it is the same width or slightly narrower than the head (Fig. 2.1, 2.2)]. Only one species of Isojulus, L constans, is recognized here as there are insufficient diagnostic characters to maintain the three species diagnosed by Fritsch (1899). Fritsch diagnosed L setipes based on what appear to be numerous setal sockets on the appendages. However, the same distribution of setal sockets is seen on the better-preserved appendages of the holotype of L constans, so we synonymize these two species. Isojulus marginatus was diagnosed by Fritsch (1899, p. 26) based on the ridges along the anterior and posterior margins of the tergites and a tergal ornament consisting of fine punctae. Both of these characters are also present in the holotype of L constans and the two species are here synonymized. The specimen upon which Fritsch based his description of Pleurojulus longipes has no obvious tergal ornament and has the longitudinal striations near the lateral margins of the tergites characteristic of Isojulus. For these reasons it is also synonymized with L constans. Pleurojulus falcifer was diagnosed by Fritsch based on a single specimen that he described as having very small pleurites and fine punctae on the tergites. He even suggested that it was possible that a new genus would need to be erected for this specimen because the pleurites appeared to be st significantly reduced relative to other species t5 of Pleurojulus. However, the pleurites are not completely preserved in the holotype, with only fragments preserved at the anterior of the speci- tU t8 t9 til men, which is preserved in dorsal aspect. The rest of the features, including the finely punctate surface ornamentation, is consistent t4 9 PI with an Isojulus constans identity, and the two species are here synonymized. 1112 13 TRUNK-RING ARCHITECTURE It is generally agreed that the millipede trunk ring is composed Q2 sp 6 primitively of discrete, articulated, exoskeletal components-tergites, pleurites, and sternites-and these components have been variously fused in different lineages. In both extant and extinct helminthomorph millipedes, there are three basic trunk-ring architectures: the colobognathan architecture in which the tergites, pleurites, and sternites are free (Fig. 9.2); the nematophoran architecture in which the tergites and pleurites are fused but the 0 1 2 mm sternites remain free (Fig. 9.3); and the juliformian/polydesmidan architecture in which the all the trunk-ring components are fused4-Pleurojulus cf. biorna FIGURE (Fig. 9.4). In colobognathans the pleurites are held underneath the Shale Member (Carboniferous: Pe mation, tergites in a near-horizontal position, though in some cases theyIllinois. 1, Enlarged view anterior are slightly angled ventrad or dorsad, giving a near semicircular legs and possible gonop pretative cross section. In nematophorans, juliformians, and polydesmi- drawing of 1. an, anten oc, ocellarium; pl, pleurite; sp, spi dans, the trunk-ring cross section is essentially circular to subcircular, modified in some cases by the presence of lateral paranota. As noted in the introduction, Pleurojulus has been compared previously to both juliformian (Fritsch, 1899, p. trunk-ring 25) and colobogarchitectures described above, were exper nathan millipedes, particularly polyzoniids (Silvestri, 1903,We p. found that the majority of pleurojulid compressed. 