THE MILLIPEDE GENUS CYLINDROIULUS VERHOEFF, 1894 (DIPLOPODA: JULIDA; JULIDAE) IN SPAIN: RECENT RECORDS AND DESCRIPTION OF THREE NEW SPECIES FROM THE BASQUE COUNTRY, NAVARRA, ANDALUCÍA AND CATALUÑA

Introduction[Up]

The diplopod fauna of the Iberian Peninsula is relatively under studied in comparison with much of Europe. Following work by Attems and Verhoeff in the early part of the 20^th century there was a relatively long gap until work in the 1970s and 1980s on cave fauna (‍Mauriès & Vicente, 1977a), species from north-west Spain and the Pyrenees (‍Mauriès, 1975; ‍Mauriès & Vicente, 1977b) and some ecological studies by Vicente & Ascaso (‍1990) and Serra et al. (‍1997) in Cataluña and Mauriès & Barraqueta (‍1985) in Biscay. Vicente (‍1981) published a list of species from Cataluña including details to help with identification.

In recent years there has been a resurgence in interest of Iberian millipedes with a variety of collections made and papers published. These include descriptions of new species in the Polydesmida (‍Gilgado et al. 2015b; ‍Djursvoll, 2019), Platydesmida and Polyzoniida (‍Mauriès, 2015, ‍2018), Chordeumatida (‍Gilgado et al., 2015a, ‍2017; ‍Mauriès, 2014; ‍Serra & Mauriès, 2018), Callipodida (‍Gilgado et al., 2020) and the Julid genera Ommatoiulus Latzel, 1884 (‍Akkari & Enghoff, 2012; ‍Akkari et al. 2018) and Cylindroiulus (‍Read, 1989, ‍2007; ‍Reboleira & Enghoff, 2018).

The French fauna is comparatively well studied and some Cylindroiulus species relatively abundant further north have a range that extends down to and/or over to the south side of the Pyrenees, such as Cylindroiulus londinensis (Leach, 1814), Cylindroiulus parisorum (‍Brölemann & Verhoeff, 1896) and Cylindroiulus punctatus (Leach, 1815) (‍Kime & Enghoff, 2017).

Systematics of the genus Cylindroiulus

The genus Cylindroiulus is large, with over 100 species described. Read (‍1990) clarified a previously confusing situation between Cylindroiulus and Allajulus C.L. Koch, 1847 which are now considered separate genera, distinguished by the presence of setae on the apodous body rings in Allajulus and details of the gonopods. Both genera are found on the Iberian Peninsula, although Allajulus is currently represented only by A. spinosus (‍Ribaut, 1904). A few species groups can be identified within the genus, with those relevant to Iberia being: the perforatus-group (Cylindroiulus perforatus, (‍Vehoeff, 1905) Cylindroiulus fenestratus, ‍Read, 1989, Cylindroiulus ventanaea Read, 2007 and Cylindroiulus villumi ‍Reboleira & Enghoff, 2018), distinguished by usually having a perforate mesomerite, a feature which has so far not been seen on any species outside the Iberian peninsular, and denticles on the anterior flagellum-conducting lamella (‍Read, 1989; ‍Reboleira & Enghoff, 2018), a feature which has so far not been seen on any species outside theIberianPeninsula; some of the truncorum/luscus-group (arborum Verhoeff, 1928, bellus (Lignau, 1903), britanicus (Verhoeff, 1892), latestriatus (Curtis, 1845), parisiorum, and truncorum (Silvestri, 1896) and a group of large bodied species with similar gonopods consisting of caeruleocinctus (Wood, 1864), chalandei (‍Ribaut, 1904), finitimus, (‍Ribaut, 1905), ibericus, ‍Brölemann, 1913, londinensis (Leach, 1814) and sanctimichaelis Attems, 1927, (londinensis-group).

The genus Cylindroiulus on the Iberian Peninsula

Vicente (‍1981) listed 14 species of Cylindroiulus from the Iberian Peninsula: C. britannicus, C. chalandei, C. finitimus, C. franzi Attems 1952, C. ibericus, C. latestriatus, C. londinensis, C. perforatus, C. pyrenaicus Brölemann, 1897, C. sagittarius, Brölemann 1897, C. sanctimichaelis, C. silvarum (now C. punctatus Leach, 1815), C. spinosus and C. teutonicus (Pocock, 1900) (now C. caeruleocinctus). Of these, spinosus is now considered to be in the genus Allajulus (‍Read, 1990), but the list omitted Cylindroiulus dahli Demange, 1970, C. iluronensis Brölemann, 1912 and C. unciger Attems, 1952 as well as C. propinquus (Porat, 1870) and C. truncorum Silvestri, 1896 both from Portugal. In recent years this list has been added to by Cylindroiulus anglilectus Read, 2007, C. boreoibericus Read, 2007, C. gregoryi Read, 2007, C. fenestratus and C. ventanaea. All of the species listed above were listed for Spain in the European Atlas of Julids (‍Kime & Enghoff, 2017). Subsequently an additional species has been described, Cylindroiulus villumi. Spelda (‍2015) also provides a complete list of Cylindroiulus species from Spain including Macaronesia which includes an additional two species from the Iberian Peninsula, C. hispanicus Ceuca, 1974 and C. simplex Ceuca, 1974 but these have previously been synonymised with C. sagittarius and S. chalandei respectively.

