On some surface structures of potential taxonomic importance in families of the suborders Polydesmidea and Dalodesmidea (Polydesmida, Diplopoda).

Surface structures have rarely been the subject of a comprehensive study in Polydesmida despite their tremendous variety within this order. A number of these peripheral structures are here studied in most families of the suborders Polydesmidea and Dalodesmidea (sensu Hoffman 1980), using scanning electron microscopy. An illustrated description of the surface sculpture of the prozonite, the limbus and the intercalary cuticular micro-scutes on the metazonite is given for the first time for the studied families, together with an account of some other poorly known surface structures. Taken together, these characters allow us to recognize two main groupings of families. The families Ammodesmidae, Cryptodesmidae, Cyrtodesmidae, Haplodesmidae, Oniscodesmidae and Pyrgodesmidae have knobs on the posterior part of the prozonites, a toothed to lobed limbus, and no micro-scutes on the metazonites, wheras the families Fuhrmannodesmidae, Polydesmidae, Dalodesmidae, Macrosternodesmidae, Nearctodesmidae, Opisotretidae and Trichopolydesmidae have no knobs on the posterior part of the prozonites, a spiky or reduced limbus, and intercalary micro-scutes on the metazonites. The results are complemented with literature records and compared with current taxonomic and phylogenetic interpretations of the group.

Introduction

Whereas the gonopods have hitherto been acknowledged to be the most reliable source of characters for millipede identification, the details of external morphology have in most cases remained under-prospected. This is also true for the order Polydesmida although it is by far the most diverse millipede order in terms of non-gonopodal morphology. The taxonomy of several polydesmidan families, notably Fuhrmannodesmidae and Pyrgodesmidae, is in a deplorable state, and new taxonomic characters are badly needed. Rowe and Sierwald (2006) drew attention to the fact that the external morphology has been studied in only a few cases in millipedes, and gave an overview of the major works which dealt with this topic.

Scanning electron microscopy has, in many contemporary works, significantly helped to illustrate fine surface structures in millipedes. Mesibov (2009a) studied and described the several shapes of spiracles in species of the families Paradoxosomatidae and Dalodesmidae, demonstrated the great variation in the shape, location and density of the sphaerotrichomes on male legs in Dalodesmidae and noted the presence of different patterns in the integument sculpture of the metatergal tuberculation in two genera of the same family (see also Mesibov 2008). Moreover, the different arrangements and structures of the spinnerets in 16 families of Polydesmida were studied by Shear (2008), who delineated a notable variation of these structures within the studied taxa (see also Mesibov 2009a). The structure of the spinnerets was furthermore studied by Mesibov (2008, 2009a) who suggested a possible synapomorphy for some of the 10 studied dalodesmid genera even if recognizing that the use of spinneret structure in taxonomy of Polydesmida requires more sampling (see Mesibov 2009a).

The limbus, or posterior margin of the metazonites, was investigated by Schmidt (1962) who systematically described and compared the variation of shapes within numerous families, using only light microscopy.

Some polydesmidans of the families Cryptodesmidae, Haplodesmidae, and Pyrgodesmidae are earth-incrusted, i.e., adult specimens bear a coat of dirt. Shear (1973, 1977) described and illustrated special “boxes” and “branched tree-like setae” (Shear 1977) which supposedly keep the dirt on the cuticle.

During the study of a new pyrgodesmid species from Tunisia using scanning electron microscopy (Akkari and Enghoff 2011), we found a number of cuticular structures which have not hitherto been described. In order to assess their significance, we made a comprehensive survey of 22 species belonging to all the families of the suborders Polydesmidea and Dalodesmidea (see the list below and Table 1) except for the Dorsoporidae (Polydesmidea) and Vaalogonopodidae (Dalodesmidea) of which material was inaccessible for study.

Table 1.

The studied species and the states of the three main characters examined.