105; Dzik, 1981), producing two very different reconstructions of and Mazon Creek are preserved in late at both Nyfany trunk-ring architecture (Fig. 9.1). In an attempt to distinguish be-partially coiled. The specimens preser either fully or have typically folded along the natural line of tween these competing hypotheses, we surveyedaspect the pattern of provided by the median dorsal suture on the tergites trunk-ring deformation in the Nffany and Mazon Creek pleurooften have visible pleurites, which are typically pres julid fossils. We also investigated the patterns of deformation exhibited when extant millipedes, representative of the the ventral three main margin of the tergites (Figs. 2.1, 3.1, 5). In 41 20 1 2 4 14 2 FIGURE 3-Pleurojulus cf. biornatus Fritsch, 1899, Francis Creek 1, FMNH PE 29445, latex cast of natural mold of left side of a showing detail of podomeres; 3, FMNH PE 32312 showing de head, collum, and anterior trunk segments in dorsal view; 5, en for 3, 4, 2 mm. c, collum, cl, claw; cx, coxa; fe, femur; h, head pl, pleurite; prfe, prefemur; t, tarsus; tg, tergite; ti, tibia. This content downloaded from 205.173.218.15 on Tue, 30 Mar 2021 15:35:15 UTC All use subject to https://about.jstor.org/terms 1112 JOURNAL OF PALEONTOLOGY, V. 79, NO. 6, 2005 'tw. .4, If .. 464 it'' 41 _ JII 141;a it~9 'vk ~ ~ r-y ~~ It-lei ~ [ 4t k ., ? I ?nl -4-1% 'PIK t.-l~b ir ?r-1 ,- ? ?YI . 4:iL V11; A At This content downloaded from 205.173.218.15 on Tue, 30 Mar 2021 15:35:15 UTC All use subject to https://about.jstor.org/terms WILSON AND HANNIBAL-CARBONIFEROUS PLEUROJULID MILLIPEDES 1113 FA -W T tg; S| -.60 7 ~tL.~ a~s~. ~ .-7 , /,~` Nk A-~l %Al __ ,ot-. e Zjkid" 4 + e 7 .-.-'f-" ... - - ,o_` $4L - 41 II. + +AV' 19,, 'A,. N6 IV ra , FIGURE 6-Pleurojulus cf. biornatus, Francis Creek Shale Member ( of FMNH PE 51789, part and counterpart, showing details of the st from tergites and slid dorsad over the tergites; pleurites cover mo center of the figure; anterior is to the left; 2, pleurites, sternites, tergites; 4, FMNH PE 32301, posterior of specimen preserved in ven trunk appendages. Scale bar for 1-3, 1 mm; for 4, 2 mm. av, an spiracle; st, sternite; tg, tergite; tr, trochanter. the pleurites remain articulated with the tergites (Figs. are 2.3 aspect, some of the pleurites and in others the pleurites have disarticulated from the terg beneath the tergites (e.g., Isoj often sliding over or under 3.1, the 5.1). tergites (Figs. 2.4, 3.1, 5.3 A small proportion of 6.1). In a small proportion of the specimens preserved inall la dorsal or ventral aspect. In FIGURE 5-Pleurojulus cf. biornatus, Francis Creek Shale Member PE 32282, natural mold of left side; 2, FMNH PE 32312, natural m stem branches which may represent a layer of flotsam; 3, FMNH of FMNH PE 32274 showing detail of the pleurites and trunk app This content downloaded from 205.173.218.15 on Tue, 30 Mar 2021 15:35:15 UTC All use subject to https://about.jstor.org/terms 1114 JOURNAL OF PALEONTOLOGY, V. 79, NO. 6, 2005 O,~. A q rIF " )2r . , Aw fT ' ? j, 2 qe 3op 4V vi~C LCI~d:( FIGURE 7-Isojulus constans Fritsch, 1899, Westphalian D, Nrf specimen preserved in dorsal aspect showing head, collum, an K1040, anterior of specimen preserved in dorsal aspect sho preserved in lateral aspect showing legs, tergites, and partial pl oz, ozopore; pl, pleurite; ta, trunk appendages; tg, tergite. pattern strikingly th preserved ventral to the tergites (Figs. 2.5,similar 3.3, 6,to 7.1, Sternites are either not well preserved obscured in all N (Fig. 10). In or the polyzoniid spe shown in Figure 10, the head h specimens. In extant millipedes with free sternites, the cu the sternites is often less robust than that of with the tergites aspect (compare Pleuro 2.2). TheWhile pleurites at theare anter was probably true in pleurojulids. sternites pr in many of the Mazon Creek specimens, they are gener swung out and downwards to r scured in all but a few specimens. Where they are of visib tergites. At the posterior th sternites are typically