The millipede order Julida (Myriapoda: Diplopoda) in - Senckenberg

The millipede order Julida (Myriapoda: Diplopoda) in Tunisia, with

an overview of the North African species

Nesrine Akkari1*, Pavel Stoev2, Henrik Enghoff3 & Said Nouira1

1Research Unit of Biodiversity and Biology of Populations, Institut Supérieur des Sciences Biologiques Appliquées de

Tunis, 9 Avenue Dr. Zouheir Essafi, La Rabta, 1007 Tunis, Tunisia; e-mail: [email protected]

2 National Museum of Natural History, Tsar Osvoboditel Blvd. 1, 1000 Sofia, Bulgaria

3 Natural History Museum of Denmark, University of Copenhagen, 15, DK-2100 Copenhagen, Denmark

*Corresponding author

Abstract

An overview of the millipedes of the order Julida in Tunisia based on both literature data and newmaterial is provided. Fifteen species from two families and five genera are presently known, of whichOmmatoiulus aumalensis (Brolemann, 1925) and Brachyiulus pusillus (Leach, 1814) are new. recordedfor the country. All old records of Cylindroiulus distinctus (Lucas, 1846) from Tunisia are probablyerroneous and refer to either C. algerinus (Brölemann, 1897) or C. attemsi Read, 2005. Notes on thespecies distribution are given along with geographical maps and comments on their habitat preferences.Ommatoiulus punicus (Brölemann, 1894) is the most euryecious species of all Tunisian julidans. B.

pusillus is the sole member of the order occurring in the oases in the south, as its presence there iscertainly due to human introduction. An illustrated dichotomous key for identification of the Tunisianspecies is provided. An annotated checklist of North African Julida comprising 58 species and subspeciesfrom 12 genera and three families, as well as a historical overview of their exploration supplements thisstudy. Ommatoiulus punicus, O. malleatus Akkari & Voigtländer, 2007 and Brachyiulus stuxbergi

(Fanzago, 1875) are herewith reported for the first time from Algeria, while Proteroiulus hispanicus

Schubart, 1959, which was hitherto known only from Spain, is now reported from Morocco. A concisebiogeographic analysis outlines the similarities in the distribution of some species at regional andcontinental scales.

Keywords: millipedes, taxonomy, Julida, Tunisia, Morocco, Algeria, distribution patterns,habitats

1. Introduction and historical account

Although better known than Subsaharan Africa, the millipede fauna of North Africa is stillunsatisfactorily studied. Contemporary taxonomic reviews, revisions or catalogues aregenerally lacking, and the bulk of the information is still scattered through several oldfaunistic or taxonomic papers. The only exceptions are the contemporary revisions ofCylindroiulus distinctus group (Read 2005) and genus Glomeris Latreille, 1802 in NorthAfrica (Golovatch et al. 2009).

SOIL ORGANISMS Volume 81 (3) 2009 pp. 453–488

ISSN: 1864 - 6417

The earliest record of a julid millipede from North Africa was that of Julus varius Fabricius,1781 (now Pachyiulus varius) which Brandt (1841) recorded from Algeria. In this paperBrandt noted some differences between the Sicilian and the Algerian specimens, which raisesdoubts about the actual occurrence of P. varius in Africa. The study of the Algerian Julidacontinued later with Lucas (1846) who described 6 new species from Algeria: Iulus lapidarius

(now Ommatoiulus lapidarius) from western Algeria; Iulus affinis (now considered asynonym of Ommatoiulus fuscounilineatus from Alger, Philippeville (now Skikda),Constantine, Bône (now Annaba) and La Calle; Iulus fusco-unilineatus (now Ommatoiulus

fuscounilineatus) without exact localities but specifying that it is a common species inAlgeria; Iulus distinctus (now Cylindroiulus distinctus) from eastern Algeria, Alger,Philippeville, Bône, Constantine and La Calle; Iulus corticalis (probably a species ofBlaniulidae) from Philippeville and Blaniulus fuscopunctatus from ‘Ruines d’Hippône’ (inAnnaba). The taxonomic position of latter two remains uncertain since both were describedonly from females and juveniles (Brölemann 1897).

Studying the millipedes collected by Dr Anderson in Algeria and Tunisia, Pocock (1892)recorded two species already known from Algeria: Ommatoiulus fuscounilineatus andCylindroiulus distinctus. Brölemann (1897) described two further Cylindroiulus species: C.

algerinus (as Iulus (Phalloiulus) algerinus) and C. truncorum (as Iulus (Anoploiulus)africanus). Subsequently, Giard (1899) described Blaniulus drahoni (now Archichoneiulus

drahoni) from Ifri Semedane Cave. Verhoeff (1900) described Pachyiulus (Geopachyiulus)oraniensis (now Afropachyiulus oraniensis) from ‘Hammam Bou Hadjar’. All these specieswere later listed in Brolemann’s (1921a) checklist of North African myriapods. Brolemann(1921b, 1923) described 6 new species and subspecies of Blaniulidae from Algeria amongwhich three cavenicolous taxa: Archichoneiulus drahoni bouarab, A. drahoni maareb and A.

brevicornis, found in caves in Alger (Ifri Bouarab Cave near Aït Ali; Ifri Maareb Cave nearAzerou Tidjer and Tessereft Guiril Cave near Djebel Heidzer, respectively). In the same work,he also described the epigeous species Archichoeneiulus crebresulcatus, Microchoneiulus

gracilis (both from Argiles de Zaouia, Mouzaïa) and M. barbaricola from Mont Baborbetween Sétif and Bougie (now Béjaia). Brolemann (1925b) described two subspecies ofSchizophyllum (now Ommatoiulus), namely Schizophyllum, fusco-unilineatum aumalense

from Aumale and S. diplurum appendiculatum from Alger, respectively. The former was laterraised to a full species and placed in the genus Archiulus, together with two new species,Archiulus albosignatus from Kabylie and Archiulus gauthieri from Jebel Bou Zegza (bothnow placed in the genus Ommatoiulus) (Brolemann 1931). In the same paper he (Brolemann1931) also described Cylindroiulus (Phalloiulus) distinctus gauthieri, which Read (2005)later considered a full species.

Schubart (1932, 1960, 1963) recorded C. truncorum (already known from Algeria) and twofurther species, Schizophyllum tetuanum and Brachyiulus lusitanus the latter with a questionmark. In another publication, Schubart (1952) updated Brolemann’s (1921) list of NorthAfrican millipedes, quoting twenty julid species for Algeria, excluding Iulus corticalis andadding Thalassisobates littoralis previously recorded by Brolemann (1925b) from algeriancoast.

In their revision of the Cylindroiulus truncorum group Korsós & Enghoff (1990) reportedC. truncorum from Annaba in Algeria. Abrous-Kherbouche and Mauriès (1996) published achecklist of the Algerian millipedes in which they mentioned 19 (sub)species of Julida (not

Nesrine Akkari et al.454

including both A. drahoni subspecies) and reported ‘Blaniulus’ corticalis as dubiouslyoccurring in the country. A few years later, Read (2005) revised the Cylindroiulus distinctus

group, describing 6 new species from Algeria: C. attemsi, C. djebelensis, C. jijelensis, C.

ouridae, C. pavoalgerinus and C. maurus. The last addition to the list of Algerian Julida isthat of Afropachyiulus mauriesi which was based on Meinert’s collection from Annaba(Akkari & Enghoff 2008b). With this last species the number of Algerian Julida has reached29 (1 doubtful) (sub)species.

In Morocco the order Julida has not been the subject of study for 50 years. Attems (1903)was the first to record the order in the country describing Schizophyllum (Bothroiulus)tetuanum. This remained the sole Moroccan julid for almost 20 years until Brolemann (1924)described Schizophyllum gravieri (now Ommatoiulus gravieri) from Jebel Tachdirt in the‘Grand Atlas Marocain’. Two subspecies were described later by the same author on the baseof the millipede collection held in the Institut Scientifique Chérifien (Brolemann 1928), viz.,Schizophyllum lapidarium marrocanum (now O. lapidarius marrocanus) from severallocalities (Mçoun, Tiflet, Rabat, Oued Ykem, forest of Maâmoura, Dar Bel Amri, Dayet erRoumi, Tanger, Oued Mellah, Casablanca and Mazagan) and S. albolineatum parvum (nowO. albolineatus) from Tanger. Subsequently, Schubart (1932) described a new genus andspecies Solaenoiulus lohmanderi from Tanger, but the genus Solaenoiulus was synonymisedlater with Cylindroiulus (Read 1990). The tribe Pachyiulini was first recorded from Moroccoby Verhoeff (1936) who described Geopachyiulus (Nesopachyiulus) lepineyi (nowAfropachyiulus lepineyi) from Daia Chiker Cave in Taza. The most important contribution tothe study of Moroccan Julida is due to Schubart who in two papers (Schubart 1952, 1960)described altogether 11 new species: Nesopachyiulus hercules (now Dolichoiulus hercules)from South Alger, Cylindroiulus rifanus from Le Rif, nine species of the genus Schizophyllum

(now Ommatoiulus), viz., Schizophyllum ademinum Schubart, 1952 from the forest ofAdemine, S. atlanteum from Haut Atlas, S. cristatum from northwest of Chemaïa, Safi, S.

curvum from south of Marrakech, S. hamatum from southwest of Agadir, S. haouense fromnorthwest of Marrakech, S. malhommei from Haut Atlas, S. quadridentatum from south ofMarrakech and S. panousei from Mechra Sfi and Aouinet n’Ait Oussa. Moreover, he(Schubart 1960) gave two further new records for Morocco: Schizophyllum hispanicum (C. L.Koch, 1847) (now Ommatoiulus rutilans) and S. lapidarium. Thus, the number of MoroccanJulida has reached 19 (sub)species.

