ZOOTAXAISSN 1175-5326 (print edition)

ISSN 1175-5334 (online edition)Copyright © 2010 · Magnolia Press

Zootaxa 2667: 51–63 (2010) www.mapress.com/zootaxa/ Article

Discovery of the centipede family Plutoniumidae (Chilopoda) in Asia: a new species of Theatops from China, and the taxonomic value of spiracle distributions in Scolopendromorpha

ZHIYONG DI1, ZHIJIAN CAO1, YINGLIANG WU1, SHIJIN YIN2, GREGORY D. EDGECOMBE3,4 & WENXIN LI1,4

1College of Life Sciences, Wuhan University, Wuhan 430072, China2South Central University for Nationalities, Wuhan Institute for Neuroscience and Neuroengineering, Wuhan 430074, China3The Natural History Museum, Department of Palaeontology, Cromwell Road, London SW7 5BD, United Kingdom. 4Corresponding authors. E-mails:(Gregory D. Edgecombe) g.edgecombe@nhm.ac.uk, (Wenxin Li) liwxlab@whu.edu.cn

Abstract

Theatops chuanensis n. sp. is proposed for a Chinese representative of Plutoniumidae, the first Asian record of this otherwise North American and southern European chilopod family. Its ultimate legs lack prefemoral and femoral spines, and the species is unique within Theatops in possessing spiracles on segment 7. The reliability of segment 7 spiracles as a criterion for diagnosing scolopendromorph genera is challenged by various lines of evidence. An updated key to genera and species of Plutoniumidae is provided.

Key words: Chilopoda, Plutoniumidae, Theatops chuanensis, taxonomy

Introduction

The discovery of species diversity of Plutoniumidae and generic concepts within the family date from the mid-19th through mid-20th centuries. Though Plutoniumidae has been monographed in modern times (Shelley 1997) and its taxonomy and biogeographic data continue to be refined (Shelley 2002), no new species have been discovered in nearly 60 years. Plutoniumidae is widely endorsed as a Palaearctic-Nearctic clade, until now known only from southern Europe and widely distributed in the United States and northern Mexico, with a total of six species in two genera. Our study exposes a major lacuna in previous knowledge of the family’s distribution; we describe a new species of Plutoniumidae from Sichuan Province, China, and assign it to a revised concept of the North American/southern European genus Theatops Newport, 1844. The Sichuan discovery extends the geographic range of Plutoniumidae by thousands of kilometers, the closest occurrences being in the Balkan Peninsula and in the western United States (Figure 1).

The taxonomic history of Plutoniumidae was recounted in detail by Shelley (1997: 58–62). Bollman (1893) established Plutoniinae as the first suprageneric grouping uniting Plutonium Cavanna, 1881, and Theatops Newport, 1844. This membership has remained nearly constant through subsequent concepts of Plutoniuminae (correction from Plutoniinae proposed by Shelley & Backeljau 1995) or Plutoniumidae (elevated to familial rank by Zapparoli 2009) to the present day. Plutonium remains known only from its type species, P. zwierleini Cavanna, 1881, originally proposed for material from Sicily, and later found in Sardinia, the Italian mainland, and southern Spain (Cavanna 1881; Würmli 1975; Shelley 1997; Zapparoli 2009). Bollman (1893) collated geographic data for the then known North American species of Theatops, T. posticus(Say, 1821) and T. spinicaudus (Wood, 1865). Additional species were later added by Chamberlin (T. californiensis Chamberlin, 1902, from California; T. phanus Chamberlin, 1951, from Texas). The taxonomic history of T. californiensis relative to the sole European congener, T. erythrocephalus (C.L. Koch, 1847), is

Accepted by A. Minelli: 18 Sep. 2010; published: 4 Nov. 2010 51

recounted by Shelley (1990, 1997: 60–61). Modern distribution maps for the five valid species of Theatopsare available in Shelley (1997, 2002).

FIGURE 1. Distribution of Plutoniumidae. New record in Sichuan Province, China, indicated by star (see Figure 19 for more detailed map). The dashed line indicates 40˚ North latitude.

The evolving classification of Plutoniumidae reflects a general trend towards recognition of more families within Scolopendromorpha. At one extreme, Kraepelin (1903) placed all scolopendromorphs in a single family, Scolopendridae, with three subfamilies—Scolopendrinae and Otostigminae, both with four ocelli on each side of the cephalic plate, and Cryptopinae, lacking ocelli. At the other extreme, Verhoeff (1906, 1907) proposed three superfamilies, each comprising two families: Cryptopina (Cryptopidae + Newportiidae), Theatopsina (Theatopsidae + Plutoniidae), and Scolopendrina (Scolopendridae + Scolopocryptopidae). Attems (1930) reverted to few families, an arrangement that became standard for the rest of the 20th century. Attems classification of Scolopendromorpha, like that of Kraepelin (1903), reverted to using ocelli as the basis for recognizing families, i.e., Scolopendridae (Scolopendrinae and Otostigminae) and Cryptopidae, and