	Species	Knobs on the posterior surface of the prozonites	Limbus	Intercalary micro-scutes <br/> on metazonites
Suborder Polydesmidea
Ammodesmidae	Elassystremma sp.	+	lobes and spikes	‒
Cryptodesmidae	Aporodesmus sp.	‒	jagged lobes and spikes	‒
	Elythesmus enghoffi	+	jagged lobes and spikes	_
Cyrtodesmidae	cyrtodesmid gen. sp.	+	lobes and spikes	‒
Fuhrmannodesmidae	Fuhrmannodesmus lividus	–	reduced	+
	fuhrmannodesmid sp.	‒	reduced	+
	Gyrophallus sp.	‒	reduced	+
Haplodesmidae	Prosopodesmus jacobsoni	+	lobes	‒
Macrosternodesmidae	Ophiodesmus albonanus	‒	reduced	+
Nearctodesmidae	nearctodesmid sp.	‒	reduced	+
Oniscodesmidae	Amphitomeus attemsi	+	lobes and spikes	‒
Opisotretidae	Corypholophus sp.	‒	reduced	?+
	Solaenaulus butteli	–	reduced	+
Polydesmidae	Brachydesmus superus	‒	ramified spikes	+
	Propolydesmus laevidentatus	–	ramified spikes	+
Pyrgodesmidae	Cryptocorypha ornata	+	lobes	‒
	Cynedesmus sp.	+	lobes	–
	Rharodesmus tabarkensis	+	lobes	–
	Tonodesmus sp.	+	lobes	–
Trichopolydesmidae	Napocodesmus endogeus	‒	reduced	+
	trichopolydesmid sp.	–	reduced	+
Suborder Dalodesmidea
Dalodesmidae	Icosidesmus sp.	‒	ramified spikes	+

Our study was mainly focused on three sets of characters: a) surface sculpture of the prozonite (anterior, cylindrical part of body ring); b) the limbus (posterior margin of body rings); c) intercalary micro-scutes on the surface of the metazonite. A few additional structures such as the cuticular outgrowths in earth-incrusted species and the ozopores are briefly presented below and compared. Moreover, recent literature accountsPageBreak of 29 species from relevant families (see Table 2), including SEM illustrations or descriptions were checked and compared with our results.

Table 2.

Literature records

	Species	Reference
Suborder Polydesmidea
Ammodesmidae	Elassystremma laeve Vandespeigel and Golovatch, 2003. Elassystremma prolaeve VandenSpeigel and Golovatch, 2003	VandenSpeigel and Golovatch 2003
Cryptodesmidae	Aporodesmus gabonicus (Lucas, 1858)	Schmidt 1962
	Tarmadesmus azucarensis Kraus, 1959
Fuhrmannodesmidae	Fuhrmannodesmus carli Kraus, 1955	Schmidt 1962
	Salvadoria sagittalis Kraus, 1954
	Cutervodesmus similis Kraus, 1959
Haplodesmidae	Agathodesmus steeli Silvestri, 1910	Mesibov 2009b
	Cylindrodesmus hirsutus Pocock, 1889	Golovatch et al. 2001
	Eutrichodesmus armatocaudatus Golovatch et al., 2009	Golovatch et al. 2009a
	Eutrichodesmus basalis Golovatch et al., 2009
	Eutrichodesmus communicans Golovatch et al., 2009
	Eutrichodesmus inciues Golovatch et al., 2009
	Eutrichodesmus similis Golovatch et al., 2009
Oniscodesmidae	Oncodesmoides rectus Kraus, 1954	Schmidt 1962
Opisotretidae	Opisotretus kraepelini (Attems, 1907)	Schmidt 1962
Polydesmidae	Polydesmus complanatus (Linneaus, 1871)	Schmidt 1962
Pyrgodesmidae	Lobiferodesmus vanuatu Golovatch, et al., 2008	Golovatch et al. 2008
	Poratia digitata (Porat, 1889)	Adis et al. 2000
	Muyudesmus obliteratus Kraus, 1960
	Cryptocorypha hoffmani Golovatch et al. 2011	Golovatch et al. 2011
	Myrmecodesmus hastatus (Schubart, 1945)	Bergholz et al. 2004
	Monachodesmus albus Kraus, 1958	Schmidt 1962
Suborder Dalodesmidea
Dalodesmidae	Ginglymodesmus tasmanianus Mesibov 2005	Mesibov 2009c
Not assigned to any family	Asphalidesmus bellendenkerensis Mesibov, 2011	Mesibov 2011
	Asphalidesmus golovatchi Mesibov, 2009
	Noteremus infimus Mesibov, 2009	Mesibov 2009c
	Noteremus summus Mesibov, 2009
	Procophorella innupta Mesibov, 2003	Mesibov 2003

Materials and methods

The studied material is preserved in 70 % ethanol and deposited in the Natural History Museum of Denmark (Zoological Museum, University of Copenhagen, ZMUC). Earth-incrusted specimens were cleaned for scanning electron microscopy by soaking in a solution of commercial detergent (©Biotex) and/or by ultrasound then air dried. SEM pictures were made with a JEOL JSM-6335F scanning electron microscope, then processed and assembled with Adobe Photoshop CS5 software.

