Revision of Schizomeritus 111

Internation Journal of Myriapodology 1 (2008) 111-122 Sofi a–Moscow

Revision of the Iberian Schizomeritus Verhoeff , 1931 (Diplopoda: ), with the description of three new species

Per Djursvoll Th e Natural History Collections, Bergen Museum, University of Bergen, Muséplass 3, N-5020 Bergen, Norway. E-mail: [email protected]

Abstract Th e genus Schizomeritus Verhoeff , 1931 is revised and the type species S. phantasma (Verhoeff , 1925) is redescribed. Th ree new species from Spain, S. ortizi sp. n., S. esgrimidor sp. n. and S. andalusius sp. n. are described and fi gured. mauriesi Vicente, 1979 is redescribed and included in Schizomeritus. Polydesmus armatus Machado, 1946 is evaluated based on the original description and also included in Schizomeritus. A key to the six species is given.

Key words Diplopoda, , Polydesmidae, Schizomeritus, new species, new synonym, new combination, Iberia, Spain

Introduction Verhoeff (1931) established Polydesmus subgenus Schizomeritus to accommodate Polydesmus phantasma Verhoeff , 1925, described by Verhoeff (1925) based on a single male collected in the Cercedilla, Sierra de Guadarrama, Spain in 1905. Attems (1940) included P. phantasma, when he established Polydesmus subgenus Normarchus Attems, 1940. However, both Jeekel (1971) and Hoff man (1980) kept P. phantasma in Schi- zomeritus and raised Schizomeritus to genus level, which is the present status used e.g. by Enghoff & Kime (2004). When studying unidentifi ed polydesmid material housed in the Museo Nacional de Ciencias Naturales in Madrid, several additional specimens of S. phantasma were discovered. Further, three new congeners were discovered, and are described below.

© Koninklijke Brill NV and Pensoft, 2008 DOI: 10.1163/187525408X316776

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Type material of Polydesmus mauriesi Vicente, 1979 was examined and the species is an obvious member of Schizomeritus. Th e type material of Polydesmus armatus Machado, 1946, stated to be stored in “Museu de Zoologia do Porto”, could unfortunately not be located. However the original description describes characters diff erent from the typi- cal Polydesmus species and it is thus regarded as a Schizomeritus species here.

Material and methods Th e examination of the specimens was done using a Leica MZ Apo stereomicroscope. When making Scanning Electron Micrographs (SEM), gonopods where gently mount- ed on stubs using sticky tabs and the air-dried stubs were sputter coated with gold. Schizomeritus phantasma was observed with a JSM 6400 scanning electron microscope, while a Zeiss Supra 55 UP fi eld emission scanning electron microscope was used for S. esgrimidor, S. ortizi, and S. andalusius. Photographs of tergal structures were taken using a Nicon Coolpix 4500 on the Leica MZ Apo stereomicroscope. Th e terminology concerning gonopod morphology and body characters mostly follows Djursvoll et. al. (2001), corrected in accordance with Golovatch and Wytwer (2007). Th e anterior/posterior moveable telopodite part may be retracted into the cox- ite (in situ) or remain erected. Th e retracted condition serves here for the defi nition of the dorsal and ventral sides of the telopodite. As the telopodite cannot move laterad or mediad, these sides are more obvious to name. In accordance with the traditional gonopod outlines, the erected position serves as basic when describing the direction of gonopod parts. Th e material examined is housed in Museo Nacional de Ciencias Naturales, Ma- drid, Spain (MNCN); Zoologische Statssammlung München, Germany (ZSM) and Muséum National d’Histoire Naturelle, Paris, France (MNHN).

Results Schizomeritus Verhoeff

Schizomeritus Verhoeff , 1931: 419; as subgenus of Polydesmus Latreille, 1803

Type species: Polydesmus phantasma Verhoeff , 1925, by original designation and monotypy. Other species included: Schizomeritus ortizi sp. n., S. esgrimidor sp. n., S. andalusius sp. n., S. mauriesi (Vicente, 1979), comb. n., and S. armatus (Machado, 1946), comb. n. Generic diagnosis: Body with longitudinal furrow on most paraterga (Fig. 2). Gono- pod exomere pointed, not extending beyond and subordinate to well developed, some- times strongly armed distofemoral part (acropodite) (Fig. 5). “Femorite” (i.e. seminal groove piece) with a conspicuous large bulb-like elevation laterally (Figs 4 & 5). Lateral ridge of coxite with dense setation (Figs 3 & 4).