preserved near medial margin outwards to the be 'preserved' alo rites. position The pleurites did pleurites (Fig. 6.1-6.3). The exact relative tonot the sw p varies, suggesting that they werewhere relatively loosely conn region a large longitudin the pleurites by membrane.gites to slide laterad. An inf In the compression studybetween of extant millipedes, we the orientation offoun the in taxa with free sternites, the membranous attachmen the polyzoniid in Figure 10.3 6.1-6.3). It is interesting not pleurites provided a significant point of structuralto weak this particular polyzoniid wer lowing the tergites to either swing outwards when dorsov served specimens, a large propo compressed or inwards when laterally compressed. The along the ventral margin th niid millipede Siphonotus Brandt, 1837 exhibited a of defo This content downloaded from 205.173.218.15 on Tue, 30 Mar 2021 15:35:15 UTC All use subject to https://about.jstor.org/terms WILSON AND HANNIBAL-CARBONIFEROUS PLEUROJULID MILLIPEDES 1115 b-.. -ONO[-- V, Jr, All> 1'rv,* IS. oz -m s oz I4r tg m 10 46 A s 4 ooo? 71 l 00 . op ?lw W. it jw . a owe or Ar All 10; - ol -zoo a 40' ed0o% 10* Now Ak 4?, ir'r St .00, __JW J* A POW- -AW Z . .- _90". _??* 411 ., . - - :P-? .1 .. '_ 'o, opr 3P MI-4, t Af A_ PF, 40r^ .4r ,b .0, 10% jol 44" It FIGURE 8-Isojulus constans (NMP IFG 600 Me 15), Westphalian D, N"fiany Member, Kladno Formation, Czech Republic. 1, Part preserving posterior tergites and legs; 2, counterpart preserving ventral morphology including legs and pleurites posteriorly. Scale bars = 2 mm. Ms, middorsal suture; oz, ozopore; pl, pleurite; st, sternite; ta, trunk appendages; tg, tergite. Intuitively one might have thought that the pleurites would swing inwards and upwards when such a specimen is laterally compressed (as in oniscomorph fossils; see Hannibal and Feldmann, 1981, pl. 1, figs. 1, 2, 6). In contrast to the polyzoniid, when the extant juliformian and nematophoran millipedes were compressed, there were no obvious parallels in the deformation pattern with that of the pleurojulid fossils. Given the lack of free pleurites in these taxa, these results are not surprising. Of course these compression experiments provide only a very rough analog to the compression that would occur during taphonomy, as the pleurojulid specimens would have been supported to a degree by sediment matrix during compression. However, the compression experiments do suggest that millipedes with a polyzoniid-type trunk-ring architecture do have the propensity to deform to produce a configuration of ring elements similar to that seen in the pleurojulid fossils. The irregular boundary between the pleurites and tergites seen on many Mazon Creek specimens suggests that it is possible that the articulation was not as mobile as that in extant polyzoniids and the two structures may have been more firmly attached. This content downloaded from 205.173.218.15 on Tue, 30 Mar 2021 15:35:15 UTC All use subject to https://about.jstor.org/terms - 1116 JOURNAL OF PALEONTOLOGY, V. 79, NO. 6, 2005 ""'?~?~-?. ?? .::::~?X O 'SO t r Z Wr ~ H ~ /I ~ U ~ ~ ~ ~? ?? // ~ nY u~ ur/ II n n .. .. ~u ~n ~r r I r N H" .:::' 1 2 3 4 FIGURE 9-Mi grey and cox ventral posit colobognatha juliformian Basic based engin ing m a cylindrica ancest perspective that t tazonian mi an arch lowing (th segme sternites). M pleuri pass dorsad eightmuscles orig tenna preceding s and n arched desig length arches com rings in an arched curve of the (discu Engho rites have a The lo ends of the ever it compression other ventral mar such as in S telopo such as Pol either mens, desmus Ger femal muscle conn times If the free a comp position, as or du the pleurote capabl