When not otherwise indicated, we have followed the classification proposed by Hoffman (1980) as updated by Shelley (2003).

Studied specimens

Suborder

Ammodesmidae Cook, 1896

sp., Tanzania, Unzungwa Mts, Iringa Region, Uzungwa Scarp Forest Res., above Chita village, 1600–1650m, 8–13.ix.1984, pitfall Traps in Montane Rain Forest, N. Scharff leg. (ZMUC 00020487).

Cryptodesmidae Karsch, 1879

sp., female, Cameroun, Northwest Province, Menchum Div. Near L. Oku forest, in litter, ca. 2150m, N6°12', E10°27', 7–13.ii.1992, C. Griswold, S. Larcher, N. Scharff and C. Wanzie leg. (ZMUC 00020478).

Hoffman, 1978, female, Tanzania, W. Usambara Mts, Mazumbai Forest Reserve, 19–29.ix.1992, M. Andersen leg. (ZMUC 00020477).

Cyrtodesmidae Cook, 1896

Cyrtodesmid sp., female, Colombia, Páramo de Sumapaz, soil under grasses, 3600m, 5.x.1978, H. Sturm leg. (ZMUC 00020494).

Fuhrmannodesmidae Brölemann, 1916

Carl, 1914, male, Colombia, Par de Monserrate, near Bogotá, 3250m, dead leaves of , 18.iv.1969, H. Sturm (ZMUC 00020483).

fuhrmannodesmid sp. (Arndt et al. 2008), female, Spain, Canary Islands, La Palma, Pared Vieja, 21.ii.-5.iii.2002, E. Arndt leg. (ZMUC 00020492).

sp., female, Colombia, 1991, H. Sturm leg. (ZMUC 00020484).

Haplodesmidae Cook, 1895

Silvestri, 1910, female, Fiji Isl, Viti Levu Suva, in garden, 2–3.ix.1995, A van Hart leg. (ZMUC 00020476).

Macrosternodesmidae Brölemann, 1916

(Latzel, 1895), male, Denmark, NE Zaland, Copenhagen, Utterslev Mose, 22.iv.1973, H. Enghoff leg. (ZMUC 00020491).

Nearctodesmidae Chamberlin and Hoffman, 1950

nearctodesmid sp., male, Calif. Humboldt Co. Jolly Giant Canyon, 300–650m, Arcata Comm. For 13.i.1979, A.K. Johnson, R. M. Shelley leg. (ZMUC 00020482).

Oniscodesmidae de Saussure, 1860

(Schubart, 1934), female, Dania: Nez UB47, Copenhagen, Botanical garden, væksthus, 16.iv.1986, H. Enghoff and Z. Korsòs leg. (ZMUC 0002046).

Opisotretidae Hoffman, 1980

sp., female, Thailand, Chieng Mai Province, Doi Inthanon N. P., Mae Ya, 6–700m (ZMUC 00020479).

(Carl, 1922), male, Fiji Isl. Viti Levu Suva, in garden, 2–3.ix.1995, A van Harten leg. (ZMUC 00020480).

Polydesmidae Leach, 1815

(Latzel, 1884), female, Tunisia, NW, Jendouba Governorate, 9km of Hammam Bourguiba (West of Aïn Draham), N36°48.046, E08°39.544, 379m, Pine Forest, close to river, under stones, logs and leaf litter, 22.iii.2008, P. Stoev and N. Akkari leg. (ZMUC 00020496).

(Loksa, 1967), male, Madeira, Faja da Nogueria, N. Side, ca. 800m. a.s.l. with , 20.xi.1980, H. Enghoff and O. Martin leg. (ZMUC 00020475).

Pyrgodesmidae Cook, 1895

(Attems, 1938), unsexed fragment, Hawaii Isl., Kauai, in a grotto, moss and , 20.x.1962, M. Hammer leg. (ZMUC 00020493)

sp., female, La Gomera, Valle Gran Rey, litter, in banana plantation, 10m, 2.xii.1987, A. Fjellberg leg. (ZMUC 00020485).

Akkari and Enghoff, 2011, male, Tunisia, NW, Jendouba Governorate, Tabarka, N36°58'10.5", E8°45'35.6", alt. < 40m, coastal slope below the Genoese fort, under stones, 9.iii.2009, N. Akkari and H. Enghoff leg. (ZMUC 00020532).

sp., male, Spain, Almeria Sima terminal, T. M. Eidijo, 28.v.2000, M. Piquer and J.G. Pedro leg. (ZMUC-00020495).

