This is in contrast to the suggestion by Shelley (2001) that Hoffmanobolidae may be affiliated with Atopetholidae based on the coxite structure (Fig. For all Bayesian analyses, we consider clades with posterior probabilities of 0.69 and below to be poorly supported, of 0.70–0.89 to be moderately supported, and of 0.90–1.00 to be strongly supported. Both Spirobolidae (Keeton, 1960a) and Atopetholidae (Hoffman and Orcutt, 1960) were revised, but neither of the works addressed the position of these families in relation to other families in the order. Length of telopodite of anterior gonopod: produced ventrally beyond level of anterior portion of coxite (state 0); not produced ventrally beyond level of anterior portion of coxite (state 1). All figures derived from SEM studies are provided with scale bars. (Wesener et al., 2008, character 37). Arthrosphaera Topology resulting from Bayesian inference analysis of a dataset containing combined morphological and molecular data. 2), but he did make predictions that we can examine in light of our results. Attems (1909, 1910) split this lineage into Euspirobolidae (much the same as Spirobolidae of Cook) and Trigoniulidae (much the same as Pachybolidae of Cook) based on the presence or absence of a sternite connecting the posterior gonopods, respectively; this established the major morphological differentiation that has been used to diagnose the suborders Spirobolidea and Trigoniulidea to this day. Forms in either of these areas, if present, have yet to be discovered. Arrangement of coxite and telopodite of posterior gonopods: angled in relation to each other (state 0); linear in relation to each other (state 1). The tree produced using information from Hoffman’s (1980) taxonomy resolves the suborders as monophyletic, with each being a polytomy beyond that point (Fig. The number of rows in the pectinate lamellae: four (state 0); five (state 1); six (state 2); seven (state 3); eight (state 4); nine (state 5); eleven (state 6). In rare instances where only a single strand for a target gene region was sequenced cleanly, the single strand was included in analyses if correcting base calls was negligible due to high‐quality sequence. Ventral pads on tarsi of males, leg pairs 4–7: absent (state 0); present (state 1). Paraspirobolus lucifugus. Spirobolidea sensu this study accommodates Allopocockiidae, Atopetholidae, Floridobolidae, Hoffmanobolidae, Messicobolidae, Pseudospirobolellidae, Spirobolellidae, Spirobolidae, and Typhlobolellidae. Sternite of anterior gonopod: absent (state 0) (Fig. Male specimens exhibit prominent ventrally produced endites on the coxites of the anterior gonopod (Fig. All parameters were analysed using the program Tracer 1.4 (Rambaut and Drummond, 2003) to determine that they had also reached stationarity. Shape of telopodite of anterior gonopod: uncinate (state 0); bilobed apicomedially (state 1); simple (state 2); modified apicomedially into a flange or bulb projecting somewhat laterally (state 3). Here we follow the approach of Wesener et al. Arthropod Phylogenetics in Light of Three Novel Millipede (Myriapoda: Diplopoda) Mitochondrial Genomes with Comments on the Appropriateness of Mitochondrial Genome Sequence Data for Inferring Deep Level Relationships. The labrum provides a number of characters at various levels within higher millipede relationships. Animal kingdom systematics In 1937, Haneda found a luminous millipede at Chuuk Islands in Micronesia and sent the specimens to a myriapod researcher, Yosioki Takakuwa (Haneda, 1972, 1985), which was described as a new species Spirobolellus phosphoreus (Takakuwa, 1941). Information on outgroup taxa can be found in Table 2. (Wesener et al., 2008, character 17). Secondary structure alignment resulted in an aligned sequence of 2020 bp, with 18S and 28S being 1754 and 265 bp, respectively. Whereas there is no suture between putative sternite and pleurotergite in Polydesmida, in the latter subterclass and order, a suture is present (Sierwald et al., 2003). This family was placed in Spirobolidea and distinguished from other families by the following two characters: presence of a distinct coxite in the posterior gonopods and presence of large apodemes associated with the coxite of the anterior gonopods. The average weight, length, and width of an adult millipede range from 4.5 to 12.5 g, from 3.5 to 6.5 cm, and from 1.5 to 2.5 