The first julidan species described from Tunisia was Iulus punicus (now Ommatoiulus

punicus) from the surroundings of Tunis (Brölemann 1894). Subsequently, Silvestri (1896)recorded Iulus lapidarius (now Ommatoiulus (Apareiulus) lapidarius) from Souk el Arba, AïnDraham, Babouch and Tabarka and Diploiulus distinctus from Babouch and Tabarka whichAttems (1927a) later synonymised with Cylindroiulus algerinus. Silvestri (1896) alsodescribed Diploiulus truncorum (now Cylindroiulus truncorum) from Aïn Draham andTabarka. Attems’ contributions to the Tunisian julid fauna (1899, 1908) comprise thedescriptions of three new species, namely Pachyiulus (Typhlopachiulus) comatus (nowAfropachyiulus comatus), Typhloblaniulus verhoeffi (now Choneiulus verhoeffi) andCylindroiulus tunetanus.

In his list of North African myriapods, Brolemann (1921a) mentioned eight species ofJulida for Tunisia: Choneiulus verhoeffi, Ommatoiulus lapidarius, O. punicus, Cylindroiulus

tunetanus, C. truncorum, C. algerinus and C. distinctus. Later, he described the subspecies

The millipede order Julida in North Africa 455

Schizophyllum fusco-unilineatum seurati (now Ommatoiulus seurati) from Jebel Ichkeul nearMateur (Brolemann 1925a). Two years later, Attems (1927b) described a new subspecies ofArchiulus fuscounilineatus, A. f. denticulatus (now Ommatoiulus fuscounilineatus) on thebasis of the material of ‘Julus mediterraneus’ from ‘Tunis’ in the collection of Robert Latzel.In his list of North African millipedes Schubart (1952) rejected O. lapidarius from theTunisian fauna. Later, Turk (1955) recorded an immature specimen of Schizophyllum fusco-

unilineatus from La Marsa (misspelled La Masa) near Tunis, and Ceuca (1967) recordedBrachyiulus (Microbrachiyulus) stuxbergi from Le Kef. Recently, Akkari & Voigtländer(2007) and Akkari & Enghoff (2008a, 2008b) described two new species, Ommatoiulus

malleatus and Cylindroiulus mitta, from the country, and clarified the taxonomic status ofsome older and poorly known taxa. Thus, the overall number of Tunisian Julida has reached13 species. However, the millipede fauna of Tunisia (and the other countries in North Africa)have never been a subject of large comprehensive taxonomic or ecological surveys. In fact,many taxa are merely known from their original descriptions and their biology and ecology iscompletely unknown.

The julidan faunas of Libya and Egypt are the least studied in North Africa. In fact, thereare only two species records from Libya: Iulus rimosus described from ‘Tripolitania’ by Karsh(1881) but later synonymised with Ommatoiulus lapidarius by Silvestri (1896) and theLibyan endemic Macheiroiulus libicus described from ‘Cirenaica’ (Manfredi 1939).Likewise, only two species, Brachyiulus lusitanus and Nopoiulus kochii are known fromEgypt (Kheirallah 1980, Kheirallah et al. 2000) both certainly introduced. There are also twoenigmatic species described by Porat (1876), Iulus seticaudus and Iulus lepidus, but theirtaxonomic status remains uncertain. I. seticaudus was based on a female specimen from‘Nubia’. The description suggests a species of the julid genus Pachyiulus but in the absenceof a male, this cannot be verified, just as it remains uncertain whether the specimen came fromthe Egyptian or from the Sudanese part of Nubia. I. lepidus was based on some specimensfrom ‘Egypten’, but although the description places it in the Julidae, no guess can be madeabout its more exact identity.

Following the two recent reviews of the myriapod orders Scolopendromorpha in Tunisia(Akkari et al. 2008) and Glomerida in North Africa (Golovatch et al. 2009), we here presenta review of the millipede order Julida. The aim is to put on record the abundant material ofJulida which has been accumulated in the last five years and to provide information on thetaxonomy, distribution and habitats preferences of Tunisian julidans. For facility of users, anidentification key for the Tunisian species is presented, with new illustrations of theirgonopods. The julidan fauna of the North African countries is summarised in Table 1,including a few new country records for Algeria, Morocco and Tunisia.

2. Materials and methods

Unless stated otherwise, the material treated herein was collected by the first author in theperiod 2003–2008. All the material is preserved in 70 or 96 % ethanol and is shared betweenthe Field Museum of Natural History, Chicago (FMNH); National Museum of NaturalHistory, Sofia (NMNHS); Natural History Museum of Denmark, University of Copenhagen(ZMUC); Muséum National d’Histoire Naturelle, Paris (MNHN) and Senckenberg Museumfür Naturkunde, Görlitz (SMNG). Apart from the reference to the original description, thecitations for each species refer only to records from Tunisia. The specimens were studied


using a stereo microscope Stemi SV (Zeiss). The gonopods of some species were dissectedand mounted on slides. The drawings were made using a camera lucida mounted on LabroluxS (Leitz) microscope.

3. Results

Taxonomic part

Order Julida

Family Blaniulidae C. L. Koch, 1847

Genus Choneiulus Brolemann, 1921

Choneiulus verhoeffi (Attems, 1899)

Typhloblaniulus verhoeffi Attems 1899: 330; Plate 16, Figs 61–65, key. Type locality:Zaghouan, Tunisia.

Typhloblaniulus verhoeffi: Brolemann 1921a: 99, key.

Choneiulus verhoeffi: Brolemann 1921b: 210.

Choneiulus verhoeffi: Schubart 1952: 220.

Choneiulus verhoeffi: Enghoff 1984: 194, Figs 5–7, 17–18, key.

General distribution: Tunisia.

Distribution in Tunisia (Map 1): Zaghouan (Attems 1899).

Habitat: Unknown.

Remarks: C. verhoeffi is the only eyeless species of the genus Choneiulus. According toEnghoff (1984) this could indicate troglophily which is plausible considering the type localityof this species – Jebel Zaghouan – one of the Tunisian mountains richest in caves. Despite thelack of recent records of C. verhoeffi, the presence of Blaniulidae in the country was recentlyconfirmed through the record of an epigean ocellated subadult specimen of an undeterminedspecies [1T subadult, Béni Mtir, Kroumirie, N36º44.582, E08º44.832, alt. 503 m, oak forestdominated by Quercus suber, under bark, 19.II.2007, N. Akkari leg. (ZMUC)].

Family Julidae Leach, 1814

Tribe Cylindroiulini Verhoeff, 1930

Genus Cylindroiulus Verhoeff, 1894

Cylindroiulus algerinus (Brölemann, 1897) Figs 1–2

Iulus (Phallouiulus) algerinus Brölemann 1897: 269, Pl. 4, Figs 34–38. Type locality: Algeria(Coll. Lucas).

Diploiulus distinctus: Silvestri 1896: 160, Figs 9, 10.

Phallouiulus algerinus: Brolemann 1921a: 99, key.

Phalloiulus algerinus: Attems 1908: 113, Figs 13–16.

Cylindroiulus (Phalloiulus) algerinus: Attems 1927a: 208.

Cylindroiulus algerinus: Read 2005: 1497, Figs 8–15.

Cylindroiulus algerinus: Akkari & Enghoff 2008a: 65, Figs 7, 10.


Material examined: TUNISIA: 1T, 1R, 2 juveniles, Jendouba Governorate, Aïn Draham,N36º47, E08º41, alt. 511 m, mixed oak forest, 24.IV.2005, N. Akkari leg. (NMNHS); 1T,Jendouba Governorate, Tabarka, 10.XII.2003, N. Akkari leg. (ZMUC); 1T, 1R, BéjaGovernorate, Jebel el Jouza, close to Amdoun Village, N36º49, E09º, alt. 557 m, 5.II.2004, N.Akkari leg. (NMNHS); 1 T, same locality, 10.III.2005, N. Akkari leg. (ZMUC); 1T, 1R,Jendouba Governorate, Béni Mtir, N36°44.673, E08°44.588, alt. 493 m, under logs,18.II.2007, N. Akkari leg. (NMNHS); 1T, 1R, same locality, alt. 493 m, 19.II.2007, N. Akkarileg. (ZMUC); 1R subadult, 1R in bad condition, 2 juveniles, Jendouba Governorate, 7 kmfrom Aïn Draham, N36°44.303, E08°40.923, alt. 709 m, Quercus suber forest, slope, under stonesand leaf litter, 21.III.2008, N. Akkari & P. Stoev leg. (FMNH); 2TT, 4RR, 2 subadults, 5juveniles, Jendouba Governorate, 15 km from Tabarka, N36°49.965, E08°42.336, alt. 232 m, Q. suber forest, slope, under stones and barks, 21.III.2008, N. Akkari & P. Stoev leg.(FMNH); 1T, severalRR and juveniles, Jendouba Governorate, 9 km from HammamBourguiba (west of Aïn Draham), N36°48.046, E08°39.544, alt. 379 m, pine forest, humid,close to river, under stones, logs and leaf litter, 22.III.2008, P. Stoev & N. Akkari leg.(FMNH).

General distribution: Tunisia and Algeria.

Distribution in Tunisia (Map 2): known from the region of Kroumirie and from a locality12 km south of Thala (for latter record see Read 2005) in north-western Tunisia.

Altitudinal range in Tunisia: from approx. 230 m to approx. 700 m.

Habitats: oak forests dominated by Quercus faginea and Q. suber; coniferous forestsdominated by Pinus pinaster. Under bark, stones, and in decaying logs and leaf litter.


Map 1 Distribution of Choneiulus verhoeffi in Tunisia.

100 km

Cylindroiulus attemsi Read, 2005 Figs 3–4

Cylindroiulus (Phalloiulus) distinctus: Attems 1927a: 207, Figs 268–271. Misidentification!