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he divided the latter into three subfamilies: Cryptopinae; Theatopsinae (=Plutoniinae Bollman, 1893), and Scolopocryptopinae. In more recent times, the blind scolopendromorphs are classified as families based on segment numbers: Scolopocryptopidae (23 segments) is identified as a separate family from Plutoniuminae and Cryptopinae (21 segments), the latter two comprising Cryptopidae (Shelley 2002). The Chinese species Mimops orientalis Kraepelin, 1903, forms the basis for a separate monotypic family, Mimopidae (Lewis 2006).

Shelley (1997) reviewed the subfamily Plutoniuminae (here family Plutoniumidae), and like most previous workers (e.g., Attems 1930; Würmli 1975) recognized two genera: the monotypic Plutonium, and Theatops, with five species. He affirmed the taxon as monophyletic and regarded the different number of spiracles in Plutonium and Theatops to be a generic level feature. At the same time, Shelley (1997) reconstructed the relationships among the species of Plutoniuminae as P. zwierleini + (T. spinicaudus + (T. phanus + (T. erythrocephalus + (T. posticus + T. californiensis)))). The implicit assumption that Theatops is monophyletic is not, however, well established; the genus differs from Plutonium only in its spiracle distribution and cladistic analyses indicate that the condition in Theatops is plesiomorphic (Edgecombe & Koch 2008; Koch et al. 2010).

Shelley’s (1997) classification contradicted the proposal by Schileyko (1992, 1996) and Schileyko & Pavlinov (1997) that Plutonium and Theatops are not each others’ closest relatives. This was based on a hypothesis that the spiracle distribution of Plutonium, on all leg-bearing trunk segments except the first and last as in Geophilomorpha, was likely to be primitive for Scolopendromorpha as a whole. The Schileyko system favoured a high-ranking taxonomic separation of Plutonium from Theatops and other scolopendromorphs.

More recent cladistic analyses have corroborated the monophyly of Plutoniumidae, resolved Theatops as paraphyletic with respect to Plutonium (a relationship anticipated by Kraepelin 1903: 27, who depicted Theatops as ancestral to Plutonium), and led to new proposals about the phylogenetic position of Plutoniumidae within Scolopendromorpha. Edgecombe & Koch (2008) investigated the higher-level systematics of Scolopendromorpha to explore the phylogenetic importance of such traditional (but conflicting) characters as segment number, segmental position of spiracles, and the presence or absence of ocelli. The shortest cladograms resolved blind Scolopendromorpha as a basal grade within which a novel clade uniting Plutoniumidae with Scolopocryptopidae (sensu Shelley (2002)) and supported by new characters from the preoral chamber, was sister to all remaining scolopendromorphs. These relationships have varied in more recent analyses that included foregut characters (Koch et al. 2009) and addition genera and species (Edgecombe & Koch 2009; Koch et al. 2010). Blind Scolopendromorpha are either monophyletic or paraphyletic according to character weighting schemes, and Plutoniumidae cluster either within Scolopocryptopidae (sensu Shelley 2002) or resolve as its sister group. The most recent analyses corroborate monophyly of a clade with 23 segments, Scolopocryptopidae sensu Shelley (2002), to which Plutoniumidae is sister group (Koch et al. 2010). Recognising Plutoniumidae at the familial rank permits Scolopocryptopidae to be delimited as by Shelley (2002), and is followed herein.

Nearly all workers have identified Plutoniumidae as strictly limited to the Nearctic and Palaearctic. The only previous assignment of Asian diversity to the group was Tonkinodentus Schileyko, 1992, a monotypic genus from Vietnam. This assignment was dismissed by Shelley (1997) because the description lacked data from the taxonomically important ultimate leg pair and because the type locality was incongruous with the existing familial distribution. Restudy of the type species, T. lestes Schileyko, 1992, incorporating data from the ultimate legs, showed that Tonkinodentus actually belongs to the Scolopendridae (Schileyko 2007).

We propose herein Theatops chuanensis n. sp. for the first authentic Asian and Chinese plutoniumid. Seven genera in five families of Scolopendromorpha are therefore known to China (Wang & Mauriès 1996): Cryptops (Cryptopidae, Cryptopinae), Scolopocryptops and Dinocryptops (Scolopocryptopidae, Scolopocryptopinae), Theatops (Plutoniumidae, Plutoniuminae), Scolopendra (Scolopendridae, Scolopendrinae), Otostigmus and Rhysida (Scolopendridae, Otostigminae), and Mimops (Mimopidae).