weakness, le produ ative to an a ulatio favor a colo popul lids in whic would (Fig. 9.1, so possib Phylogenet nearly clades of ch anteri + Platydes served desmida + S (Figs. chitecture i relativ ever, it is d preser phylogeneti gap in of nearly a pretat shown in a the si ancestral ch This content downloaded from 205.173.218.15 on Tue, 30 Mar 2021 15:35:15 UTC All use subject to https://about.jstor.org/terms WILSON AND HANNIBAL-CARBONIFEROUS PLEUROJULID MILLIPEDES 1117 't, i 1 1 1 unmodified proximal portio sternite, representing legs that its distal portion is ob distal portion of leg nine is Leg 10, associated with terg may be a gonopod. It is broa posterior projection that opp mented structure that may an incompletely preserved, appears to be associated wit legs to tergites is not an ex millipedes because it is diff millipede, to determine wh Recent work on segmentati gests that dorsal and ventr group, leading to dorsoventr mental elements because t directly to either ventral or 2004). Without additional sp parison, it is impossible to d FMNH *: Pbx ~us?~ 19"e-)4i d 2 j =rS~i~?) ~rIr ta being .st aa */ . 29445 is indeed more closely relate Sphaerotheriida + Glomeri tates that pleurojulids lack their posteriormost legs m dence for telopods in pleur point, it would be surprisin lipede the length of a pleu an *B PE or partially preserved, unm lent state of preservation specimen (Fig. 3.1), the pos opod warrants serious consi All extant male helmintho notable exceptions (e.g., Lo least leg pair 9 modified, eith gonopods. While leg pair 10 thomorphs, including colob in no extant taxa is this leg a summary of leg modifica derson (2004, fig. 12). The lipede Archidesmus macnico lipede known in which onl Anderson, 2004). In the mi oticopodus Wilson and And polypodan affinities, only l sible that Pleurojulus belong millipedes in which the go posterior position relative to We also need to consider t not helminthomorph millipe . * Glomeridesmida has been is not know documented in (Haacker, 1974). In some on female and grasp her anteri proceed to enroll such that posterior side of the coxae 3C~ I- -** " contact with the female's g stridulating millipede Lobo male grasps one antenna a FIGURE 10--The extant millipede Siphonotus Brandt, 1837 telopods and then proceeds t 1, Right side of millipede in lateral aspect-the pleurites are tive (Haacker, 1970). In a m ventrally; 2, same millipede compressed between two she grasping the posterior of t the head has rotated to be preserved in dorsal aspect and th enrollment trick both seem have swung outwards to be preserved adjacent to the ven female for sperm transfer. of the tergites; 3, interpretative drawing of the flattened p identified in pleurojulids antenna; oz, ozopore; pl, pleurite; st, sternite; ta, trunk eit app tergite. Scale bar = 2 mm. iferous Euphoberiida (Hann ?I ? --- This content downloaded from 205.173.218.15 on Tue, 30 Mar 2021 15:35:15 UTC All use subject to https://about.jstor.org/terms 1118 JOURNAL OF PALEONTOLOGY, V. 79, NO. 6, 2005 commun., and HMW, personal observation, 2001) by has idensupported thealso Kirtlandia Society (CMNH). Helpful reviews tified an extant fuhrmannodesmidan millipede (Polydesmida: Tri-and W. Shear. The authors were were provided by G. Edgecombe chopolydesmoidea) with a pair of midbodysupported legs that appear to be by National Science Foundation grant DEB-0075605 modified for clasping, although this behavior has beenthe obto HMW andnever grants from Smead Staff Enrichment Fund served. So the position of modified appendages inRose Pleurojulida (CMNH) and M. Louer Fund (FMNH) to JTH. cannot be determined at this point with any certainty, however, REFERENCES limited