Cylindroiulus attemsi Read 2005: 1502, Figs 16–20, 58. Type locality: Bona, Algeria.

Cylindroiulus attemsi: Akkari & Enghoff 2008a: 65, Figs 8, 11.

Material examined: TUNISIA: 3TT, 4RR, 1 juvenile, Jendouba Governorate, Feidja Nat.Park, N36º29, E08º18, alt. 687 m, mixed oak forest dominated by Quercus faginea and Q.

suber, 8.V.2005, N. Akkari leg. (ZMUC); 1T, same locality, 7.III.2004, N. Akkari leg.(NMNHS); 1T, 2RR, Jendouba Governorate, Béni Mtir, N36º44, E08º44, alt. 444 m, mixedoak forest, in litter, 18.II.2007, N. Akkari leg. (NMNHS).

General distribution: Algeria and Tunisia.

Distribution in Tunisia (Map 3): known only from two localities in the Kroumirie region inthe north-western part of the country.

Altitudinal range in Tunisia: from approx. 450 m up to 700 m.

Habitats: humid oak forests dominated by Quercus faginea and Q. suber with Phillyrea

angustifolia and Cistus trifolius. Under stones, and in decaying logs and leaf litter.

Remarks: While recording C. distinctus from ‘Tunis’ (it is unclear whether he referred tothe country or to the city of Tunis), Attems (1927a) provided illustrations of a specimencollected from Philippeville, Algeria, which according to Read (2005) differs significantlyfrom the original description of Lucas (1846) and the subsequent one of Brölemann (1897).Having additional material from Annaba and Jijel in Algeria (kept in ZMUC) Read (op. cit.)described the new species C. attemsi, in which she accommodated Attems’ erroneous recordof distinctus from Tunisia. The new material from Kroumirie constitutes the first formalrecord of C. attemsi in Tunisia.


Map 2 Distribution of Cylindroiulus algerinus in Tunisia.

100 km

Cylindroiulus mitta Akkari & Enghoff, 2008 Fig. 5

Cylindroiulus mitta Akkari & Enghoff 2008a: 2, Figs 1–15. Type locality: JebelAbderrahman, Tunisia.

Material examined: 2TT, 11RR, Cap Bon Peninsula, Nabeul Governorate, JebelAbderrahman, 12.XI.2006 (ZMUC), 15RR and juveniles, Zaghouan Governorate, OuedEzzit, N36° 30’.933, E10°18’831, 180 m, Quercus forest, 27.III.2005, N. Akkari leg. (ZMUC)[data published in Akkari & Enghoff 2008a]; 5RR, Cap Bon Peninsula, Nabeul Governorate,near Oued El Abid Village, N36°51.804, E10°44.711, alt. 79 m, mixed Eucalyptus and Pinus

forest, under stones, 24.III.2008, P. Stoev & N. Akkari leg. (FMNH).


Distribution in Tunisia (Map 4): known only from a few localities in the Cap Bon Peninsulaand the area of Zaghouan (Akkari & Enghoff 2008a; present paper). This species was alsorecently collected from the surroundings of Tunis, between Carthage and Sidi Bou Said(Akkari & Enghoff unpublished, material in ZMUC).

Altitudinal range in Tunisia: from approx. 10 m to approx. 200 m.

Habitats: mixed forests dominated by Quercus coccifera and Pinus halepensis with scatteredtrees of Eucalyptus. Grassy slope in a suburban area close to the sea shore. Under stones andlogs.

Remarks: C. mitta is the only species of the C. distinctus group known from north-easternTunisia in contrast to the two congeners of the same group (C. algerinus and C. attemsi),which seem to be confined to the north-west.


Map 3 Distribution of Cylindroiulus attemsi in Tunisia.

100 km

Cylindroiulus truncorum (Silvestri, 1896) Fig. 6

Diploiulus truncorum Silvestri 1896: 160, Figs 11–13. Type locality: Aïn Draham, Tunisia.

Iulus (Anoploiulus) africanus Brölemann, 1897: 271, Pl. 4, Figs 39–41. Type locality:Algeria: surroundings of Alger, viz., Skikda, Constantine, Bona and La Calle (Coll. Lucas).Synonymy proposed by Attems 1927a, see also Korsόs & Enghoff 1990.

Cylindroiulus truncorum: Attems 1908: 112, Figs 10–12.

Cylindroiulus truncorum: Brolemann 1921a: 99, key.

Cylindroiulus truncorum: Korsόs & Enghoff 1990: 347, Figs 5–8.

Cylindroiulus truncorum: Akkari & Enghoff 2008a: 66, Fig 14.

General distribution: Austria, Belgium, British Isles, Canary Isles, Danish mainland,Finland, France, Germany, Hungary, Lithuania, Luxembourg, Madeira, Norwegian mainland,Poland, Romania, Sweden, Switzerland, The Netherlands, Ukraine (Enghoff & Kime 2004),Algeria, Tunisia.

Distribution in Tunisia (Map 5): hitherto known from the area of Aïn Draham (Attems 1908,Silvestri 1896). C. truncorum was recently collected also from a locality west of Babouch atAlgerian border, and from Aïn Snoussi (Akkari & Enghoff unpublished, material in ZMUC).

Altitudinal range in Tunisia: from approx. 400 to approx. 600 m.Habitats: oak forests dominated by Quercus suber or mixed with Erica arborea and Pinus

halepensis. Under barks of Quercus suber.


Map 4 Distribution of Cylindroiulus mitta in Tunisia.

100 km

Cylindroiulus tunetanus Attems, 1908 Fig. 7

Cylindroiulus tunetanus Attems 1908: 111, Figs 6–9. Type locality: Aïn Draham, Tunisia.

Cylindroiulus tunetanus: Brolemann 1921a: 99, key.

Cylindroiulus (Phalloiulus) tunetanus: Schubart 1952: 220, key.

Cylindroiulus tunetanus: Read 2005: 1529.

Cylindroiulus tunetanus: Akkari & Enghoff 2008a: 67, Figs 13, 15.

Material examined: 1T, Jendouba Governorate, Aïn Draham, oak forest, N36º47, E08º41,alt. 511 m, 5.II.2004, N. Akkari leg. (ZMUC).


Distribution in Tunisia (Map 6): a rare species, which was hitherto known only from AïnDraham. It was recently collected also from Babouch at the Algerian border (Akkari &Enghoff unpublished, material in ZMUC).

Altitudinal range in Tunisia: found only at approx. 500 m altitude.

Habitats: oak forests dominated by Quercus faginea and Q. suber. Under logs.


Map 5 Distribution of Cylindroiulus truncorum in Tunisia.

100 km

Tribe Pachyiulini Verhoeff, 1899

Genus Afropachyiulus Schubart, 1960

Afropachyiulus comatus (Attems, 1899) Figs 8–9

Pachyiulus (Typhlopachyiulus) comatus Attems 1899: 328; pl. 16, Figs 75–78. Type locality:

Medjez el Bab, Tunisia.

Pachyiulus (Typhlopachyiulus) comatus: Brolemann 1921a: 99, key.

Geopachyiulus (Geopachyiulus) comatus: Schubart 1952: 220.

Afropachyiulus (Dolichoiulus) comatus: Schubart 1960: 191.

Afropachyliulus comatus: Strasser 1970: 2.

Afropachyiulus comatus: Enghoff 1992: 13.

Afropachyiulus comatus: Akkari & Enghoff 2008b: 31, Figs 5–9.

Material examined: 1T, 4RR, Béja Governorate, Nefza, under stones, 2.X.2005 (ZMUC)

[data published in Akkari & Enghoff 2008b], 2TT, 1R syntypes, Medjez el Bab, Attems det.

(Natural History Museum Wien).


Distribution in Tunisia (Map 7): rare species, which is known only from Medjez el Bab

(Attems 1899) and Nefza (Akkari & Enghoff 2008b) in the Kroumirie region.

Altitudinal range in Tunisia: found only at low altitudes (50–150 m).

Habitats: open grassland with sparse crops of Olea europaea, Pistacia lentiscus and Myrtus

communis. Under stones.


Map 6 Distribution of Cylindroiulus tunetanus in Tunisia.

100 km


Figs 1–7 Shape of gonopods of the species of Cylindroiulus in Tunisia: 1–2. C. algerinus: 1:promerite and mesomerite, mesal view; 2: posterior gonopods, mesal view; 3–4. C. attemsi:3: promerite and mesomerite, posterior view; 4: posterior gonopods, anterior view; 5. C.

mitta, anterior and posterior gonopods, mesal view; 6: C. truncorum, anterior and posteriorgonopods, mesal view; 7: C. tunetanus, anterior and posterior gonopods, mesal view. Scalebars: 0.1 mm. Abbreviations: b: brachite; m: mesomerite; p: promerite; px: paracoxalprocess; rx: coxal rim; s: solenomerite.

1 2

654

3

7


Figs 8–9 Afropachyiulus comatus, gonopods: 8: promerite, posterior view; 9: posterior gonopod,mesal view.

Map 7 Distribution of Afropachyiulus comatus in Tunisia.

8 9

100 km

Tribe Schizophyllini Verhoeff, 1909

Genus Ommatoiulus Latzel, 1884

Ommatoiulus aumalensis (Brolemann, 1925) Figs 10–11

Schizophyllum fusco-unilineatum aumalanse Brolemann 1925b: 245, pl. Figs 1–5, key. Typelocality: Aumale, Algeria.

Material examined: 2TT, 2RR, Béja Governorate, Jebel El Jouza close to Amdoun Village,N36º49, E09º, alt. 557 m, oak forest, 4.XI.2004, N. Akkari leg. (ZMUC).

General distribution: Algeria, Tunisia.