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Material and methods

Illustrations and measurements were produced using a Motic K-700L stereomicroscope with an Abbe drawing tube and an ocular micrometer. Morphological terminology follows recommendations by Lewis et al. (2005) and Bonato et al. (2010). The type material of the new species is deposited in the Museum of Wuhan University, Wuhan, China (MWHU).

Taxonomy

Order Scolopendromorpha Pocock, 1895

Family Plutoniumidae Bollman, 1893

Plutoniinae: Bollman 1893: 165,168.Plutoniidae: Verhoeff 1906: 433. Theatopsidae: Verhoeff 1906 432; Ribaut 1915: 338.Theatopsina: Verhoeff 1906: 431; Verhoeff 1907: 246.Theatopsini: Attems 1926: 376. Theatopsinae: Attems 1930: 249–250; Bücherl 1941: 325; Schileyko 1992: 13. Theatopinae: Crabill 1960: 9; Schileyko 1992: 13.Plutoniuminae: Shelley & Backeljau 1995: 150; Shelley 1997: 70–71; Shelley 2002: 80.Plutoniumidae: Zapparoli 2009: 112.

Diagnosis: Moderate-size to large-bodied (apparent adult lengths 45–150 mm) Scolopendromorpha, with 21 leg-bearing segments. Colour generally yellowish, with pale or depigmented patches lateral to bases of antennae in ocellar positions. T1 with anterior transverse sulcus; longitudinal median suture on anterior part of T1, bifurcating into two oblique sutures posteriorly. Forcipular coxosternite with tooth plates; poison calyx extending into coxosternite. Complete paramedian sutures on T2–20. Gizzard with stiff anteriorly-directed projections; projections evenly curved, covered by multifurcating scales that spirally encircle the projection, branching into slender, needle-like spines. T21 nearly twice as long as T20, with or without median suture. Ultimate legs swollen, forcipulate, heavily sclerotized, basal podomeres nearly as wide as long, prefemora and femora with or without dorsal and ventral spines.�

Distribution: (Modified from Shelley 1997, figure 1). China, northern Sichuan; coastal Croatia and Montenegro, Bosnia-Herzegovina, southern Italian Peninsula (Campania), Sardinia, Sicily, Spain, and Portugal (see Serra 1983, fig. 3 for Iberian Peninsula; Shelley 1997, fig. 44, p. 103 for other European records); eastern North America generally east of the Central Plains and south of the Great Lakes, from western New England and the vicinity of Chicago, Illinois, to the south Florida Keys and central, southern, and western Texas; western North America from western Chihuahua and northern Baja California Norte, Mexico, to the southern Basin and Range Province in southern Nevada and south-western Utah, westward to the Pacific Ocean in northern California with a small segregated area in southwestern Oregon (see Shelley 1997: 81, 84, 91, 92, 97, figs. 12, 18, 31, 32, 39; Shelley 2002: fig. 148).

Remarks: Shelley’s (1997) comprehensive revision of Plutoniuminae remains up to date in most respects. The grouping of Plutonium and Theatops is here classified at the familial rank following elevation by Zapparoli (2009), which will also be adopted in the forthcoming Treatise on Zoology (Edgecombe & Bonato 2010).

Theatops Newport, 1844

Theatops Newport, 1844: 409. Attems 1930: 250–251. Shelley 1997: 71–74 (see full synonymy p. 71); Shelley 2002: 81.Opisthemega Wood, 1862: 35 (synonymy by Pocock 1888: 287).Opisthomega Saussure & Humbert, 1872: 200.

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Type species: Cryptops postica Say, 1821, by subsequent monotypy of Newport (1845) (see Shelley 1997: 72–73).

Diagnosis: Plutoniumidae with 9–10 pairs of spiracles, on segments 3, 5, 7 (variable), 8, 10, 12, 14, 16, 18 and 20.

Assigned species: T. californiensis Chamberlin, 1902; T. chuanensis n. sp., T. erythrocephalus (C.L. Koch, 1847), T. phanus (Chamberlin, 1951), T. spinicaudus (Wood, 1862).

Discussion: The concept of Theatops employed here differs from previous diagnoses in allowing for variability with respect to the absence or presence of a spiracle on segment 7. All previously known species have nine pairs of spiracles but the morphological spectrum of Plutoniumidae is altered by the discovery of T. chuanensis, described below, that corresponds to Theatops in every respect apart from having an extra pair of spiracles, on segment 7. The presence or absence of a spiracle on segment 7 is a character that has traditionally been interpreted as a generic-level feature in Scolopendromorpha (Attems 1930) or even at higher taxonomic levels, such as subfamilies (Schileyko 1992). Examples of genera that are distinguished solely by this character include Scolopocryptops Newport, 1844 (without it) and Dinocryptops Crabill, 1953 (with the spiracle) in the Scolopocryptopidae; and Otostigmus Porat, 1876 (without it) and EthmostigmusPocock, 1898, and Rhysida Wood, 1862 (with the spiracle) in the Scolopendridae (Kraepelin 1903; Attems 1930). Following traditional taxonomic practice, Theatops chuanensis would be regarded as a separate, monotypic genus, diagnosed by the spiracle on segment 7. Here we advance arguments for not following this course.

