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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: nesrineakkari@gmail.com
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
Nesrine Akkari et al.456
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.
The millipede order Julida in North Africa 457
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.
Nesrine Akkari et al.458
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.
The millipede order Julida in North Africa 459
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).
General distribution: Tunisia.
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.
Nesrine Akkari et al.460
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.
The millipede order Julida in North Africa 461
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).
General distribution: Tunisia and Algeria.
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.
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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).
General distribution: Tunisia.
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.
The millipede order Julida in North Africa 463
Map 6 Distribution of Cylindroiulus tunetanus in Tunisia.
100 km
Nesrine Akkari et al.464
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
The millipede order Julida in North Africa 465
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.
Nesrine Akkari et al.466
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).
General distribution: Tunisia and Algeria.
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.
The millipede order Julida in North Africa 467
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).
General distribution: Tunisia and Algeria.
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.
Nesrine Akkari et al.468
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).
General distribution: Tunisia.
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.
The millipede order Julida in North Africa 469
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).
Nesrine Akkari et al.470
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.
The millipede order Julida in North Africa 471
Nesrine Akkari et al.472
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.
The millipede order Julida in North Africa 473
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.
Nesrine Akkari et al.474
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
The millipede order Julida in North Africa 475
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)
Nesrine Akkari et al.476
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
The millipede order Julida in North Africa 477
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)
Nesrine Akkari et al.478
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)
The millipede order Julida in North Africa 479
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)
Nesrine Akkari et al.480
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 millipede order Julida in North Africa 481
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).
Nesrine Akkari et al.482
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
The millipede order Julida in North Africa 483
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
Nesrine Akkari et al.484
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.
The millipede order Julida in North Africa 485
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Accepted 13 October 2009
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