Distribution in Tunisia (Map 8): known only from Kroumirie.

Altitudinal range in Tunisia: collected at approx. 600 m altitude.

Habitats: mixed oak forests dominated by Quercus faginea and Q. suber with anundergrowth of Erica arborea, Cistus trifolius and Clematis flammula. Under stones.

Remarks: This species was initially described as a subspecies of Schizophyllum fusco-

unilineatum on the basis of a single specimen from Aumale, Algeria (Brolemann 1925b).Shortly after that, having further material available from Médéa, Marengo (forest of SidiSliman), Djebel Bou Zerga, Rusgubiae and ‘jardins des Facultés d’Alger’ Brolemann (1931)raised it to a full species and placed it in the genus Archiulus Berlese, 1884. It was later listedunder the genus Schizophyllum in Shubart’s (1952) checklist of North African millipedes andas Ommatoiulus aumalensis in Abrous-Kherbouche & Mauriès (1996). It is reported here forthe first time from Tunisia.


Map 8 Distribution of four Ommatoiulus species in Tunisia: O. malleatus (filled circle), O. seurati

(triangle), O. aumalensis (pentagon), O. fuscounilineatus (filled lozenge).

100 km

Ommatoiulus fuscounilineatus (Lucas, 1846)

Iulus fusco-unilineatum Lucas 1846: 286. Type locality: Algeria.

Schizophyllum (Bothroiulus) fusco-unilineatum: Brölemann 1897: 256, Pl. 3, Figs 1–10.

Archiulus fuscounilineatum denticulatus n. subsp. Attems 1927b: 285, Figs 76–78. Typelocality: Tunis.

Schizophyllum fusco-unilineatus: Turk 1955: 277.

General distribution: Algeria, Tunisia.

Distribution in Tunisia (Map 8): reported from the area of the city of Tunis (see Brölemann1897, Turk 1955 and Attems 1927b).

Habitat: unknown.

Remarks: The former records of Brölemann (1897) and Turk (1955) were based onfemales and immatures only. The status of the Attems’ (1927b) Archiulus fuscounilineatus

denticulatus will remain unclear until type specimens are re-examined or topotypic materialbecomes available for study.

Ommatoiulus malleatus Akkari & Voigtländer, 2007 Figs 12–13

Ommatoiulus malleatus Akkari & Voigtländer 2007: 60, Figs 1–5. Type locality: IchkeulNational Park, Tunisia.

Material examined: TUNISIA: 11TT, 14RR, 3 juveniles, Bizerte Governorate, IchkeulNational Park, N37°9, 09°39, alt. 11 m, 24.IX.2005 (SMNG) [data published in Akkari &Voigtländer 2007]; 2TT, 2RR, same locality (MNHN); 2TT, 6 juveniles, JendoubaGovernorate, Béni Mtir, N36°44, E08°44, alt. 408 m, 18.II.2007, N. Akkari leg. (NMNHS);1T, 4RR, same locality, alt. 472 m, 19.II.2007, N. Akkari leg. (ZMUC); ALGERIA: 3TT,Bona, 1869, F. Meinert leg. (ZMUC).


Distribution in Tunisia (Map 8): hitherto known only from Ichkeul National Park and BéniMtir.

Altitudinal range in Tunisia: from sea level up to approx. 500 m.

Habitats: mixed Quercus suber and Q. faginea forests; mixed forest with Olea europaea andPistacia lentiscus. Under stones and logs.

Remarks: The study of samples collected from Bona (now Annaba) by F. Meinert in 1869(kept in ZMUC) has revealed some specimens of O. malleatus, which constitute the firstrecord of this species from Algeria.

Ommatoiulus punicus (Brölemann, 1894) Figs 14–15

Iulus punicus Brölemann 1894: 24, pl. 12, Figs 23–26. Type locality: surroundings of Tunis,Tunisia.

Schizophyllum (Bothroiulus) punicum: Attems 1903: 143, Pl. 11, Figs 77–81.

Schizophyllum (Bothroiulus) punicum: Brolemann 1921a: 99, key.

Schizophyllum (Bothroiulus) punicum: Schubart 1952: 221.

Ommatoiulus punicus: Akkari & Voigtländer 2007: 63, Figs 6, 12, key.


Material examined: TUNISIA: 2TT, Tunis Governorate, Faculty of Sciences of Tunis,15.X.2003, N. Akkari leg. (ZMUC); 1T, 5RR, Ariana Governorate, Nahli Park, 6. XII.2003,N. Akkari leg. (ZMUC); 3TT, 4RR, same locality, 18.IX.2005, N. Akkari leg. (ZMUC); 1T,1 juvenile, Bizerte Governorate, Aïn Ghelal, 21.XI.2003, N. Akkari leg. (ZMUC); 1T, 5RR,2 juveniles, Nabeul Governorate, Jebel Abderrahman, 28.XI.2004, N. Akkari leg. (ZMUC); 1T, Nabeul Governorate, Oued el Abid, 20.X.2004, N. Akkari leg. (ZMUC); 11TT, TunisGovernorate, Jebel Rsas, 21.IX.2005, N. Akkari leg. (ZMUC); 2TT, 4RR, 4 juveniles,Jendouba Governorate, Aïn Draham, N36º47, E08º41, alt. 511 m, 3.X.2005, N. Akkari leg.(ZMUC); 2TT, 2RR, 5 juveniles, Béja Governorate, Nefza, 3.X.2005, N. Akkari leg.(ZMUC); 3TT, 4RR, Tunis Governorate, Bir el Bey, Bou Kornine, 1.X.2006, N. Akkari leg.(NMNHS); 1T, Zaghouan Governorate, Jebel el Fahs, 9.X.2005, N. Akkari leg. (ZMUC); 1T, 3RR, Ariana Governorate, Jebel Ammar close to Sidi Thabet, 6.X.2006, N. Akkari leg.(NMHNS); 2TT, 15RR and juveniles, Jendouba Governorate, Aïn Draham, Col des Ruines,N36º47, E08º41, alt. 511 m, 14.IV.2007, N. Akkari leg. (ZMUC); 1T, severalsubadults/juveniles, Zaghouan Governorate, Jebel Zaghouan, along the track Sidi BouGabrine (N36°22.423, E10°06.328) – Sidi Abdel kader Cave (N36°22.419, E10°06.371) –Saida Mannoubia (N36°22.650, E10°06.332) – the asphalt road to Zaghouan (N36°22.924, E10°06.789), alt. 650–780 m, mixed forest, under stones and leaf litter, 19.III.2008, P. Stoev &N. Akkari leg. (FMNH); 1T, 3RR, Kairouan Governorate, 6 km from Oueslatia, N35°51.785,E09°30.972, alt. 581 m, sparse olive trees, Roman ruins, bush, open area, stone debris, understones, 6.III.2008, N. Akkari & P. Stoev leg. (FMNH); 1T, Jendouba Governorate, Tabarka,the Genoese fort and surroundings, N36°57.838, E08°44.680, alt. 20–30 m, slope facing thesea, grass, rocks, under stones and logs, 22.III.2008, P. Stoev & N. Akkari leg. (FMNH).ALGERIA: 1T, Bona, 1869, F. Meinert leg. (ZMUC).


Distribution in Tunisia (Map 9): a widespread species; in northern Tunisia it ranges fromMogods-Kroumirie Mts in the west to the Gulf of Tunis area, the eastern part of the Ridge(Jebel Zaghouan, Jebel el Fahs) and the Cap Bon Peninsula (Jebel Abderrahman, Oued elAbid) in the east. In central Tunisia it is known from the plain of Kairouan (Oueslatia).Brölemann (1894) and Silvestri (1896) recorded it from Tunis and Aïn Draham.

Altitudinal range in Tunisia: known from sea level up to approx. 800 m.

Habitats: an euryecious species found in oak forests dominated either by Quercus faginea

or Q. suber with Cistus trifolius and Clematis flammula. It inhabits also: coniferous forests ofPinus halepensis; mixed forests of P. halepensis and Quercus ilex; heterogeneous woods ofOlea europaea, Ceratonia siliqua, and Pistacia lentiscus; forests dominated by Tetraclinis

articulata, Pinus halepensis and Eucalyptus; open areas with shrubs and rocks; ruins; coastalgrasslands close to sea; urban and suburban areas. Under stones, leaf litter and in decayinglogs.

Remarks: This species was hitherto known only from Tunisia. Male specimens collectedfrom Annaba in 1869 were found in the collection of F. Meinert (ZMUC), which is the firstrecord of this species from Algeria.


Ommatoiulus seurati (Brolemann, 1925) Figs 16–17

Schizophyllum fusco-unilineatum seurati Brolemann 1925a: 61, Pl. II, Figs 1–4; Pl. III, Figs5–7. Type locality: Ichkeul National Park, Tunisia.

Archiulus seurati: Brolemann 1931: 122.

Schizophyllum (Bothroiulus) seurati: Schubart 1952: 221.

Schizophyllum (Ischnophyllum) seurati: Schubart 1963: 89.

Ommatoiulus seurati: Akkari & Voigtländer 2007: 63, Figs 7–13, key.

Material examined: 16TT, 8RR, Bizerte Governorate, Ichkeul National Park, inside thePark, N37°08.301, 09°41.03, alt. 14 m, mixed broad-leaf forest with Olea europaea, Pistacia

lentiscus, under stones, 24.IX.2005; 1R, same locality, N37°08.301, 09°41.031, alt. 53 m,under stones, 23.III.2008, N. Akkari & P. Stoev leg. (FMNH); 2TT, 2RR, ZaghouanGovernorate, Jebel Zaghouan, N36°21, E10°05, alt. 500 m, mixed forest, 4.X.2008, N. Akkarileg. (NMNHS).