Evidence is increasingly accumulating to indicate that the presence or absence of a spiracle on segment 7 is less reliable as a phylogenetic and taxonomic character than previously thought. In the case of Scolopocryptops versus Dinocryptops, the absence of a spiracle on segment 7 maps onto morphology-based cladograms as a shared primitive character and the group lacking the spiracle (Scolopocryptops) is paraphyletic with respect to the one that possesses it (Dinocryptops) (Edgecombe & Koch 2008; Koch et al.2009, 2010). The evolutionary lability of the spiracle on segment 7 in Scolopocryptopinae was noted by Crabill (1955), who reported an otherwise normal specimen of Scolopocryptops sexspinosus that had a small but evidently functional spiracle on segment 7 (Crabill based his interpretation of function on the presence of tracheae in segment 7 of the specimen). Crabill (1955: 134), however, interpreted character polarity opposite to that optimised on cladograms: he “assume[d] the presence of seventh somite spiracles to be primitive possessions”, and despite “the loss of seventh somite spiracles… as evidence of parallel evolution in this character”, he maintained their taxonomic value (“That this is a sound generic distinction cannot, at the present state of knowledge, be seriously doubted”). This confidence is undermined by the discovery that Scolopocryptops nigrimaculatus Song et al., 2004, may exceptionally show a small segment 7 spiracle on one side of the body only versus its (normal) absence on the other side (Song et al. 2004: fig. 3C).

With respect to non-monophyly at least, the situation is not dissimilar in Otostigminae. Apart from lacking the autapomorphies of other genera, Otostigmus is diagnosed only by a single character (absence of the segment 7 spiracle), and analyses of molecular data in progress retrieve both Otostigmus and Rhysida as polyphyletic groups. Although genera (or subfamilies: Schileyko 1992) distinguished by the presence or absence of a spiracle on segment 7 are easily identified and taxonomically convenient, in the case of Otostigminae they do not reflect phylogeny accurately. Lewis (2004) provided a possibly relevant observation from Otostigminae in Tanzania, wherein Otostigmus tanganjikus Verhoeff, 1941, and Rhysida intermediaAttems, 1910, are apparently distinguished by no character apart from the spiracle on segment 7; in these instances it might be anticipated that the similar species pairs are more closely related than either is to most species assigned to the same genus under the standard classification. In this case, the segment 7 spiracle may remain a reliable character at the species level, but likely not at the generic level.

In Plutoniumidae, the morphological analyses of Edgecombe & Koch (2008) and Koch et al. (2009, 2010) retrieved Theatops as a paraphyletic group, i.e., Plutonium is nested within Theatops. As in the previous examples in which spiracle distributions are the principal basis for diagnosing genera, paraphyly of one genus is not surprising because Theatops is diagnosed only by primitive characters (the primitive spiracle arrangement, contra Crabill’s inferred ancestral condition). A three-genus classification (Theatops: lacking

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spiracles on segment 7; a new genus for T. chuanensis with spiracles on segment 7; and Plutonium: with spiracles on segments 2–20) would increase paraphyly rather than lessen it, i.e., Theatops would be paraphyletic to two genera rather than one. We do not place Plutonium in synonymy under Theatops, which would eliminate non-monophyletic taxa from Plutoniumidae, but we do discard the option of erecting a monotypic genus for T. chuanensis.

Knowledge of Theatops chuanensis is limited to a single specimen and it can rightly be questioned whether its segment 7 spiracle could be an abnormality in a population or species that more commonly lacks a spiracle, by comparison to the examples cited above in which Scolopocryptops species only exceptionally have a spiracle on that segment. The likelihood that the segment 7 spiracle is an aberration in T. chuanensis is contradicted by the fact that the spiracles on segment 7 are as large as those on segment 8, are equally developed on both sides of the body, and the two segments have identical proportions of the stigmatopleurites. In contrast, in Scolopocryptops species that only rarely display a segment 7 spiracle, it is either conspicuously smaller than that on other segments (Crabill 1955) or both smaller than that on segment 8 and restricted to only one side of the body (Song et al. 2004).