evidence points to the presence of gonopods modified from leg pair 10 and associated with tergite eight. BAIRD, G. C. 1997a. Geologic setting of the Mazon Cre A second synapomorphy of Pentazonia is deposit, the presence p. 16-20.of In diC. W. Shabica and A. A. Hay (eds.), vided sternites (Enghoff, 1984). As the sternites Pleurojulus Guide to thein Fossil Fauna of Mazon Creek. Northeastern Illinois Uniare undivided, the Pleurojulida cannot be accommodated within versity, Chicago. Pentzonia and only a sister-group position can G. beC.considered. In BAIRD, 1997b. Paleoenvironmental setting of the Mazon Creek addition, Pentazonia are characterized by the lack of an internal biota, p. 35-51. In C. W. Shabica and A. A. Hay (eds.), Richardson's Guidepresence to the Fossil Fauna Creek. Northeastern Illinois Unisagittal phragma on the cranium and by the ofofaMazon large versity, Chicago. postantennal organ (Hoffman, 1982). However, a sagittal phragma G. C., AND J. of L. ANDERSON. is present in Pleurojulus, as evidenced by BAIRD, the presence a sag- 1997. Relative abundance of different Mazon Creek organisms, ittal suture on the cranium (well preserved in NMP 54125, Fig.p. 27-29. In C. W. Shabica and A. A. Hay (eds.), Richardson's Guide to the Fossil Fauna of Mazon Creek. 2.2) and Pleurojulus does not appear to have a large postantennal Northeastern Illinois University, Chicago. organ (Figs. 2.2, 4). However, if it turnedBAIRD, out that G. C., C.Pleurojulida W. SHABICA, J. L. ANDERSON, AND E. S. RICHARDSON lack gonopods, they could possibly represent a lineage of basal muddy coast: Habitats within the JR. 1985. Biota of a Pennsylvanian Chilognatha that branched off before the Pentazonia-HelminthoMazonian Delta complex, northeast Illinois. Journal of Paleontology, 59:253-281. morpha split. The only other Paleozoic millipede with aBRANDT, trunk-ring J. E 1837.architecNote sur un ordre nouveau de la classe des Myr podes et sur 10 tablissement des sections de cette classe d'animaux e ture similar to that of Pleurojulus is Zosterogrammus stichostescientifique thus Wilson, in press, from Mazon Creek, general. whichBulletin belongs to ande l'Acad6mie imp6riale des Sciences Saint-P6tersbourg, 5:307-315. extinct order, the Zosterogrammida, which ranges from the MidBURKE, J. J. 1979. new millipede genus, Myriacantherpestes (Dip dle Silurian through the Pennsylvanian of Europe andA North poda, Archipolypoda) and a new species, Myriacantherpestes brade birksi, from the English Coal Measures. Kirtlandia, 30:1-24. stichostethus lacks a middorsal suture, the trunk is shorter than COOK, O. E 1895. Introductory notes on the families of Diplopoda, p America. The main differences between the two taxa are that Z. Pleurojulus, the pleurites are much broader than those of Pleu1-7. In O. E Cook and G. N. Collins (eds.), The Craspedosomatida rojulus, and the sternites are divided (Wilson, in press). Although of North America. Annals of the New York Academy of Sciences, 1-100. the trunk-ring architecture is similar, there are no characters that suggest a phylogenetic proximity between Zosterogrammida andJ. 1981. An early Triassic millipede from Siberia and its evoluDZIK, tionary significance. Neues Jahrbuch fir Geologie und Palaiontologie Pleurojulida, with the similarities pointing to a diversity of PaleoMonatshefte, 7:395-404. zoic millipedes with primitive trunk-ring architectures. ENGHOFF, H. 1978. Parthenogenesis and spanandry in millipedes. AbGiven the available evidence, we hypothesize that the Pleuro- handlungen und Verhandlungen des Naturwissenschaftlichen Vereins julida form the sister group to the Colobognatha. The Colobogin Hamburg, 21/22:73-85. natha are characterized