Distribution in Tunisia (Map 8): known only from Ichkeul National Park and JebelZaghouan. In Ichkeul National Park O. seurati occurs sympatrically with O. malleatus.

Altitudinal range in Tunisia: known from sea level up to 500 m.

Habitats: mixed forests with Olea europaea and Pistacia lentiscus; mixed forest dominatedby Quercus ilex with Ceratonia siliqua and Olea europaea. Under stones.

Remarks: Initially regarded as a subspecies of O. fusco-unilineatus, it was raised bySchubart (1952) to a full species.


Map 9 Distribution of Ommatoiulus punicus in Tunisia.

100 km

Tribe Brachyiulini Verhoeff, 1909

Genus Brachyiulus Berlese, 1884

Brachyiulus pusillus (Leach, 1814) Fig. 18

Material examined: 2TT, 2RR, Tozeur Governorate, Tozeur City, ‘Ancien Oasis deTozeur’, palm trees, 25.III.2006, N. Akkari leg. (ZMUC); 2TT, 2RR, Gabès Governorate,Kattena Oasis, 23.III.2006, N. Akkari leg. (NMNHS); 10TT, 27RR, same locality,N33°45.046, E10°11.183, alt. 22 m, under stones and in leaf litter, 15.III.2008, N. Akkari &P. Stoev leg. (FMNH); numerous specimens, 6 km from Kébili Governorate, Tombar, oasis,N33°43.463, E08°54.349, alt. 14 m, palm trees, under stones, 12.III.2008, N. Akkari & P.Stoev leg. (NMNHS).


Figs 10–17 Shape of gonopods of the species of Ommatoiulus in Tunisia: 10–11. O. aumalensis: 10:promerite, posterior view; 11: posterior gonopods, posterior view (redrawn from Brolemann1925b); 12–13: O. malleatus: 12: promerite, anterior view; 13: posterior gonopods, posteriorview; 14–15: O. punicus: 14: promerite, anterior view; 15: posterior gonopods, posteriorview; 16–17: O. seurati: 16: promerite, posterior view; 17: posterior gonopods, posteriorview (redrawn from Brolemann 1925a). Scale bar: 0.1 mm. Abbreviations: a: promeriteapex; m: mesomerite; s: solenomerite.

General distribution: Albania, Austria, Azores, Bosnia and Herzegovina, British Isles,Canary Islands, Channel Islands, Croatia, Danish mainland, Estonia, France, Germany,Greece, Ireland, Italy, Kaliningrad region of Russia, Lithuania, Luxemburg, Madeira,Monaco, Northern Ireland, Poland, Romania, Spain, Sweden, Switzerland, Yugoslavia (sensuFauna Europaea = Serbia, Kosovo, Voivodina, Montenegro) (Enghoff & Kime 2004), Tunisia.

Distribution in Tunisia (Map 10): B. pusillus is the only millipede reported from thesouthern part of Tunisia. So far it is known from few oases around the saline depression Chottel Jérid and from an oasis in the region of Gabès.

Altitudinal range in Tunisia: hitherto recorded only from 14–22 m.

Habitats: inhabits coastal and mountainous oases dominated by Phoenix dactylifera withorchards of Punica granatum and Prunus amygdalus. Found in decomposing fruits ofpomegranate, under stones and in leaf litter.

Remarks: Although known from several countries in the North Mediterranean realm(Greece, Italy, France, Spain), B. pusillus is here recorded for the first time from Tunisia andNorth Africa. As yet, it is the only millipede recorded in southern Tunisia, and exclusivelyfrom oases, where it has most likely been introduced by human activity.

Brachyiulus stuxbergi (Fanzago, 1875) Fig. 19

Brachyiulus (Microbrachyiulus) stuxbergi: Ceuca 1967: 553.

Material examined: TUNISIA: 1T, Béja Governorate, Jebel El Jouza, close to AmdounVillage, N36º49, E09º, alt. 557 m, oak forest, 5.II.2004, N. Akkari leg. (ZMUC) 1T, 9RR,Ariana Governorate, Sidi Thabet, open area with stands of Olea europaea, 17.III.2004, N.Akkari leg. (ZMUC); 2TT, 2RR, Jendouba Governorate, Béni Mtir, N36°44, E08°44, alt.472 m, 18.II.2007, N. Akkari leg. (NMNHS); 1T, 1R, same locality, alt. 500 m, 18.II.2007,N. Akkari leg. (ZMUC); 3RR, same locality, N36°43.888, E08°44.105, alt. 404 m, Quercus

suber, close to the road, under stones, 21.III.2008 P. Stoev & N. Akkari leg. (FMNH); 1T, 1R, Zaghouan Governorate, Jebel Zaghouan, surroundings of the marabout Sidi Bou Gabrine,N36°22.423, E10°06.328, alt. 642 m, meadows, scattered trees, under stones and leaf litter,17.III.2008 N. Akkari leg. (FMNH); 1T, 1R, same locality, 17.III.2008, N. Akkari leg.(NMNHS). ALGERIA: 1T, Aokas (Bejaia), 12.V.1983, G. Osella leg. (Museo Civico di StoriaNaturale, Verona); 1T, Setif, Djebel Babor, 2100 m, 5.XI.1984, Boffa, Casale, Cavzzuti,Gavetti & Giachino leg. (Museo Regionale di Scienze Naturali, Torino).

General distribution: Italy, Greece, Malta (Enghoff & Kime 2004), Tunisia, Algeria.

Distribution in Tunisia (Map 10): B. stuxbergi occurs in the northern part of Tunisia, fromthe western humid forests of the Kroumirie to the eastern part of the Dorsale Ridge (mountainof Zaghouan). The species is also known from Le Kef in the High Tell (Ceuca 1967) and wasrecently collected from the surroundings of Tunis, in parks of the ancient city of Carthage(Akkari & Enghoff unpublished, material in ZMUC).

Altitudinal range in Tunisia: known from approx. 400 m up to approx. 600 m.

Habitats: oak forests dominated by Quercus suber, mountain meadows, stands of Olea

europaea and Pistacia lentiscus, sometimes mixed with Ceratonia siliqua. Under stones, logsand in leaf litter.

Remark: This Mediterranean species seems much more widespread in North Africa thenpreviously thought. Here it is confirmed from Tunisia and recorded for the first time fromAlgeria.



Map 10 Distribution of the Brachyiulus species in Tunisia: B. pusillus (open square), B.

stuxbergi (filled square).

100 km

4. Discussion

4.1. Taxonomy and faunistics

The order Julida is the richest and most diverse millipede order in North Africa. Fifty-eight(sub)species belonging to 12 genera and 3 families are currently known from Morocco,Algeria, Tunisia, Libya and Egypt (Tab. 1). The old record of Pachyiulus varius (Brandt,1841) from Algeria is yet to be confirmed. Likewise, the two enigmatic species Iulus

seticaudus and Iulus lepidus are of uncertain taxonomic placement and are presently excludedfrom the list of North African julids.

The greatest diversity is observed in Algeria where the order is represented by threefamilies, seven genera and 32 (sub)species. Twenty species from eight genera and twofamilies were hitherto known in Morocco, while these figures for Tunisian julids are 15, fiveand two, respectively. The Libyan and Egyptian julid faunas remain very poorly known withonly two species being reported from each country. The most species-rich genera in NorthAfrica are Ommatoiulus and Cylindroiulus known with 24 and 14 species respectively. Thefamily Blaniulidae is represented by 10 (sub)species, of which the genus Archichoneiulus

Brolemann, 1921, contains five species and subspecies, all of them known only from theiroriginal descriptions (Brolemann 1921b, 1923). Among Julidae, the tribe Pachyiulini, isrepresented by 2 (3?) genera and 6 species, of which Afropachyiulus comprises 4 species witha vicariant distribution from Morocco to Tunisia. While in Europe the tribe Brachyiulini isquite diversified and comprises large genera such as Megaphyllum Verhoeff, 1894 withapproximately 50 species (Enghoff and Kime 2004). In North Africa, this tribe is representedby only three species of the genus Brachyiulus.


Figs 18–19 Shape of gonopods of the species of Brachyiulus in Tunisia: 18: B. pusillus (redrawn fromBlower 1985); 19: B. stuxbergi. Abbreviations: p: promerite, s: solenomerite, ph: phylacum.


Tab.

1A

n an

nota

ted

list

of

the

Nor

th A

fric

an s

peci

es o

f Ju

lida

.

Moro

cco

Alg

eria

Tu

nis

iaL

ibya

Egyp

t

Fam

ily B

lan

iuli

dae

Koch

, 1847

Arc

hic

honei

ulu

s bre

vico

rnis

Bro

lem

ann,

192

1

+ (

Bro

lem

ann

1921

b,19

23, S

chub

art

1952

,A

brou

s-K

herb

ouch

e &

Mau

riès

199

6)

Arc

hic

honei

ulu

s cr

ebre

sulc

atu

s

Bro

lem

ann,

192

1

+ (

Bro

lem

ann

1921

b,19

23, S

chub

art

1952

,A

brou

s-K

herb

ouch

e &

Mau

riès

199

6)

Arc

hic

honei

ulu

sdra

honi

dra

honi

(Gia

rd, 1

899)

+ (

Gia

rd, 1

899,

Sch

ubar

t 19

52, A

brou

s-K

herb

ouch

e &

Mau

riès

1996

)

Arc

hic

honei

ulu

s dra

honi

bouara

bB

role

man

n, 1

921

+ (

Bro

lem

ann

1921

b,19

23, S

chub

art

1952

)

Arc

hic

honei

ulu

s dra

honi

maare

b

Bro

lem

ann,

192

1+

(B

role

man

n 19

21b,

1923

, Sch

ubar

t 19

52)

Chonei

ulu

s ve

rhoef

fi (

Att

ems,

1899

)

+ (

Att

ems

1899

,B

role

man

n 19

21a,

1921

b, S

chub

art

1952

,E

ngho

ff 1

984)

Mic

roch

onei

ulu

s babori

cola

Bro

lem

ann,

192

1

+ (

Bro

lem

ann

1921

b,19

23, S

chub

art

1952

,A

brou

s-K

herb

ouch

e &

Mau

riès

199

6)

Mic

roch

onei

ulu

s gra

cili

s

Bro

lem

ann,

192

1

+ (

Bro

lem

ann

1921

b,19

23, S

chub

art

1952

,19

63)

Nopoiu

lus

koch

ii (

Ger

vais

, 184

7)+

(K

heir

alla

h et

al.