Key to genera and species of Plutoniumidae

1. Spiracles on segments 2–20 ..................................................................................Plutonium zwierleini Cavanna, 1881- Spiracles on only nine or ten segments bearing the long tergites ................................................................................. 22. European species, occurring in the Iberian (Portugal, Spain) and Balkan (Croatia, Bosnia-Herzegovina, Montene-

gro) peninsulas ..................................................................................................................erythrocephalus (C.L. Koch)- Non-European species (A, USA and Mexico; C, China) ............................................................................................. 33. Ultimate prefemur with dorsal distomedial spur (Shelley 1997: figs. 7–9).................................................................. 4- Ultimate prefemur without dorsal distomedial spur (Shelley 1997: fig. 6) .................................................................. 54. Tergite of ultimate leg-bearing segment with complete longitudinal median suture dorsally (Shelley 1997: figs. 33–

34); A: Texas, USA .........................................................................................................................phanus Chamberlin - Tergite of ultimate leg-bearing segment without median suture or with only minute remnant anteriad (Shelley 1997:

figs. 7–9); A: eastern United States from northern Illinois and central Iowa to southwestern Arkansas, and from northwestern North Carolina, northeastern Tennessee and adjacent periphery of Virginia to central North and South Carolina, and eastcentral and northwestern Alabama ..................................................................... spinicaudus (Wood)

5. Spiracle present on segment 7; C: Sichuan, China............................................................................. .chuanensis n. sp- Spiracle absent on segment 7, American species ......................................................................................................... 66. Coxopleuron distinctly elongate and apically acuminate, usually with blackened terminal spur; ventral surfaces of

ultimate prefemora and femora usually with one spine each (four spines total) distal to midlength (Shelley 1997: figs. 13–17); A: Kern County, California, to Douglas County, Oregon, USA ..................... californiensis Chamberlin

- Coxopleuron at most only slightly prolonged, apically rounded, without blackened spurs; ventral surfaces of ulti-mate prefemora and femora with or without spines, rarely with one on each podomere, total almost always less than four (Shelley 1990: figs. 6–11); A: eastern United States from eastern Connecticut to eastern and southern coastal Texas, eastern Oklahoma, and south Florida Keys, and western United States and northwestern Mexico from west-ern Chihuahua, southern Sonora, and northern Baja California Norte to southwestern Utah, southern Nevada, and southern California.................................................................................................................................. .posticus (Say)

Theatops chuanensis n. sp.Figs. 2–19

Type specimens: Holotype: China: Sichuan Province, Nanping County, Jiuzhai Valley; leg. Zhiyong Di, viii.2009; MWHU (Ar.-MWHU-SCJZ0908).

Diagnosis: Prefemur and femur of ultimate leg lacking spines on ventral side; dorsal distomedial prefemoral spur absent; medial border of coxopleuron slightly prolonged posteriorly, acuminate, bearing a small spine at apex; pore field extending near to posterior margin of coxopleuron; 10 pairs of round spiracles, on segments 3, 5, 7, 8, 10, 12, 14, 16, 18 and 20.

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FIGURES 2–7. Theatops chuanensis n. sp. holotype. 2–3. Habitus of holotype (Ar.-MWHU-SCJZ0908), dorsal and ventral views. 4–5. Head and tergites 1–5, dorsal and ventral views. 6–7. Tergites 19–21 and ultimate legs, dorsal and ventral views. Scale bars of figs. 2–3: 10.0 mm. Scale bars of figs. 4–7: 5.0 mm.

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FIGURES 8–11. Theatops chuanensis n. sp. holotype. 8. Head and tergites 1–5. 9. Tergites 9–15. 10–11. Tergites 19–21 and ultimate legs, dorsal and ventral views. Scale bars: 5.0 mm.

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FIGURES 12–18. Theatops chuanensis n. sp. holotype. 12–13. Head and tergite 1 and 2, dorsal and ventral views. 14. Right lateral view of 7th and 8th segments. 15. Spiracle. 16–18. Legs 1 (left), 18 (right) and 20 (left). Scale bars (except 15): 2.5 mm, scale bar(15): 0.25 mm.

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Etymology: The specific epithet refers to the locality where the specimen was collected. The prefecture name “Chuan” is compounded with the suffix “ensis”.

Description: Length 76.6 mm (Figures 2–3).Cephalic plate, coxosternite and forcipules, T1–2, ultimate segment and ultimate legs pale orange (Figures

2–3). Antennae, T3–20, all sternites, and dorsal aspects of legs light yellow. Margin of coxosternal tooth plates, tibial and tarsal spurs, and distal parts of pretarsi red-brown. Ventral aspects of legs pale yellow with nearly white patches.

Tegument smooth. Cephalic plate rounded, about as long as wide, overlapping T1; two pale areas instead of lateral ocelli at base of antennae (Figures 4, 12); pair of unequal, longitudinal sutures on posterior part of cephalic plate, the longer less than half length of cephalic plate, terminating slightly anteriad of posterior margin (Figure 12). Antenna extending to posterior end of T5, with 17 articles, the basal four bearing very few scattered setae dorsally; 1st article sparsely setose ventrally, increasing to moderately setose by 3rd and densely setose at 5th article (Figure 13). Coxosternal tooth plate narrowing distad, with 1–2 teeth, where paired, situated near medial and lateral margins (Figure 13); trochanteroprefemoral process thorn-like.