Trichopolydesmidae Verhoeff, 1910

Ceuca, 1974, female, Moldavian SSR, Tiraspol, deep in orchard soil, 1985, S. Golovatch leg. (ZMUC 00020481).

trichopolydesmid sp. Slovakia, Slovak Karst, Ardouská Cave A-04-47, 5.x.2004, A. Mock leg. (ZMUC 00020490).

Suborder

Dalodesmidae Cook, 1896

sp., male, New Zealand, South Isl, Christchurch Banks Peninsula, Hinewai Reserve, Big Kanuka Trail, 3 iii.2010, S43°48'38.0", E173°01'15.6", 508m, sifted leaf litter and mosses, N. Scharff and G. Hormiga leg. (ZMUC 00020488).

Results

Fine sculpture of the prozonite

Figs 1–9

Figures 1–6.

Fine sculpture of the prozonite in the families Ammodesmidae, Cryptodesmidae, Cyrtodesmidae, Haplodesmidae, Oniscodesmidae and Pyrgodesmidae 1 sp. 2 , 3 cyrtodesmid sp. 4 5 6 . Abbreviations: a anterior part of the prozonite, b posterior part of the prozonite, r ridge, s spherical knobs.

Figures 7–9.

Fine sculpture of the prozonite in pyrgodesmid species. 7 sp. 8 sp. 9 . Abbreviations: a anterior part of the prozonite, b posterior part of the prozonite, r ridge, s spherical knobs.

The prozonite of the studied species is divided into two main parts separated by a transverse ridge. While the anterior part is rather uniform, showing a scaly aspect, the posterior surface displays varied patterns within the studied families.

In the examined species of Ammodesmidae, Cryptodesmidae, Cyrtodesmidae, Haplodesmidae, Oniscodesmidae and Pyrgodesmidae the anterior part of the prozonite (a) displays a covering of small scaly units, juxtaposed in series of irregular transverse rows, becoming elongated in the posteriormost row to form a transverse ridge (r) marking the border of this part (Figs 1–9). In front of the ridge, the general aspect is quite uniform in all the above cited families, with lozenge-shaped units. These units could sometimes be star-like, furrowed, and marginally jagged (cyrtodesmid sp.) (Fig. 3). In (Oniscodesmidae), the units are more elongated and strongly prominent, interconnected with parallel cuticular ridges (Fig. 5). On the other hand, the microsculpture of the anterior part of the prozonite in (Haplodesmidae) takes the shape of hollow chambers separated by walls of “microvilli-like” structures (Fig. 4). The posterior surface of the prozonite (b) is, in all examined species of these six families, characterized by a regular covering of sub-spherical knobs (s) placed on a smooth to microgranulated background. The cover of knobs is regularly dense in most cases (Figs 1, 3–9)though fairly dispersed in (Cryptodesmidae) (Fig. 2). These knobs are generally uniformly smoothPageBreak (Figs 1–3, 6–9) but sometimes exhibit special configurations: lobed in and spiky in (Figs 4, 5).

The examined species of the families Fuhrmannodesmidae, Polydesmidae, Dalodesmidae, Macrosternodesmidae, Nearctodesmidae, Opisotretidae and Trichopolydesmidae show an anterior surface of the prozonite with polygonal units serrated marginally. However, these units are much less conspicuous and prominent than in the species of the first set of families (Figs 10–17). The posterior border of the anterior part of the prozonite is similarly marked by a transverse ridge but its units are nevertheless only slightly modified. The posterior surface of the prozonite remarkably differs by the complete absence of the sub-spherical knobs described above; instead the surface is smooth to scaly (Figs 10–17).

Figures 10–13.

Fine sculpture of the prozonite in 10 (Fuhrmannodesmidae) 11 sp. (Fuhrmannodesmidae) 12 (Polydesmidae) 13 sp. (Dalodesmidae). Abbreviations: a anterior part of the prozonite, b posterior part of the prozonite, r ridge.

Figures 14–17.

Fine sculpture of the prozonite in 14 (Macrosternodesmidae) 15 nearctodesmid sp. (Nearctodesmidae) 16 (Opisotretidae) 17 (Trichopolydesmidae). Abbreviations: a anterior part of the prozonite, b posterior part of the prozonite, r ridge.

The limbus

Figs 18–34

Figures 18–22.