2000

)

Pro

tero

iulu

s his

panus

Sch

ubar

t,19

59+

new

rec

ord


Tab.

1co

nt.

Moro

cco

Alg

eria

Tu

nis

iaL

ibya

Egyp

t

Fam

ily N

emaso

mati

dae

Boll

man

, 1893

Thala

ssis

obate

s li

ttora

lis

(Sil

vest

ri, 1

903)

+ (

Sch

ubar

t 19

52,

Abr

ous-

Khe

rbou

che

&M

auri

ès 1

996)

Fam

ily J

uli

dae

Lea

ch,

1814

Su

bfa

mil

y P

aec

top

hyll

inae

Ver

hoef

f, 1

900

Tri

be

Paec

top

hyll

ini

Ver

hoef

f,

1900

Mach

eiro

iulu

s li

bic

us

Man

fred

i,19

39+

(Man

fred

i 19

39)

Su

bfa

mil

y J

uli

nae

Lea

ch,

1814

Tri

be

Pach

yiu

lin

i V

erh

oef

f,

1899

Afr

opach

yiulu

s co

matu

s (A

ttem

s,18

99)

+ (

Att

ems

1899

,V

erho

eff,

190

0,B

role

man

n 19

21a,

Sch

ubar

t 19

52, 1

960,

Stra

sser

197

0, A

kkar

i &

Eng

hoff

200

8b)

Afr

opach

yiulu

s le

pin

eyi

(Ver

hoef

f, 1

936)

+ (

Ver

hoef

f 19

36,

Sch

ubar

t 19

52, 1

960,

Stra

sser

197

0, A

kkar

i &

Eng

hoff

200

8b)

Afr

opach

yiulu

s m

auri

esiA

kkar

i&

Eng

hoff

, 200

8+

(A

kkar

i &

Eng

hoff

2008

b)


Tab.

1co

nt.

Moro

cco

Alg

eria

Tu

nis

iaL

ibya

Egyp

t

Afr

opach

yiulu

s ora

nie

nsi

s

(Ver

hoef

f, 1

900)

+ (

Ver

hoef

f 19

00,

Bro

lem

ann

1921

a,S

chub

art

1952

, 196

0,St

rass

er 1

970,

Abr

ous-

Khe

rbou

che

& M

auri

ès19

96, A

kkar

i &

Eng

hoff

2008

b)

Doli

choiu

lus

her

cule

s(S

chub

art,

1960

)+

(S

chub

art

1960

)

Tri

be

Sch

izop

hyll

ini

Ver

hoef

f,

1909

Om

mato

iulu

sla

pid

ari

us

lapid

ari

us

(Luc

as, 1

846)

+ (

Sch

ubar

t 19

52, 1

960)

+ (

Luc

as 1

846,

Sch

ubar

t 19

52, A

brou

s-K

herb

ouch

e &

Mau

riès

1996

)

+ (

Kar

sch

1881

,M

anfr

edi

1939

,S

chub

art

1952

, 196

0)

Om

mato

iulu

s (A

.)la

pid

ari

us

maro

ccanus

(Bro

lem

ann,

192

8)+

(B

role

man

n 19

28,

Sch

ubar

t 19

52, 1

960)

Om

mato

iulu

s adem

inus

(Sch

ubar

t, 19

52)

+(S

chub

art

1952

)

Om

mato

iulu

s alb

oli

nea

tus

(Luc

as, 1

845)

+(B

role

man

n 19

28,

Sch

ubar

t 19

52)

Om

mato

iulu

s alb

osi

gnatu

s

(Bro

lem

ann,

193

1)

+ (

Bro

lem

ann

1931

,S

chub

art

1963

, Abr

ous-

Khe

rbou

che

& M

auri

ès19

96)

Om

mato

iulu

s atl

ante

us

(Sch

ubar

t, 19

60)

+(S

chub

art

1960

)

Om

mato

iulu

s aum

ale

nsi

s

(Bro

lem

ann,

192

5)

+ (

Bro

lem

ann

1925

b,19

31, S

chub

art

1952

,19

63, A

brou

s-K

herb

ouch

e &

Mau

riès

1996

)

+ n

ew r

ecord


Tab.

1co

nt.

Moro

cco

Alg

eria

Tu

nis

iaL

ibya

Egyp

t

Om

mato

iulu

s cr

ista

tus

(Sch

ubar

t,19

60)

+ (

Sch

ubar

t 19

60)

Om

mato

iulu

s cu

rvus

(Sch

ubar

t,19

60)

+ (

Sch

ubar

t 19

60)

Om

mato

iulu

s dip

luru

s

appen

dic

ula

tus

(Bro

lem

ann,

1925

)

+ (

Bro

lem

ann

1925

b,S

chub

art

1952

, Abr

ous-

Khe

rbou

che

& M

auri

ès19

96)

Om

mato

iulu

s fu

scounil

inea

tus

fusc

ounil

inea

tus

(Luc

as, 1

846)

+(L

ucas

184

6, P

ococ

k18

92, B

röle

man

n 18

97,

1921

, Sch

ubar

t 19

52,

1963

, Abr

ous-

Khe

rbou

che

& M

auri

ès19

96)

+?

(Brö

lem

ann

1897

,T

urk

1955

)

Om

mato

iulu

s fu

scounil

inea

tus

den

ticu

latu

s (A

ttem

s, 1

927)

+

(A

ttem

s 19

27,

Sch

ubar

t 19

60)

Om

mato

iulu

sgauth

ieri

(Bro

lem

ann,

193

1)

+ (

Bro

lem

ann

1931

,S

chub

art

1952

, 196

3,A

brou

s-K

herb

ouch

e &

Mau

riès

199

6)

Om

mato

iulu

s gra

vier

i

(Bro

lem

ann,

192

4)+

(B

role

man

n 19

24,

Sch

ubar

t 19

60)

Om

mato

iulu

s ham

atu

s(S

chub

art,

1960

)+

(S

chub

art

1960

)

Om

mato

iulu

s haouze

nse

(Sch

ubar

t, 19

60)

+ (

Sch

ubar

t 19

60)

Om

mato

iulu

s m

alh

om

mei

(Sch

ubar

t, 19

60)

+ (

Sch

ubar

t 19

60)

Om

mato

iulu

s m

all

eatu

s A

kkar

i&

Voi

gtla

nder

, 200

7+

new

rec

ord

+

(A

kkar

i &

Voi

gtla

nder

200

7)


Tab.

1co

nt.

Moro

cco

Alg

eria

Tu

nis

iaL

ibya

Egyp

t

Om

mato

iulu

s panouse

i

(Sch

ubar

t, 19

52)

+(S

chub

art

1952

)

Om

mato

iulu

s punic

us

(Brö

lem

ann,

189

4)+

new

rec

ord

+ (

Brö

lem

ann

1894

,19

21, A

ttem

s 19

03,

1908

, Sch

ubar

t 19

52,

Akk

ari

& V

oigt

land

er20

07)

Om

mato

iulu

s quadri

den

tatu

s

(Sch

ubar

t, 19

60)

+ (

Sch

ubar

t 19

60)

Om

mato

iulu

s ru

tila

ns

(C. L

.K

och,

184

7)+

(S

chub

art

1960

)

Om

mato

iulu

s se

ura

ti

(Bro

lem

ann,

192

5)

+ (

Bro

lem

ann

1925

a,S

chub

art

1952

, Akk

ari

& V

oigt

land

er 2

007)

Om

mato

iulu

s te

tuanus

(Att

ems,

1903

)

+ (

Att

ems

1903

,B

role

man

n 19

21,

Sch

ubar

t 19

52, 1

960)

(Sch

ubar

t 19

60)

Tri

be

Cyli

nd

roiu

lin

i V

erh

oef

f,

1930

Cyl

indro

iulu

s alg

erin

us

Brö

lem

ann,

189

7

+ (

Brö

lem

ann

1897

,19

21a,

Sch

ubar

t 19

52,

Abr

ous-

Khe

rbou

che

&M

auri

ès 1

996,

Rea

d20

05)

+ (

Sil

vest

ri 1

896,

Bro

lem

ann

1921

a,A

ttem

s 19

08, 1

927a

,R

ead

2005

, Akk

ari

&E

ngho

ff 2

008b

)

Cyl

indro

iulu

s att

emsi

Rea

d, 2

005

+ (

Rea

d 20

05)

+ (

Att

ems

1927

a,A

kkar

i &

Eng

hoff

2008

b)


Tab.

1co

nt.