T1 with anterior transverse sulcus and cruciform sutures (Figures 2, 8, 12); former straight for most of its length, forming obtuse angle at midline; anterior halves of cruciform sutures bissected by longitudinal median furrow continuing anterior to the anterior transverse sulcus. T2–20 with pair of complete paramedian sutures (Figure 9); T2–5 additionally with bilobate transverse suture between anterior part of paramedian sutures and oblique sutures across more than half of tergites lateral to paramedian sutures (Figure 8). T21 with longitudinal median suture terminating just short of caudal margin; length of tergite nearly twice width. Sternites 1–20 with round median depressions. S21 attenuated posteriorly, elongate subtriangular, lateral margins weakly concave, posteromedian margin with slight indentation (Figures 7, 11). Segments 3, 5, 7, 8, 10, 12, 14, 16, 18 and 20 with rounded spiracles (Figures 14–15); spiracle on segment 7 as large as that on segment 8. Each coxopleuron with ovoid field of several hundred small pores, pore field extending close to posterior margin; medial border of coxopleuron slightly prolonged posteriorly, acuminate, bearing a small red-brown spine at apex (Figure 11).

Two tibial spurs on legs 1–18, leg 19 damaged (tibia absent), leg 20 without tibial spurs (Figure 18). Legs 1–20 each with a tarsal spur near distal end of tarsus (Figures 14, 16, 17) and a pair of pretarsal accessory spurs. Ultimate legs forcipulate, prefemora subquadrate in cross-sections, ventral sides of prefemora and femora without spines (Figures 6–7, 10–11).

Ecology: Under stones in moist, warm mixed forest.Distribution: Known only from the type locality (Figure 19).Discussion: Based on the shared absence of a dorsal distomedial prefemoral spur on the ultimate leg and

the suppression of ventral spines on the prefemur and femur of that leg, T. chuanensis appears to be closely related to T. californiensis and T. posticus. The most pronounced difference between them is the presence (T. chuanensis) or absence (North American species) of the segment 7 spiracle. The holotype of T. chuanensiswholly lacks ventral spines on the prefemur and femur of the ultimate leg, a state variably shared with T. californiensis (Shelley 1997: fig. 17) and T. posticus (Shelley 1990: fig. 6), notably the eastern population of the latter, though both of the American species more often have a single spine on each prefemur and femur. T. chuanensis is further distinguished from T. posticus by the distinct prolongation of the coxopleuron and the presence of a spine at its apex, as well as by the closer approximation of the pore field to the posterior margin (compare Figure 11 and Shelley 1990: figs. 6–11). The pore field of T. chuanensis is longer than in any of four North American species of Theatops, and is most closely approached by that of T. erythrocephalus (Shelley 1990: fig. 12; Shelley 1997: figs. 40–41).

The discovery of Theatops in China demonstrates that Plutoniumidae is much more widely distributed geographically than had been thought to be the case. The (then reasonable) inference by Shelley (1997) that taxa in Asia are unlikely to belong to Plutoniumidae is no longer tenable on geographic grounds alone. It is likely that this family had a very widespread distribution that is now reduced to highly disjunct parts of the world (North America, the Mediterranean Basin, and China), pruned by extinction. The type locality of Theatops chuanensis lies near the limits of published myriapod localities in China mapped by Wang &

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Mauriès (1996). Their map showed vast areas of China that remained to be surveyed by myriapodologists, and they judged that probably only a small percentage of Chinese myriapod species had yet to be discovered and described. The completely unexpected discovery of a Chinese plutoniumid during a field excursion as recently as 2009 demonstrates the enormous potential for further surveys of previously unexplored or under-sampled parts of China to find species that are of global biogeographic significance.

FIGURE 19. Map of southwestern China, showing the type locality (red star) for Theatops chuanensis n. sp.

Acknowledgements

Our thanks are extended to Dr. Jun Hu for photographing the specimen, and to Dr. Lucio Bonato for providing the base map used in Fig. 1. Reviews by Drs. John Lewis and Rowland Shelley helped us to improve the manuscript. This work was supported in part by Grants from the National Natural Sciences Foundation of China (30900239)

References

Attems, C. (1910) Myriopoden von Madagaskar den Comoren und den Inseln Ostafrikas. In: Voeltzkov (ed.), Reise in Ostafrika, 3. Schweizerbart’sche E. Nägele & Dr. Sproessner, Stuttgart, pp.73–115.

Attems, C. (1926) Chilopoda. In: Kükenthal, W., Krumbach, T. (eds.), Handbuch der Zoologie, 4 (Myriapoda). Walter de Gruyter, Berlin, Leipzig, pp. 239–402.