The structure of the limbus in 18 (Haplodesmidae) 19 (Pyrgodesmidae) 20 sp. (Pyrgodesmidae) 21 sp. (Pyrgodesmidae) 22 (Pyrgodesmidae). Abbreviations: L limbus, lo lobe, pa palette-like lobe, sp spike, t tooth-like lobe.

Figures 23–26.

The structure of the limbus in 23 (Oniscodesmidae) 24 cyrtodesmid sp. (Cyrtodesmidae) 25 sp. (Ammodesmidae) 26 (Cryptodesmidae). Abbreviations: L limbus, lo lobe, sp spike, pa: palette-like lobe.

Figures 27–30.

The structure of the limbus in 27 sp. (Dalodesmidae), 28 (Macrosternodesmidae) 29 (Polydesmidae) 30 (Polydesmidae). Abbreviations: L limbus, rs ramified spike.

The limbus (L) displays three major patterns of shapes in the studied families:

1) A regular set of rounded lobes (lo) placed in one row as in (Haplodesmidae), (Pyrgodesmidae) (Figs 18, 19) or two superposed rows of 'palette-shaped' lobes (pa) separated by spikes (sp) as in sp. (Pyrgodesmidae) (Fig. 20) although in some species of the latter family (e.g. sp. and ) the lobes are more tooth-like (t) (Figs 21, 22). The lobes are also surmounted by fine spikes as in (Oniscodesmidae), cyrtodesmid sp. (Figs 23, 24), sp. (Ammodesmidae) in which the lobes are moreover stocky or ‘palette-like’(pa) and serrated (Fig. 25) and in (Cryptodesmidae) where the spikes are more elongate and inserted between the jagged lobes (Fig. 26).

2) A series of ramified spikes (rs) in sp. (Dalodesmidae) (Fig. 27) or “icicles” in (Macrosternodesmidae), and (Polydesmidae) (Figs 28–30).

3) Reduced. In (Opisotretidae), (Trichopolydesmidae) and (Fuhrmannodesmidae) the limbus is hardly developed, taking the shape of a regular edge bearing a few scattered bulges (bu) which could be isolated or grouped, e.g. sets of three bulges in furhmannodesmid sp. (Figs 31–34).

Figures 31–34.

The structure of the limbus 31 (Opisotretidae) 32 (Trichopolydesmidae), 33 (Fuhrmannodesmidae) 34 fuhrmannodesmid sp. (Fuhrmannodesmidae). Abbreviations: bu bulges, L limbus.

A peculiar structure of the metazonites: intercalary cuticular micro-scutes

Figs 35–41

Figures 35–41.

Intercalary micro-scutes on the metazonitesof 35 (Fuhrmannodesmidae) 36 (Polydesmidae) 37 sp. (Dalodesmidae) 38 (Macrosternodesmidae) 39 nearctodesmid sp. (Nearctodesmidae) 40 (Opisotretidae) 41 (Trichopolydesmidae). Arrows point to the micro-scutes.

The studied species of the families Fuhrmannodesmidae, Polydesmidae, Dalodesmidae, Macrosternodesmidae, Nearctodesmidae, Opisotretidae and Trichopolydesmidae (cf. Table 1) present a peculiar structure on the metazonital surface: between the normal polygonal cuticular scutes which cover the metazonital surface there are rows of small ovoid “intercalary scutes”. The placement of these structures is unlikely to be accidental or indicating any kind of bacterial infection as they seem well arranged in a regular pattern, appearing like spaced nodes or pearls aligned on strings crossing the surface of the metazonites (Figs 35–41). These structures have never been documented. However, they are visible on an illustration in Mesibov [2003, fig. 3, (right)], for Mesibov, 2003 (Dalodesmidea).

Some other poorly known surface structures

Figs 42–49

Figures 42–43.

Cuticular outgrowths 42 (Pyrgodesmidae) 43 sp. (Ammodesmidae).

Figures 44–49.

Ozopores 44 (Pyrgodesmidae) 45 sp. (Pyrgodesmidae) 46 sp. (Ammodesmidae) 47 (Polydesmidae) 48 sp. (Fuhrmannodesmidae) 49 sp. (Opisotretidae).