Moro

cco

Alg

eria

Tu

nis

iaL

ibya

Egyp

t

Cyl

indro

iulu

s dis

tinct

us

(Luc

as,

1846

)

+ (

Luc

as 1

846,

Poc

ock

1892

, Brö

lem

ann

1897

,19

21a,

193

1, S

chub

art

1952

, 196

3, A

brou

s-K

herb

ouch

e &

Mau

riès

1996

, A

brou

s-K

herb

ouch

e, 1

996,

Rea

d 20

05)

Cyl

indro

iulu

s dje

bel

ensi

sR

ead,

2005

+ (

Rea

d 20

05)

Cyl

indro

iulu

s gauth

ieri

(Bro

lem

ann,

193

1)

+ (

Bro

lem

ann

1931

,S

chub

art

1952

, Abr

ous-

Khe

rbou

che

& M

auri

ès19

96,

Abr

ous-

Khe

rbou

che,

199

6,R

ead

2005

)

Cyl

indro

iulu

s ji

jele

nsi

sR

ead,

2005

+ (

Rea

d 20

05)

Cyl

indro

iulu

s lo

hm

ander

i

(Sch

ubar

t, 19

32)

+ (

Sch

ubar

t 19

32, 1

960,

Rea

d 19

90, 2

005)

Cyl

indro

iulu

s m

auru

sR

ead,

2005

+ (

Rea

d 20

05)

Cyl

indro

iulu

s m

itta

Akk

ari

&E

ngho

ff, 2

008

+ (

Akk

ari

& E

ngho

ff20

08a)

Cyl

indro

iulu

s ouri

dae

Rea

d,20

05+

(R

ead

2005

)

Cyl

indro

iulu

s parv

oalg

erin

us

Rea

d, 2

005

+ (

Rea

d 20

05)


Tab.

1co

nt.

Moro

cco

Alg

eria

Tu

nis

iaL

ibya

Egyp

t

Cyl

indro

iulu

s ri

fanus

Sch

ubar

t,19

60+

(Sch

ubar

t 19

60, R

ead

2005

)

Cyl

indro

iulu

s tr

unco

rum

(Sil

vest

ri, 1

896)

+ (

Brö

lem

ann

1897

,S

chub

art

1932

, 195

2,19

63, A

brou

s-K

herb

ouch

e &

Mau

riès

1996

)

+ (S

ilves

tri 1

896,

Atte

ms

1908

, Bro

lem

ann

1921

a,19

31, S

chub

art 1

952,

Kor

sόs

& E

ngho

ff 1

990,

Akk

ari &

Eng

hoff

200

8a)

Cyl

indro

iulu

s tu

net

anus

Att

ems,

1908

+ (

Bro

lem

ann

1921

a,S

chub

art

1952

)

+ (

Att

ems

1908

,B

role

man

n 19

21a,

Sch

ubar

t 19

52, R

ead

2005

)

Tri

be

Bra

chyiu

lin

i V

erh

oef

f,

1909

Bra

chyi

ulu

s lu

sita

nus

Ver

hoef

f,18

98+

(S

chub

art

1963

) +

(K

heir

alla

h 19

80)

Bra

chyi

ulu

s pusi

llus

(Lea

ch,

1814

)+

new

rec

ord

Bra

chyi

ulu

s st

uxb

ergi

(Fan

zago

,18

75)

+ n

ew r

ecord

+ (

Ceu

ca 1

967)

20

32

15 (

1 d

ou

btf

ul)

22

Proteroiulus hispanus Schubart, 1959 (Blaniulidae) is here reported for the first time fromMorocco. Material studied: Morocco: Azrou/ Ifrane area, 1400–2000 m, in rotten wood,17–19.IV.1989, Zool. Mus. Copenh. Exp. Leg. (ZMUC). The species was hitherto knownonly from southern Spain (Sierra Nevada).

In Tunisia, the most species-rich genera are again Ommatoiulus and Cylindroiulus,

represented by 5 species each. However, this number is likely to increase in the near futureafter the identification of unknown (new) taxa of Ommatoiulus from remote regions, viz. LaGalite Islands and Chambi Mts. (study in progress). Studying some julidans from ‘Tunis’, inthe collection of Latzel (Wien Museum), Attems (1927b) described the subspeciesOmmatoiulus fuscounilineatus denticulatus which he distinguished from the nominate by thepresence of an ‘irregularly’ serrated notch separating the two characteristic apical lobes of thepromerite, and by setae being visible in posterior view. Until further material, preferablyincluding the type specimens, is studied the taxonomic status of this subspecies remainsuncertain. Ommatoiulus punicus as currently delimited is quite a variable taxon, and adetailed analysis may well necessitate splitting it into several (sub)species.

Several authors (e.g. Silvestri 1896, Brolemann 1921, Attems, 1927a, Schubart 1952)recorded Cylindroiulus distinctus (Lucas, 1846) from the Aïn Draham region of Tunisia.However, these records seem erroneous and may refer either to C. algerinus (Silvestri’srecord) or C. attemsi. Since none of these authors provided illustrations of the gonopods oftheir material and no samples were obtained during the intensive recent collection in thecountry the presence of C. distinctus in Tunisia remains dubious.

4.2. Distribution patterns

The majority of North African Julida are local or regional endemics with a very few specieshaving larger ranges. These are mainly species of European or Mediterranean origins, such asBrachyiulus lusitanus, B. pusillus, B. stuxbergi and Nopoiulus kochii. Thalassisobates

littoralis is a widespread halophilous millipede, which occurs under stones and seaweeds, inrock crevices and shingle in or shortly above the tidal zone (Enghoff 1987). It has anamphiatlantic distribution with several localities in the coastal areas of Western Europe and afew on the Atlantic coast of North America. Whether its presence in North America is due tonatural dispersal or facilitated by man is still questionable (op. cit.). Thalassisobates littoralis

was also recorded from coastal biotopes in northern Algeria by Brolemann (1925b). Somespecies like Ommatoiulus albolineatus, O. rutilans and Proteroiulus hispanus only occur onboth sides of the Gibraltar strait and thus probably represent remains of formerly continuouspopulations. At least one julidan, B. pusillus, is alien to North Africa, being hitherto knownonly from man-made habitats. Perhaps the same hold true for Nopoiulus kochii which issynanthropic in Europe, and has also been introduced to North and South America and NewZealand (Lee 2006). Arthropod introductions from Europe to North Africa are quite possiblegiven the similar climate and habitat conditions and the long historical connections of humanactivities between these two regions. This process seems bidirectional, from Europe to Africaand from Africa to Europe. Cylindroiulus truncorum, apparently native to Algeria and Tunisiagained a wide distribution in Europe (mainly known from greenhouses, parks, gardens) andmight represent an example of the latter case (see Korsós & Enghoff 1990).


The genera Cylindroiulus, Afropachyiulus (Julida) and Glomeris (Glomerida) showremarkably similar distribution patterns in North Africa. Species of these genera are mainlydistributed in a narrow belt along the Mediterranean coast, comprising the Atlas Mountains.The greatest similarity in the faunal composition is observed between Tunisia and Algeria, asnine Julida species occur in both countries (Table 1). This is also the case for the species ofGlomeris, notably G. punica, G. sublimbata and G. klugii (Golovatch et al. 2009). The faunalcomposition of Moroccan Julida is quite different from the remaining Maghreb countries, asonly Ommatoiulus tetuanus and O. lapidarius are known also from Algeria and Algeria andLibya respectively. The specific composition of the Moroccan fauna could be explained bythe isolated position of the regions where most of the species come from. Most Moroccanspecies occur in the Rif Mountains in the north (continuation of the Baetic Cordilleramountains in the South of Spain) and in the High and Anti-Atlas mountains (see Brolemann1924, Schubart 1960). These massifs lie in western Morocco and are subject to a differentclimate, being also influenced by the Atlantic Ocean and the European continent.

The species Macheroiulus libicus, described from Cirenaica, northeastern Libya, is aninteresting biogeographic case worthy of mention. It is the sole member of the tribePaectophyllini in North Africa, a tribe whose distribution ranges from Greece, Cyprus andLibya in the west to the Caucasus and northern Iran in the east, and is thought to have itscenter of origin in Turkey (Enghoff 1995). The genus comprises two further species, M.

compressicauda (Verhoeff, 1901) and M. martini Enghoff, 1995, known from Greece andTurkey, respectively (Enghoff 1995, 2007).

In Tunisia, Julida are mainly distributed in northern and central areas, with a singleexception, the European species Brachyiulus pusillus, introduced to the southern oases (Map10). Autochthonous Julida are virtually unknown south of the Steppe Zone, which constitutesthe transition area to the desert region of Tunisia. The mountains of Kroumirie and Mogodsin the north and northwest show the highest species diversity. Being the easternmost part ofthe Tell Atlas, this region is characterised by a strong Mediterranean influence and a mosaicof dense mesophilous oak and pine forests. Such habitats are among the few in the countrythat best correspond to the food and humidity requirements of most millipedes. Thus, most ofCylindroiulus, Afropachyiulus and Ommatoiulus species occur in this region. Higher speciesdiversity among other myriapod taxa such as the genus Glomeris (Golovatch et al. 2009) andthe order Scolopendromorpha (Akkari et al. 2008) has also been registered in this part of thecountry.

Cylindroiulus mitta is the sole member of the genus that occurs in northeastern Tunisia,from Cap Bon Peninsula down to the Gulf of Tunis area. The four remaining congeners arerestricted to the western part of the country, where their ranges extend to the neighbouringparts of Algeria. Other myriapods that seem to be endemic to the mountains of north-easternTunisia or show similar distribution patterns are Choneiulus verhoeffi, O. seurati, O.

fuscounilineatus, Scolopendra morsitans Linnaeus, 1758 (Map 2 in Akkari et al. 2008), andGlomeris colorata Golovatch, Mauriès, Akkari & Stoev, 2009 (Fig. 29 in Golovatch et al.2009). The Cap Bon Peninsula constitutes a contact zone between the northern humid andsouthern arid zones and exhibits high spatial heterogeneity. This is confirmed by the presencein this particular area of species with Palaearctic and Saharo-Sindian biogeographicalaffinities which is perhaps the case of Scolopendra canidens Newport, 1844 (Map 1 in Akkariet al. 2008).