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Attems, C. (1930) Myriapoda 2. Scolopendromorpha. In: Schulze, F. E., Kükenthal,W. (eds.), Das Tierreich, 54. Walter de Gruyter, Berlin, pp. 1–308.

Bollman, C.H. (1893) The Myriapoda of North America. Bulletin of the United States National Museum, 46, 1–210.Bonato, L., Edgecombe, G.D., Lewis, J.G.E., Minelli, A., Pereira, L.A., Shelley, R.M. & Zapparoli, M. (2010) A common

terminology for the external anatomy of centipedes (Chilopoda). Zookeys.Bücherl, W. (1941) Catálogo dos Quilópodos da zona neotrópica. Memórias do Instituto Butantan, 15, 251–372. Cavanna, F. (1881) Nuovo genere (Plutonium) e nuova specie (P. zwierleini) di Scolopendridi. Bullettino della Società

Entomologica Italiana, 13, 169–178.Chamberlin, R.V. (1902) A new genus and three new species of chilopods. Proceedings of the Academy of Natural

Sciences, Philadelphia, 54, 39–43. Chamberlin, R.V. (1951) A new species in the chilopod genus Theatops. Psyche, 58, 100–101. Crabill, R.E., Jr. (1955) On the reappearance of a possible ancestral characteristic in a modern chilopod (Chilopoda:

Scolopendromorpha: Cryptopidae). Bulletin of the Brooklyn Entomological Society, 50, 133–136.Crabill, R.E., Jr. (1960) A new American genus of cryptopid centipede, with an annotated key to the scolopendromorph

genera from America north of Mexico. Proceedings of the United States National Museum, 3422, 1–15.Edgecombe, G.D. & Bonato, L. (2010) Scolopendromorpha. In: Minelli, A. (ed.), Treatise on Zoology. Myriapoda:

Chilopoda. Brill, Leiden, pp. 392–407.Edgecombe, G.D. & Koch, M. (2008) Phylogeny of scolopendromorph centipedes (Chilopoda): morphological analysis

featuring characters from the peristomatic area. Cladistics, 24, 872–901. Edgecombe, G.D. & Koch, M. (2009) The contribution of preoral chamber and foregut morphology to the phylogenetics

of Scolopendromorpha (Chilopoda). Soil Organisms, 81, 295–318.Koch, C.L. (1847) System der Myriapoden. In: Herrich-Schäffer, L. (ed.), Kritische Revision der Insectenfauna

Deutschlands, 3. Pustet, Regensburg, pp. 1–270. Koch, M., Edgecombe, G.D. & Shelley, R.M. (2010) Anatomy of Ectonocryptoides (Scolopocryptopidae:

Ectonocryptopinae) and the phylogeny of blind Scolopendromorpha (Chilopoda). International Journal of Myriapodology, 3, 51–81.

Koch, M., Pärschke, S. & Edgecombe, G.D. (2009) Phylogenetic implications of gizzard morphology in scolopendromorph centipedes (Chilopoda). Zoologica Scripta, 38, 269–288.

Kraepelin, K. (1903) Revision der Scolopendriden. Mitteilungen aus dem Naturhistorischen Museum in Hamburg, 20, 1–276.

Lewis, J.G.E. (2004) Redescription and reassignment of the type material of the African centipedes Otostigmus reichardtiKraepelin and Otostigmus tanganjikus Verhoeff, with remarks on the genera Digitipes Attems, Otostigmus Porat and Rhysida Wood (Chilopoda: Scolopendromorpha). African Invertebrates, 45, 279–286.

Lewis, J.G.E. (2006) On the scolopendromorph centipede genus Mimops Kraepelin, 1903, with a description of a new family (Chilopoda: Scolopendromorpha). Journal of Natural History, 40, 1231–1239.

Lewis J.G.E., Edgecombe G.D. & Shelley R.M. (2005) A proposed standardised terminology for the external taxonomic characters of the Scolopendromorpha (Chilopoda). Fragmenta faunistica, 48, 1–8.

Newport, G. (1844) [Conclusion of the monograph on the Myriapoda Chilopoda]. Proceedings of the Linnean Society of London, 1, 191–197.

Newport, G. (1845) Monograph of the Class Myriapoda, Order Chilopoda; with observations on the general arrangement of the Articulata, Parts 1 and 2. Transactions of Linnean Society of London (1844), 19, 265–302, 349–439.

Pocock, R.I. (1888) On the genus Theatops. Annals and Magazine of Natural History, Series 6, 1, 283–290.Ribaut, H. (1915) Biospeologica XXXVI. Notostigmophora, Scolopendromorpha, Geophilomorpha (Myriapodes).

Première Série. Archives de Zoologie expérimentale et générale, 55, 323–346.Saussure, H. de & Humbert, A (1872) Études sur les myriapodes. In: Milne-Edwards, H. (ed.), Mission scientifique au

Mexique et dans l’Amérique Centrale, recherches zoologiques. Mémoires du Muséum national d’Histoire naturelle, Paris,�6, 2, 1–211.