cm, respectively, (Figure 2). Cycle sequencing reactions were purified and sequencing was conducted using an Applied Biosystems 3730 DNA Analyzer (Applied Biosystems). (Wesener et al., 2008, character 14). For ClustalX aligned data, mean base composition was A = 0.25785, C = 0.23116, G = 0.28212, T = 0.22887 with a homogenous nucleotide frequency among taxa (χ2 = 99.1574, d.f. The main The first use of the name Spirobolida to define the order was by Chamberlin (1943), although this change was not explained until Chamberlin and Hoffman’s (1958) publication. K.M.P. We also trace the history of Trigoniulinae due to the uncertainty of its taxonomic status. (Diplopoda Spirobolida Pachybolidae)1 T. Wesener 1,3 and H. enghoff 2 1 Zoologisches Forschungsmuseum Alexander Koenig, Museumsmeile Bonn, Adenauerallee 160, D … The program PAUP*4b10 (Swofford, 2002) was used to run unweighted MP analyses on datasets consisting of morphological data alone, combined ClustalX aligned molecular data and morphological data, and combined secondary structure aligned data and morphology. Analyses consisted of two simultaneous analyses employing one heated and three cold Markov chain Monte Carlo (MCMC) chains that were run for 1 × 106 generations sampling trees every 1000 generations. This was possible for the majority of species used in our analyses. He did not provide illustration of the character states used to define these taxa, making it somewhat cumbersome to interpret his definition and description without visual representation. Currently, no detailed study of this character suite across a large sampling of millipedes and using SEM technology is available. Shape of the mentum: with a large swollen area apically, between the lamellae linguales (Fig. In this group, the coxite of the anterior gonopod is elongated along its lateral axis (Fig. Spirobolida was recovered as monophyletic with strong support (58/100). As members of this family are known mainly from arid regions of North America and few other spirobolidan millipedes are known from this habitat type, it is not surprising that atopetholids would be morphologically unique; it is unclear whether any of the previously mentioned characteristics can be correlated with life in this habitat, but it is certain that future detailed studies of this family will identify morphological adaptations to this environment. . All members of these families share a common ancestor, and the diversity of this lineage is probably a result of a single common ancestor and subsequent radiation into North and Central America. Structure of lamellae lingualis: separated by anterior portion of mentum (state 0); not separated by anterior portion of mentum (state 1). This study demonstrates that suborder Spirobolidea (as previously defined) represents a paraphyletic lineage, with Rhinocricidae recovered as sister to all other spirobolidans. Median suture line on front of head (clypeus, labrum and epipharynx), extending upwards from labrum and visible above level of antennae: absent (state 0); present only dorsally above interocular area (state 1); present both above interocular area and on clypeus (state 2). Our terminal Pachybolus ligulatus exhibits two setae on one lamella lingualis and three on the other; examination of a second specimen revealed two setae per lamella lingualis, and this taxon was coded as having two setae. The science of millipede biology and taxonomy is called diplopodology: the study of diplopods. Morphological data were recorded from the same specimen as was used to collect molecular data in all cases, with the exception of Narceus americanus (Palisot de Beauvois), where a specimen from the same collection event and locality as the one sequenced was used to code morphological data. 1). Based on Bollman’s newly established subfamily, other workers began recognizing the lineage as family Spirobolidae soon thereafter (Verhoeff, 1893; Pocock, 1894). These millipedes can grow up to 6 inches long, and sport a beautiful blackish-blue color with orange-red highlights along the back of … (Wesener et al., 2008, character 32). We provide reference to the major advancements in Spirobolida taxonomy as they relate to the order, suborder, and family levels within this lineage, followed by a focus on advances made since Keeton’s (1960a) and Hoffman’s (1980) publications. 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