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Description: Body with 20 (18+1+T) body rings, medium to large-sized Polydesmi- dae, body length 10-21 mm, coloration light yellowish brown to brownish or reddish yellow, as in typical Polydesmus s.l. Collum transverse-oval with narrowly rounded and slightly elevated caudal corners, not as wide as head, tuberculation visible and with relatively short setae. Antennae surpassing somite 2; antennomere 5/6 length ratio vari- able; numerous sensilla on antennomeres 5 and 6, few on 7; dorsoparabasal sensory knob on antennomere 7 present. Metaterga 2–4 generally with well developed tergal sculpture (tuberculation) in three transverse rows; setae clavi- to bacilliform, variable in length; paranota horizontal to rather strongly elevated in some species (Fig. 1); limbus elements at margin with sharp saw-like teeth. Metatergum with longitudinal furrow formed by well developed bosses and the lateral margin of paraterga (Fig. 2); paraterga horizontal, with 4 lat- eromarginal incisions, ozopore located laterally a short distance from caudal margin; caudolateral projections on paraterga gradually larger posteriorly. Legs 1.5–2.0x as long as midbody height and with single dorsal macroseta on tibia. Epiproct length modestly variable. Male: Gonopod coxite strongly setose, lateral edge almost two-lobed, forming two fi elds of setae, larger fi eld medially, smaller fi eld posteriorly, two macrosetae more an- teriorly; hooked cannula typical as in Polydesmus s.l.; prefemorite comparatively large and moderately excavated, length almost equal to “femorite” length; seminal groove (Figs 3-6) starting mesally, then halfway along ”femorite” turning laterad to dorsal side of “femorite” before reaching base of acropodite where it makes a short loop ending in a seminal chamber; pilose pulvillus well developed ventrally; “femorite” laterally with large, well set-off bulb; exomere branch relatively short, well set off dorsolaterally bellow recurvature point of seminal groove, gradually tapering towards needle-shaped apex, almost straight, curved or sharply bent. Distofemoral part well developed, with one to several large acropodite processes. Male legs with sphaerotrichomes and numerous modifi ed setae, prefemur conspicu- ously enlarged and covered with long setae especially on legs close to somite 7. Female: (females of ortizi, esgrimidor and armatus unknown): Epigynal ridge dis- tinct with small medial incision; vulvae relatively short in examined species.

1 2 Figures 12. Schizomeritus esgrimidor sp. n.: 1. Metaterga 2–4. 2. Midbody metaterga.

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Key to species (based on males)

1. Exomere sharply bent, forming a knee (Figs 7 & 12); endomere process present ....2 - Exomere straight or slightly curved; endomere absent; large process a2 with tooth (Figs 3, 14, 16 & 17) ...... 3 2. Distofemoral part with all acropodite processes extending well beyond tip of ex- omere; three small acropodite processes (a2-a4) directed caudad (Figs 7-10) ...... S. ortizi sp. n. - Distofemoral part with acropodite processes reaching or slightly extending beyond tip of exomere and with a1 crossing the exomere; two small acropodite processes (a2-a3) directed caudad (Figs 11-13) ...... S. esgrimidor sp. n. 3. Distofemoral part with one long acropodite process a2 distinctly longer than other acropodite processes (Figs 14-16) ...... 4 - Distofemoral part with two large acropodite processes a1 and a2 (Figs 3-6 & 17) ...... 5 4. Acropodite process a2 strongly curved cephalolaterad; with distinct tooth at curva- ture (Figs 14-15) ...... S. andalusius sp. n. - Acropodite process a2 straight, with small tooth medially (Fig. 16) ...... S. mauriesi (Vicente), comb. n. 5. Acropodite process a1 directed laterad; acropodite process a2 directed distad with distinct tooth proximally; a3 serrated; exomere slightly bent caudad (Fig. 17) ...... S. armatus (Machado), comb. n. - Acropodite process a1 directed cephalad, acropodite process a2 directed laterad and slightly curved; a4 smooth; exomere almost straight (Figs 3-6) ...... S. phantasma (Verhoeff )

Schizomeritus phantasma (Verhoeff , 1925) Figs 3-6.