Say, T. (1821) Descriptions of the Myriapodae of the United States. Journal of the Academy of Natural Sciences, Philadelphia, 2, 102–114.

Schileyko, A.A. (1992) Scolopenders of Viet-Nam and some aspects of the system of Scolopendromorpha (Chilopoda Epimorpha). Part 1. Arthropoda Selecta, 1, 5–19.

Schileyko, A.A. (1996) Some problems in the systematics of the Order Scolopendromorpha (Chilopoda). In: Geoffroy, J.-J., Mauriès, J.-P., Nguyen Duy-Jacquemin, M. (eds.), Acta Myriapodologica. Mémoires du Muséum national d’Histoire naturelle, Paris, 169, 293–297.

Schileyko, A.A. (2007) The scolopendromorph centipedes (Chilopoda) of Vietnam, with contributions to the faunas of Cambodia and Laos. Part 3. Arthropoda Selecta, 16, 71–95.

Schileyko, A.A. & Pavlinov, I.J. (1997) A cladistic analysis of the order Scolopendromorpha (Chilopoda). Entomologia Scandinavica, 51 (Supplement), 33–40.

Shelley, R.M. (1990) The centipede Theatops posticus (Say) (Scolopendromorpha: Cryptopidae) in the southwestern

DI ET AL.62 · Zootaxa 2667 © 2010 Magnolia Press

United States and Mexico. Canadian Journal of Zoology, 68, 2637–2644.Shelley, R.M. (1997) The Holarctic centipede subfamily Plutoniuminae (Chilopoda: Scolopendromorpha: Cryptopidae)

(nomen correctum ex subfamily Plutoniinae Bollman, 1893). Brimleyana, 24, 51–113.Shelley, R.M. (2002) A synopsis of the North American centipedes of the order Scolopendromorpha (Chilopoda).

Virginia Museum of Natural History Memoir, 5, 1–108.Shelley, R.M. & Backeljau, T.(1995) Case 2946. Plutoniinae Bollman, 1893 (Arthropoda: Chilopoda) and Plutoniinae

Cockerell, 1893 (Mollusca: Gastropoda): proposed removal of homonymy. Bulletin of Zoological Nomenclature, 52, 150–152.

Serra, A. (1983) Els Scolopendrinae i els Theatopsinae (Chilopoda: Scolopendromorpha) de la Península Ibèrica. Butlleti de la Institució Catalana d’Història Natural, 49, 77–83.

Song, Z.S., Song D.X. & Zhu, M.S. (2004) On a new species and a new record of the genus Scolopocryptops from China (Chilopoda: Scolopendromorpha: Scolopocryptopidae). Journal of Agricultural University of Hebei, 7, 80–85.

Verhoeff, K.W. (1906) Vergleichend-morphologische Studie über die coxopleuralen Körperteile der Chilopoden, mit besonderer Berücksichtigung der Scolopendromorpha, ein Beitrag zur Anatomie und Systematik derselben, nebst physiologischen und phylogenetischen Mitteilungen und Ausblicken auf die Insekten.� Nova Acta Leopoldina, Abhandlungen der Kaiserlich Leopoldinisch-Carolinisch Deutschen Akademie der Naturforscher, 86, 349–501.

Verhoeff, K.W. (1907) Systematik. In: Bronn, H.G. (ed.), Klassen und Ordnungen des Tierreichs, 5, Abteilung 2, Buch 1. Akademische Verlagsgesellschaft, Leipzig, pp. 217–264.

Verhoeff, K.W. (1941) Aliquid novi ex Africa 4. Zoologischer Anzeiger, 135, 196–204.Wang, D. & Mauriès, J.-P. (1996) Review and perspective of study on myriapodology of China. In: Geoffroy, J.-J.,

Mauriès, J.-P., Nguyen Duy-Jacquemin, M. (eds.), Acta Myriapodologica. Mémoires du Muséum national d’Histoire naturelle, Paris, 169, 81–99.

Wood, H.C. (1862) On the Chilopoda of North America, with a catalogue of all the specimens in the collection of the Smithsonian Institution. Journal of the Acadaemy of Natural Sciences, Philadelphia, Series 2, 5, 2–52.

Wood, H.C. (1865) The Myriapoda of North America. Transactions of the American Philosophical Society, 13, 137–248.Würmli, M. (1975) Contributo alla conoscenza del Plutonium zwierleini Cavanna, 1881 (Chilopoda:

Scolopendromorpha: Cryptoidae). Animalia, 2, 209–213.Zapparoli, M. (2009) An annotated catalogue of the epigeic and cave centipedes (Chilopoda) of Sardinia. Zootaxa, 2318,

56–168.

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