The surface structure of most earth-incrusted species is characterized by the presence of papilla-like cuticular outgrowths which are particularly abundant in Ammodesmidae, Cyrtodesmidae and Pyrgodesmidae. These papillae are boletiform (mushroom-shaped) PageBreakPageBreakand are presumed to keep the cuticular secretions and adhering soil particles in place. Displaying variation in length, shape of the apex, and density on the surface, the papillae are short, with a rounded apex in (Fig. 42) and sp., elongate and slender in sp. (Fig. 43) and in cyrtodesmid sp. The same structures have been mentioned in previous works, generally quoted as “papillis” (Silvestri 1925, 1947), “Papillen” (Attems 1940), or “microvilli” in most of Golovatch’s works ‒ illustrations can be found in Golovatch et al. (2009a, figs 24A, B, E, F). Shear (1977) was the first to provide SEM illustrations of such structures and to comment on their possible function.

The ozopores (defense gland openings) display a notable variation within the examined families. In (Haplodesmidae), sp., , sp. (Pyrgodesmidae) and sp. (Ammodesmidae), they open on small rounded discs, situated on the surface of the paratergites or on porosteles. The discs are of variable thickness and diameter, bear an apparent internal closing mechanism and are externally bordered with several whorls of papillae (Figs 44–46). On the other hand, the ozopores appear as simple sub-circular openings on the surface of the paratergites in sp. (Dalodesmidae), (Polydesmidae), and sp. (Fuhrmannodesmidae), and sp. (Opisotretidae) (Figs 47–49).

Discussion

Polydesmida is the most speciose millipede order, and despite the fact that it has remained quite stable in terms of number of families (Shelley 2003), its taxonomy is far from being satisfactory. In his attempt to classify the suborders Polydesmidea and Dalodesmidea, Hoffman (1980: 146) expressed his dissatisfaction and pessimism: “The groupings set forth in the following pages are to a large extent exercises in futility, but may have some reference value in a bibliographic sense”. Nevertheless, Hoffman’s (1980) main classification scheme still stands and has been adopted by most subsequent authors. Simonsen (1990), studying the phylogeny of Polydesmida, made a number of changes and synonymies. Notably, Simonsen (1990) placed Dalodesmidae + Vaalogonpodidae as sister-group of Polydesmidae, i.e., the suborder Dalodesmidea nested within the suborder Polydesmidea. However, most of Simonsen’s conclusions were soon after criticized by several authors because they were based on insufficient data and bold assumptions (e.g. Golovatch 1996, Shelley 2003).

Taken together, the fine sculpture of the prozonite, the structure of the limbus and the presence/absence of metazonital micro-scutes indicate two main groupings of families within the suborders Polydesmidea and Dalodesmidea. The first group (A) comprises the families Ammodesmidae, Cryptodesmidae, Cyrtodesmidae, Haplodesmidae, Oniscodesmidae and Pyrgodesmidae. These six families have a cover of sub-spherical knobs on the posterior surface of the prozonites (Figs 1–9) and a toothedPageBreak limbus constituted of a series of lobes which may or not be surmounted by spikes (Figs 18–26).

The second group (B) encompasses the families Fuhrmannodesmidae, Polydesmidae, Dalodesmidae Macrosternodesmidae, Nearctodesmidae, Opisotretidae and Trichopolydesmidae, and is characterized by 1) an absence of knobs on the posterior surface of the prozonite which is smooth to scaly (Figs 10–17), 2) an absence of lobes on the limbus which has instead a jagged margin or spikes with varied complexity (Figs 27–34), and 3) intercalary micro-scutes on the surface of the metazonites (Figs 35–41), absent in the first set of families.

An assessment of several recent species descriptions and SEM illustrations shows in most cases similar structural arrangements of the prozonites. The presence of sub-spherical knobs on the posterior surface of the prozonite has been verified in a few additional genera and species of Pyrgodesmidae, such as (Golovatch et al. 2008, fig. 4C), (Adis et al. 2000, figs 25, 26, 27; Golovatch and Sierwald 2000, figs 1, 6), (= Muyudesmus) obliterata (Adis et al. 2000, fig. 29) and (Golovatch et al. 2011, fig. 38). Moreover, comparable structures are seen in the ammodesmids and (VandenSpeigel and Golovatch 2003, figs 7, 9, 18) and for the haplodesmid (Mesibov 2009b, figs 4B, 5C, 6C). However, we noticed some differences in (Golovatch et al. 2009a, figs 1C, 3C), (Golovatch et al 2009a fig. 6F), (Golovatch et al. 2009a fig. 11F) and (Golovatch et al 2009a, fig. 22E) although low image resolution doesn’t allow us to draw any conclusion about these species, neither about those cited in Golovatch et al. (2009b).