by having eight pairs of legs in front of ENGHOFF, H. 1984. Phylogeny of millipedes-a cladistic analysis. Zeitthe simple, leglike gonopods and mouthparts variously reduced. schrift fUir zoologische systematik und Evolutionsforschung, 22:8-26. Hoffman (1979) suggested that Colobognatha is not a natural ENGHOFF, H. 1990. The ground-plan of chilognathan millipedes (external group, with the simple nature of the gonopods and their segmental morphology), p. 1-21. In A. Minelli (ed.), Proceedings of the 7th In- position being primitive helminthomorph characters and the ternational Congress Myriapodology. E. J. Brill, Leiden. FRITSCH, A. 1879. Neueof.. Ubersicht in der Gaskohle und den Kalksteinen mouthparts having been independently reduced, as evidenced by der Performation in Bbhmen vorge funde den Tierreste. Sitzungsberithe wide range of cephalic morphology in colobognaths. He also chte der Koniglichen b6hmischen Gesselschaft der Wissenschaften, suggested that Polyzoniida may have a closer affinity to GlomPrague, 1879:184-195. eridesmida than to the other colobognath taxa. However, a recent FRITSCH, A. 1895. Vorldiufiger Bericht tiber die Arthropoden und Molmolecular phylogenetic analysis of three protein-encoding genes lusken der bijhmischen Permformation. Sitzungsberichte Gesellschaft supports a monophyletic Colobognatha (Regier et al., 2005), as der Wissenschaften (Jahrgang), 1894:1-4. does a recent morphological analysis that expanded and reanaFRITSCH, A. 1899. Fauna der Gaskohle und der Kalksteine der Permforlyzed Enghoff's (1984) character set (Sierwald et al., 2003). mation B6hmens, Volume 4, Pt. 1. E Rivnic, Prague, 152 p. GERVAIS, P. 1844. Etudes sur les Myriapodes. Annales des Sciences Naturelles, series 3, 2:51-80. Thanks go to the following for hospitality and/or the of HAACKER, U. loan 1969. Spermatibertragung von Glomeris (Diplopoda). NaACKNOWLEDGMENTS turwissenschaften, 56:467. material under their care: G. Buckley, S. Lidgard, and W. Taylor HAACKER, 1970. R. Der Stridulationsapparat von Loboglomeris und seine (FMNH); C. MacClintock and T. White (YPM); V. TurekU.and Funktionand im Sexualverhalten. Vie Milieu, 20:57-64. Prokop (NMP); E. Pietrzeniuk (MfNB); J. Thompson C. LaHAACKER, U. 1974. Patterns of communication in courtship and mating bandeira (USNM); B. Paton (NMS); M. Howe (BGS); and A. Ross (NHM). Some of the photographs of Mazon Creek speci- behaviour of millipedes (Diplopoda). Zoological Society of London Symposia, 32:317-328. mens were taken by B. Frumker and printed by D. Flocke and G. HANNIBAL, J. T. 1995. Modified legs (clasping appendages?) of CarbonPetusky, Cleveland Museum of Natural History (CMNH). Helpful iferous euphoberiid millipeds (Diplopoda: Euphoberiida). Journal of discussion and comments were supplied by: J. Schneider, TU BerPaleontology, 69:932-938. gakademie Freiburg; J. Shultz, University of Maryland; and colHANNIBAL, J. T. 1997. Myriapods and arthropleurids, p. 172-183. In C. leagues at the 10th International Congress of Myriapodology, esW. Shabica and A. A. Hay (eds.), Richardson's Guide to the Fossil pecially S. Golovatch, Russian Academy of Sciences, Moscow, Fauna of Mazon Creek. Northeastern Illinois University, Chicago. and H. Enghoff, University of Copenhagen. Several of the latex HANNIBAL, J. T. 2001. Hexecontasoma, a new helminthomorph millipede casts used in this study were prepared by C. Tome, who was (Hexecontasomatidae, n. fam.) from the Mazon Creek, Illinois, fauna This content downloaded from 205.173.218.15 on Tue, 30 Mar 2021 15:35:15 UTC All use subject to https://about.jstor.org/terms WILSON AND HANNIBAL--CARBONIFEROUS PLEUROJULID MILLIPEDES 1119 (Carboniferous, North America), p. 19-35. In J. Wytwer andMULLER, S. I. Go- A. H. 1963. Lehrbuch der Paliozoologoie, Volume 2, Pt. lovatch (eds.), Progress in Studies on Myriapoda and Onychophora. Gustav Fischer, Jena, 698 p. Proceedings of the 11th International Congress of Myriapodology. MULLER, A. H. 1992. Lehrbuch der Palaozoologie. Baud 1. Allgeme Fragmenta Faunistica, 43 (Supplement 2000). Grundlagen. 