Polydesmus phantasma Verhoeff , 1925: 112; Attems 1927: 60. Polydesmus (Schizomeritus) phantasma Verhoeff : Verhoeff 1931: 419. Polydesmus (Nomarchus) phantasma Verhoeff : Attems 1940: 43. Schizomeritus phantasma (Verhoeff ): Jeekel 1971: 351; Hoff man 1980: 173.

Diagnostic characters: see the key. Material examined: Holotype male, Spain: Madrid province, Sierra de Gua darrama, Cercedilla, 15.x.1905, J. Bolivar (ZSM A20033494). Spain: Madrid province, La Silla, Herreria, El Escorial, 2 males (fragmented), 20.ii.1954, J. Alvarez (MNCN no. 20.07/1388); Spain: Àvila province, Sierra de Gredos, Casillas, 6 males and 7 females, 28.xi.1983, J. Alvarez (MNCN no. 20.07/1333). Redescription: Body 17–18 mm long. Antennae not surpassing somite 3, antenno- meres 5 and 6 almost equal in length. Metaterga 2–4 with modestly elevated paraterga,

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3 4

5 6

Figures 36. Schizomeritus phantasma (Verhoeff , 1925), right gonopod (scanning electron micrographs, SEM): 3. Anteromesial view. 4. Anterolateral view. 5. Lateral view. 6. Posterior view. Abbreviations: exm, exomere; fb, femoral bulb; a, acropodite process; t, tooth; simple arrows showing a fi eld of setae. tuberculation distinct, with short setae. Metatergum  tuberculation barely visible; paraterga horizontal. Male gonopodal exomere needle-shaped, pointed, with solid base and apex located close to base of acropodite processes a1, a2, and a4; acropodite processes a1 and a2 distinctly longer than other acropodite processes, spiniform, a1 large, projecting cepha- lad; process a2 largest, at least as long or longer than exposed part of exomere, slightly curved, projecting laterad, with single proximal tooth; a3-a4 almost shield-like, a3 slightly rounded, a4 smaller and more acute (Figs 3–6). Female epigynal ridge large, convex, with small incision. Vulvae comparatively short.

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Schizomeritus ortizi sp. n. Figs 7-10.

Type material: Holotype male, Spain: Toledo province, Sierra de Toledo, Navahermosa, 23.iv.1967, E. Ortiz (MNCN no. 20.07/1353). Etymology: Named after Dr. E. Ortiz, Madrid, who collected the specimen. Diagnostic characters: see the key. Description: Body 17 mm long. Antennae not surpassing somite 3, antennomere 6 sligthly longer than 5. Metaterga 2–4 tuberculation readily visible, setae short; parater- ga modestly elevated at edge. Metatergum 10 tuberculation barely traceable; paraterga with four lateromarginal incisions but posterior incision barely visible. Male gonopod with pointed and sharply bent exomere, directed caudad and touch- ing base of distofemoral part (Fig. 7); endomere process situated between large femoral bulb and pulvillus, distofemoral part strongly armed, extending well beyond exomere, with fi ve distinct acropodite processes (a1–a5) radiating from a shield-like base; a1

7 8

9 10

Figures 7-10. Schizomeritus ortizi sp. n., right gonopod, holotype (scanning electron micrographs, SEM): 7. Lateral view. 8. Anterolateral view. 9. Distal view. 10. Posterolateral view. Abbreviations: exm, exomere; fb, femoral bulb; endm, endomerite (broken off ); a, acropodite process.

Downloaded from Brill.com09/25/2021 06:58:11PM via free access Revision of Schizomeritus 117 large and spine-like, projecting cephalad; a2–a4 smaller processes projecting caudad; a5 roughly serrated at apex, forming the distalmost part of telopodite (Fig. 9).

Schizomeritus esgrimidor sp. n. Figs 11-13.