The fine structure of the prozonite in both species of the Tasmanian dalodesmid genus , and (Mesibov 2009c, fig. 3) perfectly agrees with what we recorded in the dalodesmid sp. (Fig. 13). The genus , in contrast, exhibits a pyrgodesmid-like pattern with a conspicuous cover of sub-spherical knobs on the posterior surface of the prozonite. In his description of , Mesibov (2009c) wrote “prozonites with narrow band of longitudinal ridges just anterior to suture, elsewhere uniformly covered with very small protuberances with blunt, rounded tips directed slightly posteriorly”. Mesibov (2011) did later describe several new species of the same genus on which the posterior prozonite protuberances were visible, especially on Mesibov, 2011 (Mesibov 2011, fig. 4A).

Although assigned to Dalodesmidea (Mesibov 2009c), the similarity of the genus to the first set of families (Ammodesmidae, Cryptodesmidae, Cyrtodesmidae, Haplodesmidae, Oniscodesmidae and Pyrgodesmidae) might not be a surprise knowing that the taxonomic position of has been matter of controversy. In fact, the genuswas originally described in Dalodesmidae then subsequently placed in Fontariidae, Vanhoeffeniidae, and Haplodesmidae, listed in Xystodesmidae, and assigned to Cryptodesmidae and Oniscodesmidae (see Mesibov 2009c). Despite PageBreaklacking sphaerotrichomes and showing a few further peripheral structures resembling more Pyrgodesmidae than Dalodesmidae and Vaalogonopodidae, was informally placed in the suborder Dalodesmidea (see Golovatch 2003). This was subsequently confirmed by Mesibov (2009c) although both authors agree in not assigning the genus to any family (Golovatch 2003, Mesibov 2009c).

Several literature records clearly support the patterns described above for the limbus. For example, the limbus shows a series of lobes with or without additional spikes in the pyrgodesmid species (Golovatch et al. 2008, fig. 4D) and (Bergholz et al. 2004, figs 5, 6), the ammodesmids and (VandenSpeigel and Golovatch 2003, figs 7, 18), and the haplodesmids (Mesibov 2009b, fig. 6C), (Golovatch et al. 2001, fig. 11) and (Golovatch et al. 2009a, fig. 19E). The limbus is, on the other hand, composed of a number of ramified spikes in, for example, the dalodesmid (Mesibov 2005, fig. 4A). Schmidt (1962, figs 22–29) provided detailed descriptions and drawings of the limbus for a number of Polydesmida species viz. (Oniscodesmidae), (Polydesmidae), (Opisotretidae), , (Pyrgodesmidae), (Cryptodesmidae), , , and (Fuhrmannodesmidae). In all these species, the limbus is strikingly similar to what we observed in the studied species belonging to the same families.

Though we are aware that the present data alone do not warrant a strict cladistic analysis, we have compared our findings (Fig. 51) with the only existing phylogenetic work on the order Polydesmida (Simonsen 1990). The two main groups (A) and (B) mentioned above agree remarkably well with the basal dichotomy in Simonsen’ cladogram (Fig 50), except for the families Haplodesmidae and Cryptodesmidae which belong to our group (A) but which according to Simonsen are in the clade which otherwise includes our group (B). However, the general knowledge about these two families and their diversity is still incomplete, and no evidence has hitherto been provided in support of their monophyly. In his revision of the Haplodesmidae, Golovatch (2009a) recognized two major ‘grades’, the haplodesmid and doratodesmid grades, according to their somatic (non-sexual) characters and capacity for “volvation” (coiling into a sphere). In the present study, only the haplodesmid genus was studied. Golovatch et al. (2009a) characterised as a “pyrgodesmid-like haplodesmid” and considered it as basal in the Haplodesmidae (together with Miyosi, 1958).

Figures 50–51.

50 Family-level cladogram of suborders Polydesmidea + Dalodesmidea according to Simonsen (1990). Haplodesmidae here corresponds to Haplodesmidae + Doratodesmidae on Simonsen’s original cladogram; families not studied here are marked with asterisks 51 Branching diagrams (not cladograms) based on Fig. 50 but modified to illustrate the distribution of the different states of the three studied characters: A. presence of knobs on the anterior part of the prozonite, B. shape of the limbus, C. presence of intercalary micro-scutes on the metazonites (see Appendix for character states).

The Cryptodesmidae studied by us alos present a complication as has knobs on the posterior part of the prozonites (Fig. 2) and a lobed limbus (Fig. 26) as in families group A, whereas has a posterior prozonite surface free of knobs (Fig. 52) and dentate leaf-shaped (le) elements and spikes (sp) on the limbus (Fig. 53).