Brachyiulus stuxbergi is widespread throughout north Tunisia. Its southern border ofdistribution is perhaps limited by the Dorsale Ridge, as is the case with some other myriapods,e.g. Glomeris punica Attems, 1900 (Glomerida), Cormocephalus gervaisianus (C. L. Koch,1841) and Cryptops trisulcatus Brölemann, 1902 (Scolopendromorpha).

The most widespread julidan in Tunisia is Ommatoiulus punicus being distributed from theKroumirie-Mogods Mts in the north-northwest, along the Gulf of Tunis, in Jebel Zaghouan,Jebel Mansour in the northeast, up to the Cap Bon Peninsula (Oued el Abid). In centralTunisia, O. punicus occurs in the Kairouan Plain (Map 9). Taking into account the highvariability of some morphological traits shown in the different populations (e.g. coloration oflegs, promerite, paracoxite and mesomerite conformation, etc.) we presume that severalcryptic species will be recognised among this polymorphic complex in the future.

4.3. Habitat preferences

In North Africa, as elsewhere, julidans are primarily confined to forests, which provide thenecessary humidity for their survival. However, there are some exceptions of speciesoccurring in more ‘extreme’ environments such as high-altitude mountain grasslands, caves,littoral, oases, etc.

Five cave-dwelling julidans are known from North Africa: Afropachyiulus lepineyi (Julida:Pachyiulini) found from Daia Chiker Cave in Taza, Morocco and four Blanulidae:Archichoneiulus drahoni, A. drahoni bouarab Brolemann 1921, A. drahoni maareb and A.

brevicornis, all described from different caves in Djurjura Mountain, Algeria (Brolemann1923).

The only species adapted to temporary submergence in salt water is Thalassisobates

littoralis, which is so far only known in the Mahgreb from Algeria but is probably much morewidespread in the littoral of North Africa. Ommatoiulus gravieri shows another interestingcase of specialisation. It occurs in Djebel Tachdirt in High Atlas in Morocco at 3000–3200 maltitude (Brolemann 1924, Beron 2008), and represents the only North African alticolousmillipede.

Studying the ecology of some myriapods in the National Park of Chréa in Algeria, Abrous-Kherbouche (1996) reported Ommatoiulus gauthieri and Cylindroiulus distinctus from anorchard with Cerasus avium and forests of Pinus halepensis, Quercus ilex, Quercus suber,Cedrus atlantica, etc. Her analysis shows that these two species are most abundant inorchards and cedar forest.

As regards the habitat preferences of Tunisian julidans, with few exceptions, most of themare associated with oak formations dominated by either Quercus faginea or Q. suber, usuallywith understorey of Erica arborea, Myrtus communis, and Cistus trifolius. Such biotopes arepreferred by species like Cylindroiulus tunetanus, C. truncorum, C. attemsi and Ommatoiulus

aumalensis, which are quite abundant in the thick humus layers of oak woods of theKroumirie and Mogods mountains. Cylindroiulus algerinus is a moderately eurytopic species,which, in addition to the humid Quercus suber and Quercus faginea forests of Kroumirie,occurs also in coniferous woods dominated by Pinus pinaster. Likewise, Brachyiulus

stuxbergi and Ommatoiulus malleatus were recorded from both humid and subhumid habitats,in oak formations as well as in mixed broad-leaved forests. Ommatoiulus punicus can beconsidered as euryecious species; it was recorded from several habitats like oak forests


dominated either by Quercus faginea or Q. suber; coniferous forests with Pinus halepensis;mixed forests with P. halepensis and Q. ilex; open areas with shrubs and rocks; ruins, urbanand suburban areas. Few other Julida, like Afropachyiulus comatus and a still undeterminedspecies of Ommatoiulus (only females and juveniles were found) occur under stones ingrasslands. Similar open habitats seem to be preferred also by Cylindroiulus sp. which wasfound on a grassy slope facing the sea, only a few metres from the water line, and by C. mitta,which was collected on a suburban polluted slope only few metres away from the seashore.Several specimens of a still undetermined species of the genus Ommatoiulus were found onthe beach of Kelibia, on sandy substratum under decaying leaves of Nerium oleander andAcacia mimosa.

Brachyiulus pusillus is so far the only undoubtedly introduced julidan known in Tunisia; itwas recorded in a few oases in the south, always associated with pomegranate trees, especiallyunder decaying fruits, leaves and logs. Brachyiulus stuxbergi has also been found insynanthropic habitats, in the managed parks of the ancient city of Carthage.

An identification key for the Tunisian species of Julida

1(2) a distinct suture between prozona and metazona; eyes when present in a subtriangulargroup; main part of male gonopods retracted into the 7th body ring

Family Julidae . . . . . . . . . . . . . . . . . . . . . . . . .3

2(1) suture between prozona and metazona indistinct; eyes in a single row or absent; malegonopods freely projecting from the 7th body ring

Family Blaniulidae . . . . .Choneiulus verhoeffi

3(8) head with frontal setae; gonopods without a free mesomerite . . . . . . . . . . . . . . . . . . .4

4(5) no distinct colour pattern, no preanal projection, telson densely pilose; anteriorgonopod (Fig. 8) large and club-shaped with 2 apical denticles, without a flagellum

Tribe Pachyiulini . . . .Afropachyiulus comatus

5(4) two longitudinal pale stripes dorsally, usually a small preanal projection, telson notdensely pilose; anterior gonopod small, without apical denticles, with a flagellum

Tribe Brachyiulini . . . . . . . . . . . . . . . . . . . . . .6

6(7) posterior gonopod with a large, striate phylacum (Fig. 19, ph) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Brachyiulus stuxbergi

7(6) posterior gonopod without a large phylacum (Fig. 18) . . . . . . . .Brachyiulus pusillus

8(3) head without frontal setae; posterior gonopods with clearly separate mesomerite andopisthomerite . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .9

9(18) ozopores opening on the suture between prozona and metazona; preanal ring withoutor with a blunt caudal projection; gonopods with a flagellum arising from the base ofpromerite Tribe Cylindroiulini . . . . .Genus Cylindroiulus

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .10

10(15) mesomerite (m) much shorter than promerite (p) (Figs 1, 3, 5); paracoxal rim (rx)large; solenomerite (s) and brachite (b) well separated; brachite apically complex;paracoxal process absent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .11


11(12) solenomerite and brachite deeply separated; brachite with two serrated lobes,solenomerite with a jagged rectangular plate (Fig. 4) . . . . . . . . . .C. (Phalloiulus) attemsi

12(11) solenomerite and brachite partly separated but linked by a lamella . . . . . . . . . . .13

13(14) adult body size: 31–37 mm (T) up to 47 mm (R); brachite with three flanges at thetip and a large beak-shaped projection distally (Fig. 2) . . . . . . .C. (Phalloiulus) algerinus

14(13) adult body size: 11–13 mm (T) up to 16.7 mm (R); brachite mitten-shaped, broadand rounded, bent apically (Fig. 5, b) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .C. mitta

15(10) mesomerite and promerite of the same length; paracoxal rim reduced; solenomeriteand brachite incompletely separated; brachite with more or less simple tip; paracoxalprocess present . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .16

16(17) paracoxite with serrated edge (Fig. 7, px) . . . . . . . . . . . . . . . . . . . . . . .C. tunetanus

17(16) paracoxite with smooth edge (Fig. 6, px) . . . . . . . . . . . . . . . . . . . . . .C. truncorum

18(9) ozopores opening behind the suture; preanal ring with a long, protruding, pointedcaudal projection; flagellum absent Tribe Schizophillini . . . . .Genus Ommatoiulus

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .19

19(20) mesomerite hammer-shaped (Fig. 13, m) . . . . . . . . . . . . . . . . . . . . . . .O. malleatus

20(19) mesomerite hook-shaped (Figs 13, 15, 17, m) . . . . . . . . . . . . . . . . . . . . . . . . . . . .21

21(22) promerite strongly narrowed in its distal part (Fig. 14) . . . . . . . . . . . . . .O. punicus

22(21) promerite slightly narrowed in its distal part (Figs 10, 16, p) . . . . . . . . . . . . . . . .23

23(24) promerite with a finger-shaped apex (Fig. 10, a) . . . . . . . . . . . . . . . .O. aumalensis

24(23) promerite without a finger-shaped apex . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .25

25(26) solenomerite short, mesomerite broad, abruptly narrowed apically . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .O. fuscounilineatus

26(25) solenomerite long, mesomerite uniformly slender (Fig. 17) . . . . . . . . . . .O. seurati

5. Acknowledgements

N.A. is much obliged to Jean-Paul Mauriès (MNHN) for the identification of a large partof the material treated herein. Karin Voigtländer and David Russell supported N.A. during hervisits to the SMNG, Mohamed Hédi Ktari (Tunis, Tunisia) is thanked for his overall supportto N.A. Hans Reip (Jena, Germany) and Jean-Paul Mauriès provided some obscure papers onNorth African Julida. Richard Hoffman (Martinsville, Virginia, USA) called our attention toIulus seticaudus, and Guiseppe Osella and E. Gavetti provided access to material in theVerona and Torino museums, respectively. Atef Belkahla (Tunis, Tunisia) kindly helped withthe preparation of maps. The collecting trip of N.A. and P.S. in Tunisia in March 2008 wassupported financially with a grant from the Field Museum, kindly arranged by Petra Sierwald(FMNH). N.A. and P.S. are grateful to Willi Xylander (SMNG) for supporting theirparticipation at the 14th congress of Myriapodology, Görlitz in 2008. Thanks are also due toHelen Read and Jean-Paul Mauriès for their useful comments on the manuscript and to JohnLewis who improved the English of the final draft.


6. References

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Accepted 13 October 2009


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The millipede order Julida (Myriapoda: Diplopoda) in - Senckenberg