Type material: Holotype male, Spain: Avila province, Sierra de Gredos, Piedralaves, E. Ortiz (MNCN no. 20.07/1334). Etymology: From Spanish esgrimidor meaning fencer, as the position of the gonopodal exomere and the process a1 recall a fancing duel; the name is to be regarded as a noun in apposition. Diagnostic characters: see the key. Description: Body 18 mm long. Antennae surpassing somite 2; antennomere 6 slightly longer than 5. Metaterga 2–4 with paraterga set relatively high and elevated at edge, tuberculation distinct with short setae. Metatergum 10 tuberculation low and barely traceable; tergal setae short; paratergum with four lateromarginal incisions, pos- terior incision barely visible. Male gonopod (Figs 11-13) with small and strongly bent exomere; endomere situ- ated between large femoral bulb and pulvillus; distofemoral part (Fig. 13) strongly armed, consisting of four distinct acropodite processes (a1–a4); a1 large and spine- like, projecting cephalad, crossing the opposite directed exomere; a2 and a3 smaller, directed caudad; a4 shield-like and slightly dentate at apex.

11 12

13

Figures 11-13. Schizomeritus esgrimidor sp. n., left gonopod, holotype (scanning electron micro- graphs, SEM): 11. Distoventral view. 12. Lateral view. 13. Distomedial view. Abbreviations: exm, exomere; fb, femoral bulb; endm, endomerite; a, acropodite process.

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Schizomeritus andalusius sp. n. Figs 14-15.

Type material: Holotype male, Spain: Huelva province, Sierra Aracena, Aracena, Gruta de las Maravillas, 11.ii.1930, F. Bonet (MNCN no. 20.07/1267). Paratypes: 1 female, 4 juveniles, same as holotype; 1 male, Spain: Sevilla province, Cazalla de la Sierra, Venta de la Angels?, 22.iii.1975, P. Gamarra (MNHN no. Jc 285). 4 males, 1 female: Spain: Cadis province, Zahara de la Sierra [36.836°N, 5.392°W], 9 km N Grazalema, inside town, un- der wooden plank, 500-550m, between 15h and 17h, 17.iii.2004, Jörg Spelda (ZSM). Etymology: Named after Andalucia Community, Spain. Diagnostic characters: see the key. Description: Body 20–21 mm long. Antennae not surpassing somite 3; antenno- mere 5 longer than 6. Metaterga 2–4 subrectangular, tuberculation distinct, paraterga elevated. Metatergum 10 with low or indistinct tubercles, paraterga with 4 indistinct lateromarginal incisions. Male gonopod (Figs 14-15) exomere thin, needle-shaped, straight to moderately curved; distofemoral part with main acropodite process a2 strongly recurved cepha- lolaterad, touching proximal end of exomere, with small teeth at curvature; acropodite processes a1 and a3 distinct, situated proximally on distofemoral part and close to pul- villus, almost forming a claw, a3 sharply bent laterad. Female epigynal ridge present, with distinct medial incision. Vulvae relatively short.

14 15

Figures 14-15. Schizomeritus andalusius sp. n., left gonopod, holotype (scanning electron micro- graphs, SEM): 14. Medial view. 15. Anteromedial view. Abbreviations: exm, exomere; fb, femoral bulb; a, acropodite process; t, tooth.

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Schizomeritus mauriesi (Vicente, 1979), comb. n Fig. 16.

Polydesmus mauriesi Vicente, 1979: 21. Propolydesmus mauriesi (Vicente, 1979): Enghoff & Golvatch 2003: 82.

Material examined: Spain: Caceres Province, Sierra de Gredos, Piornal, one male and one female paratype, 19.ix.1976, A. Serra (MNHN no. Jc 26). Diagnostic characters: see the key. Redescription: Body 13–17 mm long. Antennae reaching somite 3; antennomere 6 slightly longer than antennomere 5. Metaterga 2–4 with distinct tubercles, setae short; paraterga modestly elevated at the edge, with well developed bosses. Metatergum 10 tuberculation barely discernible, 3rd row about ½ as high as 1st row; paraterga not elevated, with four lateromarginal incisions, posterior incision barely visible. Male gonopod (Fig. 16) with a spinelike, narrow exomere projecting diagonally towards and barely intersecting the large upright directed a2 acropodite process, a2 with small medial tooth; acropodite processes a1 and a3, close to pulvillus, smaller and directed caudad. Female epigynal ridge convex with small incision medially. Vulvae relatively short.