Figures 52–53.

Prozonite and limbus in sp. (Cryptodesmidae) 52 Prozonite fine sculpture 53 The limbus. Abbreviations: a anterior part of the prozonite, b posterior part of the prozonite, L limbus, le dentate leaf-shaped element of the limbus, r ridge, sp spike.

As stated above, the second set of families (B) have no lobes on the limbus. Instead, ramified spikes and “icicles” (Figs 27–30) can be observed in the families Dalodesmidae, Macrosternodesmidae and Polydesmidae, which clearly separatesPageBreakPageBreak them from the Fuhrmannodesmidae, Trichopolydesmidae, Nearctodesmidae and Opisotretidae in which the limbus is hardly developed and bears but few scattered bulges (Figs 31–34). However, this separation fits neither with Hoffman’s (1980) classification in which the Macrosternodesmidae, Nearctodesmidae, Trichopolydesmidae and Fuhrmannodesmidae form the superfamily Trichopolydesmoidea (see also Shelley 2003), nor with the phylogenetic analysis of Simonsen (1990) in which the Macrosternodesmidae and Nearctodesmidae appear in the same clade while the Opisotretidae, a separate clade, is grouped with a different set of families (Fig. 50). The simple limbus could well be a plesiomorphic character state expressed in the Fuhrmannodesmidae, Trichopolydesmidae, Nearctodesmidae and Opisotretidae (see also Simonsen 1990).

In the present work, we do not pretend to offer a new subordinal classification of Polydesmida, or to solve any of the taxonomic problems related to families – a colossal task which definitely will require a lot more sampling and character scoring, including the gonopod structures which haven’t been considered here. However, documenting these remarkable surface structures and trying to compare them between the different families will perhaps contribute to bringing new insights, leading towards a better understanding of polydesmidean and dalodesmidean relationships.

Character matrix

Only the species studied by us have been coded.

	Char. no.	1	2	3
Suborder Polydesmidea Pocock, 1887
Ammodesmidae Cook, 1896	Elassystremma sp.	1	0	0
Cryptodesmidae Karsch, 1879	Elythesmus enghoffi Hoffman, 1978	1	0	0
	Aporodesmus sp.	0	0	0
Cyrtodesmidae Cook, 1896	cyrtodesmid sp.	1	0	0
Fuhrmannodesmidae Brölemann, 1916	Fuhrmannodesmus lividus Carl, 1914	0	2	1
	fuhrmannodesmid sp.	0	2	1
	Gyrophallus sp.	0	2	1
Haplodesmidae Cook, 1895	Prosopodesmus jacobsoni Silvestri, 1910	1	0	0
Macrosternodesmidae Brölemann, 1916	Ophiodesmus albonanus (Latzel, 1895)	0	1	1
Nearctodesmidae Chamberlin and Hoffman, 1950	nearctodesmid sp.	0	2	1
Oniscodesmidae de Saussure, 1860	Amphitomeus attemsi (Schubart, 1934)	1	0	0
Opisotretidae Hoffman, 1980	Corypholophus sp.	0	2	?+
	Solaenaulus butteli (Carl, 1922)	0	2	+
Polydesmidae Leach, 1815	Brachydesmus superus (Latzel, 1884)	0	1	1
	Propolydesmus laevidentatus (Loksa, 1967)	0	1	1
Pyrgodesmidae Cook, 1895	Rharodesmus tabarkensis Akkari and Enghoff, 2011	1	0	0
	Tonodesmus sp.	1	0	0
	Cynedesmus sp.	1	0	0
	Cryptocorypha ornata (Attems, 1938)	1	0	0
Trichopolydesmidae Verhoeff, 1910	trichopolydesmid. sp.	0	2	1
	Napocodesmus endogeus Ceuca, 1974	0	2	1
Suborder Dalodesmidea Hoffman, 1980
Dalodesmidae Cook, 1896	Icosidesmus sp.	0	1	1

Char. no.→	1	2	3
Ammodesmidae	1	0	0
Cryptodesmidae	1	0	0
Cyrtodesmidae	1	0	0
Fuhrmannodesmidae	0	2	1
Haplodesmidae	1	0	0
Macrosternodesmidae	0	1	1
Nearctodesmidae	0	2	1
Oniscodesmidae	1	0	0
Opisotretidae	0	2	1
Polydesmidae	0	1	1
Pyrgodesmidae	1	0	0
Trichopolydesmidae	0	2	1
Dalodesmidae	0	1	1