5. Auflage. G. Fischer Verlag, Stuttgart, 496 p. MULLER, HANNIBAL, J. T., AND R. M. FELDMANN. 1981. Systematics and func- A. H., AND H. ZIMMERMANN. 1962. Aus Jahrmillionen Tie der Vorzeit. Veb Gustav Fischer Verlag, Jena, 409 p. tional morphology of oniscomorph millipedes (Arthropoda: Diplopoda) NEWPORT, from the Carboniferous of North America. Journal of Paleontology, 55: G. 1844. A list of the species of Myriapoda, order Chilog 730-746. natha, contained in the cabinets of the British Museum, with descr tions of a new genus and thirty-two new species. Annals and Magaz HOFFMAN, R. L. 1969. Myriapoda, exclusive of Insecta, p. R572-R606. History, 13:263-269. In R. C. Moore (ed.), Treatise on Invertebrate Paleontology. of Pt.Natural R. PEACH, B. N. 1882. On some fossil myriapods from the Lower Old R Arthropoda 4. Vol. 2. Geological Society of America and University Sandstone of Forfarshire. Proceedings of the Royal Physical Societ of Kansas Press, Lawrence. 77:177-188. HOFFMAN, R. L. 1979. Classification of the Diplopoda. Museum POCOCK, R. I. 1887. On the classification of the Diplopoda. Annals an D'Histoire Naturelle, Geneve, 237 p. of Natural History, ser. 5, 20(35):283-295. HOFFMAN, R. L. 1982. Diplopoda, p. 689-724. In S. P ParkerMagazine (ed.), POCOCK, R. I. 1894. Contributions to our knowledge of the arthrop Synopsis and Classification of Living Organisms. Vol. 2. McGraw-Hill, fauna of the West Indies, Pt. III, Diplopoda and Malacopoda, wit supplement on the Arachnida of the class Pedipalpi. Journal of th HOFFMAN, R. L. 1990. Diplopoda, p. 835-860. In D. L. Dindel (ed.), Soil Linnean Society of London (Zoology), 24:473-544, pls. 37-40. Biology Guide. John Wiley and Sons, New York. HOLUB, V. 1988. Geology and stratigraphy of Permo-Carboniferous REGIER, conJ. C., H. M. WILSON, AND J. W. SHULTZ. 2005. Phylogenet analysis of Myriapoda using three nuclear protein-coding genes. M tinental basins of the Bohemian Massif in view of the latest research lecular Phylogenetics and Evolution, 34:147-158. and analysis of stratigraphical methods used, p. 37-43. In J. Pelek and SCHNEIDER, J. W, AND R. WERNEBURG. 1998. Arthropleura und Dip J. VozIir (eds.), Coal-Bearing Formations of Czechoslovakia. Dionyz poda (Arthropoda) aus dem Unter-Rotliegend (Unter-Perm, Assel) d Stdir Institute of Geology, Bratislava. Thtiringer Waldes (Stidwest-Saale-Senke). Ver6ffentlichungen Natu JANSSEN, R., N.-M. PRPIC, AND W. G. M. DAMEN. 2004. Gene expression historisches Museum Schleusingen, 13:19-36. suggests decoupled dorsal and ventral segmentation in the millipede SCOTESE, C. R., AND W. S. MCKERROW. 1990. Revised world maps a Glomeris marginata (Myriapoda: Diplopoda). Developmental Biology, introduction, p. 1-21. In W. S. McKerrow and C. R. Scotese (eds 268:89-104. New York. Palaeozoic Palaeogeography and Biogeography. Geological Socie KRAUS, 0. 1974. On the morphology of Palaeozoic diplopods, p.Memoir, 13-22. No. 12. In J. G. Blower (ed.), Symposia of the Zoological Society ofSCUDDER, London, S. H. 1873. On the Carboniferous myriapods preserved in t 32. sigillarian stumps of Nova Scotia. Boston Society of Natural Histor KUHN, 0. 1949. Lehrbuch der Palaozoologie. E. Schweizerbart, Stuttgart, Memoir, 2:231-239. 