Schizomeritus armatus (Machado, 1946), comb. n Fig. 17.

Polydesmus armatus Machado, 1946: 14.

Th e type material is apparently lost. However, according to the original description (Machado 1946), this is a well defi ned species of Schizomeritus. Diagnostic characters: see the key. Description: Body small and slender, 10 mm, dark brownish-reddish. Male go- nopod with a small, thin, pointed and slightly caudad bent exomere. Distofemoral part with two large processes a1 and a2. A1, referred to as “tibia-tarsus” by Machado (1946), directed laterad, a2, “solenomerito” by Machado (1946), directed distad and with proximal tooth. Two smaller acropodite processes a3 and a4 near pulvillus, both serrate. Th e original description by Machado (1946) reveals a certain resemblance to Schi- zomeritus phantasma (Verhoeff , 1925), but the diff erences, in particular in the shape of the gonopods, allows it to retain its specifi c rank. S. armatus is noticeably smaller than the remaining congeners.

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16 17

Figures 16-17. 16. Schizomeritus mauriesi (Vicente, 1979), comb. n., left gonopod, paratype, medial view (digital photo). 17. S armatus (Machado, 1946), gonopod fi gure redrawn from Machado (1946). Abbre- viations: exm, exomere; a, acropodite process; t, tooth.

Discussion Th e six known Schizomeritus species seem to form two subgroups, i.e. ortizi and esgrimidor, on the one hand, and phantasma, armatus, andalusius, mauriesi, on the other (see Key). Whereas the exomere is clearly homologous in all the species of Schizomeritus, this is not the case for all the acropodite processes. Nevertheless, in ortizi, a1 and a2 are sugges- ted to be homologous with a1 and a2 in esgrimidor, respectively. Furthermore, phantasma, armatus, andalusius and mauriesi all feature a large homologous process a2 with a tooth. Th e Schizomeritus gonopodal character state, with a small exomere, seems to have evolved through a secondary reduction, indicated by the distinct sulcus around the base of the exomere, clearly visible in phantasma, esgrimidor, and andalusius (Figs 3, 12 & 15). Golovatch (1991) suggested a transformation series going from absence of an exomere as in Pacidesmus Golovatch, 1991 (plesiomorphic) through a small exomere (derived) as in Epanerchodus Attems, 1901 and Usbekodesmus Lohmander, 1933, to a large exomere (further derived) in Polydesmus s. str. and Propolydesmus Verhoeff , 1895. Regarding the gonofemoral bulb, it must be noted that Propolydesmus testaceus (C.L. Koch, 1847) and Propolydesmus corsicus (Schubart, 1931) to some extent show a gonofemo- ral bulb but not such a prominent one as in Schizomeritus. Th e dense setation on the lateral ridge of the gonopodal coxite in Schizomerius is also found in the Iberian species Propoly- desmus dismilus (Berlese, 1891) and in Polydesmus liber Golovatch, 1991 from Hong Kong. Convergent evolution or not, and despite the apparent homoplasies mentioned above, I believe this study supports Schizomeritus as a well defi ned monophyletic genus. When redefi ning the genus Propolydesmus, Enghoff and Golovatch (2003) included P. dismilus and all former species of Polydesmus subgenus Hormobrachium Attems, 1940,

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Figure 18. Map showing the distribution of the Schizomeritus Verhoeff , 1931 species. Scale bar 200 km. where the course of the seminal groove extends distad on the telopodite before making a loop and runs into the accessory seminal chamber. Th is is a “normal” Schizomeritus and Polydesmus Latreille, 1802/03 state, too, but markedly diff erent from Propolydesmus pectiniger (the type-species of Propolydesmus), P. laevidentatus (Loksa, 1967) and P. miguelinus (Attems, 1908) where the seminal groove does not extend distad on the telopodite, but makes a larger loop immediately after the start of seminal groove, see Figs 1, 2, and 3 in Enghoff and Golovatch (2003). All six Schizomerius species are restricted to mountainous areas in the Iberian Peninsula south of 41 paralell north (Fig. 18), mainly belonging to the Iberian sclero- phyllous and semi-deciduous forest ecozone. Th e knowledge of the millipede fauna of the Iberian Peninsula is defi cient (Kime 2001), and more collecting will without doubt reveal new species.

Acknowledgements I am indebted to Jörg Spelda for the very valuable advice and comments on the manu- script, as well as some important information kindly shared. I am indebted to Miguel

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Villena Sánches-Valero, Museo Nacional de Ciencias Naturales, Madrid for the loan of a large material of unidentifi ed polydesmids; to Jörg Spelda and Stefan Friedrich, Zool- ogische Staatssammlung, München and Monique Nguyen Duy-Jacquemin, Muséum National d’Histoire Naturelle, Paris for loan of material; to both referees, and espe- cially referee 1 for kindly editing English text. I am also grateful to Egil S. Erichsen, University of Bergen, for technical assistance at the Electron Microscopy Laboratory. Th anks are due to Bjarne Meidell and Trond Andersen, both University of Bergen, for comments on the manuscript. Th anks are also due to Humberto Fonseca Mendes for translating the Portuguese text by Machado (1946).

References

Attems, C.G. (1940) Polydesmoidea III. – Das Tierreich 70: 1-577. Djursvoll, P., Golovatch, S.I., Johanson, K.A. & Meidell, B. (2000) Phylogenetic relationships within Polydesmus sensu lato (Diplopoda: Polydesmidae). – Fragmenta Faunistica 43 (Sup- plement): 37-57. Enghoff , H. & Golovatch, S.I. (2003) Th e millipede genus Propolydesmus Verhoeff , 1895 rede- fi ned, with a revision of the genus in the Canary Islands (Diplopoda, Polydesmida, Polyde- smidae). – Graellsia 59 (1): 79-86. Enghoff , H. & Kime, R.D. (2004) Fauna Europaea: Diplopoda, Polydesmidae. – Fauna Euro- paea version 1.1, http://www.faunaeur.org Golovatch, S.I. (1991) Th e millipede family Polydesmidae in Southeast Asia, with notes on phylogeny (Diplopoda: Polydesmida). – Steenstrupia 17 (4): 141-159. Golovatch, S.I. & Wytwer, J. (2007) nevoi, a new millipede from Israel (Diplo- poda: Polydesmida). – Annales Zoologici 57 (2): 205-210. Hoff man, R.L. (1980) Classifi cation of the Diplopoda. Muséum d’Histoire Naturelle, Genève, 237 pp. Jeekel, C.A.W. (1971) Nomenclator generum et familiarum Diplopodorum: A list of the genus and family-group names in the Class Diplopoda from the 10th edition of Linnaeus, 1758, to the end of 1957. – Monografi eën van de Nederlandse Entomologische Vereniging 5: 1-412. Kime, R.D. (2000) Present knowledge of the distribution of European . – Fragmenta Faunistica 43 (Supplement): 281-294. Machado, A. (1946) Contribuição para o conhecimento dos Miriápodes de Portugal. – Broteria 15 (1): 5-37. Vicente, M.C (1979) Diplopodos polydesmidos de Zamora, Salamanca y Carceres (Espana). Descripcion de una nueva especie del genero Polydesmus Latreille, 1802-03 (Diplopoda, Polydesmidae). – Miscellània zoologica 5: 21-23. Verhoeff , K.W. (1925) Neue Diplopoden-Beiträge. 95. Diplopoden-Aufsatz. – Zoologische Jahrbücher, Abteilung für Systematik, Ökologie und Geographie der Tiere 50: 61-122. Verhoeff , K.W. (1931) Chilognathen aus den Bergamasker Alpen und Nachbargebieten; auch über zwei neue Gattungen der Polydesmoidea aus Spanien und Japan. 121. Diplopoden- Aufsatz. – Zoologische Jahrbücher, Abteilung für Systematik, Ökologie und Geographie der Tiere 61: 410-452 + 2 Taf.

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