326 p. SCUDDER, S. H. 1889. Archipolypoda, a subordinal type of spined myrLANGFORD, G. 1963. The Wilmington Coal Fauna and Additions to the iapods from the Carboniferous formation. Boston Society of Natural Wilmington Coal Flora from a Pennsylvanian Deposit in Will County, History Memoir, 3:143-182. Illinois. Esconi Associates, Downers Gove, Illinois, 280 p. SCUDDER, S. H. 1890. New Carboniferous Myriapoda from Illinois. Boston Society of Natural History Memoir, 4:417-442, pls. 33-38. LATREILLE, P. A. 1802-1803. Histoire naturelle, g6n6rale et particulibre SHABICA, C. W., AND A. A. HAY (EDS.). 1997. Richardson's Guide to the des Crustac6s et des Insectes; ouvrage faisant suite aux oeuvres de Fossil Fauna of Mazon Creek. Northeastern Illinois University, ChiLeclerc de Buffon, et partie du cours complet d'histoire naturelle r6dig6 cago, 308 p. par C. S. Sonnini. Volume 2. E Dufart, Paris, 467 p. SHAROV, A. G. 1962. Class Diplopoda, p. 22-24. In Yu. A. Orlov (ed.), LATZEL, R. 1886. Les Myriapodes des la Normanider (2e liste) suivie de Osnovy Paleontologii. Tom 9: Chlenistonogie, Trakheinye, i Khelitsediagnoses d'especes et de vari6t6s nouvelles (de France, Alg6rie et rovye. Akademiya Nauk SSSR, Moscow. Tunisie). Bulletin de la Soci6t6 des Amis des Sciences naturelles, SHAROV, A. G. 1966. Basic Arthropodan Stock with Special Reference Rouen, 71:165-177. LAURENTIAUX, D. 1953. Classe des Myriapodes (Myriapoda Leach,to Insects. Permagon Press, Oxford, 271 p. SHAROV, A. G. 1991. Class Diplopoda, p. 6-10. In B.B. Rohdendorf 1814), p. 385-395. In J. Piveteau (ed.), Trait6 de Paldontologie. Volume (ed.), Fundamentals of Paleontology. Volume 9. Smithsonian Institution 3. Masson et Cie, Paris. Libraries and the National Science Foundation, Washington, DC. LEACH, W. E. 1815. A tabular view of the external characters of four SIERWALD, P, W. A. SHEAR, R. M. SHELLEY, AND J. E. BOND. 2003. classes of animals which Linnd arranged under Insecta. Transactions Millipede phylogeny revisited in light of the enigmatic order Siphonof the Linnean Society of London, 11:306-400. iulida. Journal of Zoological Systematics and Evolutionary Research, LOOMIs, H. E 1943. New cave and epigean millipeds of the United States,41:87-99. with notes on some established species. Bulletin of the Museum of SILVESTRI, P. 1903. Classis Diplopoda. Volume 1. Anatome. Vesu Comparative Zoology, 92:371-410. Portici, 272 p. LooMIs, H. E, AND R. L. HOFFMAN. 1962. A remarkable new family of VERHOEFF, K. W. 1910-1914. Die Diplopoden Deutschlands. C. E spined polydesmoid Diplopoda, including a species lacking gonopods ter, Leipzig, 640 p. in the male sex. Proceedings of the Biological Society of Washington, VERHOEFF, K. W. 1926-1932. Diplopoda, p. 1-2084. In H. G. 75:145-158. (ed.), Klassen und Ordnungen des Tier-Reichs. Akademische V MANTON, S. M. 1961. The evolution of arthropodan locomotory mech- 5(2). Leipzig, anisms. Pt. 7. Functional requirements and body design in ColobogWILSON, H. M. In press. Zosterogrammida, a new order of mil from the Middle Silurian of Scotland the Upper Carboniferous natha (Diplopoda), together with a comparative account of diplopod ramerica. Palaeontology. burrowing techniques, trunk musculature and segmentation. Journal of WILSON, H. M., AND L. I. ANDERSON. 2004. Morphology and tax the Linnean Society of London (Zoology), 44:383-461. of Paleozoic millipedes (Diplopoda: Chilognatha: Archipolypoda) MARTYNOV, A. V. 1936. O Nekotorykh novykh materialakh chelenistonScotland. ogikh zhivotnykh iz kuznetskogo basseina. Izvestiia Akademii nauk Journal of Paleontology, 78:169-184. SSSR, Seriia Biologicheskaia, 6:1258-1260. (In Russian) ACCEPTED 10 AUGUST 2004 This content downloaded from 205.173.218.15 on Tue, 30 Mar 2021 15:35:15 UTC All use subject to https://about.jstor.org/terms