Material and methods[Up]

Most of the material studied here was collected during a week-long visit by an international group of Myriapodologists who visited the Basque Country and Navarra, in 2009 and were assisted by various local biologists and foresters. The Sierra de Grazalema was visited by a German expedition, primarily collecting myriapods, in 2008 (see ‍Voigtländer & Reip, 2013 for more details). Additional material studied was from smaller collections and includes some unidentified Cylindroiulus in the collection of the NHMD and specimens collected by S. Pagola-Carte and I. Zabalegui, Basque entomologists.

Species in the genus Cylindroiulus tend to be very uniform in shape, are often of similar size, and are generally pigmented with a fairly standard pattern. Descriptions of colour or patterning are generally only given when they differ from the Cylindroiulus basic pattern. There are relatively few external characters that are generally useful for distinguishing species, making descriptions and identification of the species within the genus difficult. In the following descriptions the overall body shape is described using the terms anterior constriction, which is the reduction in body width behind the head in dorsal view. Posterior attenuation refers to the degree of reduction in body height at the posterior end of the body when looking from the side. The term body rings is used instead of segments as the rings in a millipede do not equate to true segments. Setae on the anal valves are generally evenly spaced and located just in from the margin, unless otherwise specified.

Locations given on labels were converted to a standard format (decimal degree) using coordinates-converter.com. For the small number where details of location from labels were vague, a map was used to find the decimal degree of the nearest place mentioned on the label. To produce the distribution maps Henrik Enghoff hand plotted them on the relevant species map from the European Atlas (‍Kime & Enghoff 2017) using https://mappingsupport.com/p2/gissurfer.php to help locate the positions.

Specimens were examined using a Nikon SMZ 2B binocular microscope and a Brunel Microscopes SP-1100 compound microscope. Photographs were taken using a Cannon EOS 100D using photoadapters from Brunel Microscopes and the stacking software Helicon Focus 7 was used. Photoshop was used for further picture editing.

Acronyms

The locations of type specimens are stated using the following acronyms.

Unless otherwise stated non type material is held as follows:

Specimens collected by S. Pagola and I. Zabalegui are in GEE

Specimens collected by Per Dursvoll are in ZMBN

Specimens collected by Steve Gregory and J. Paul Richards are in their private collections

Specimens collected by Karin Voigtländer are in SMNG

Specimens collected by Hans Reip and Norman Lindner are in the collection of Hans Reip

Specimens collected by Desmond Kime and Helen Read are in the private collection of Helen Read

Numbers in brackets before detailed descriptions of localities are the numbers given to locations during the collecting trips in 2009 and 2008.

Results[Up]

The collections studied yielded 10 species of Cylindroiulus. Some of these are well known with good descriptions published elsewhere but three new species were found which are described here. In addition, some species were felt worthy of fuller descriptions with the addition of more specimens, and the caeruleocinctus/sanctimichaelis/londinensis group has been discussed in more detail as they are challenging to distinguish. The abundance of some species has enabled comments to be made about life histories which have previously been unknown.

Specimens collected by Hans Reip and Norman Lindner from the Basque Country and Navarra were identified by Hans Reip.

Description of species

Cylindroiulus caramelos sp. nov.

urn:lsid:zoobank.org:act:B8498843-D247-4DE5-8BA4-A3A5100B8CBB

Figs. 1-‍18

Holotype: ♂, Spain: Cataluña: Tarragona, Alfara de Carles, El Toscar. 40.86, 0.334. 3.04.2015. J.D. Gilgado leg. (NHMD).

Paratypes: 4 ♀♀, details as above (NHMD).

Total material studied: male (1), females (4).

Diagnosis

Very similar to Cylindroiulus sanctimichaelis and Cylindroiulus caeruleocinctus, with large body size and lacking a telson projection. Differing by having a relatively long body with many body rings. Dark in colour, preserved specimens are banded in colour, with pale legs. Gonopods are similar to C. caeruleocinctus/C. sanctimichaelis, the anterior gonopods are relatively large in relation to the posterior gonopods which have a large brachite.

Description

Length: ♂ 38 mm; ♀ up to 51 mm. Body height: ♂ 2.1 mm; ♀ up to 3.06 mm. Rings: ♂ 57; ♀ up to 62. Body colour: General appearance, black with white legs (Figs. 1-‍5). Head dark brown, prozonites pale grey, metazonites black, with paler posterior edge. Thin black mid dorsal line. Appearing striped around the body (but only preserved specimens seen). Anal valves very dark brown, preanal ring dark brown/black with white posterior margin. Overall body shape: Anterior constriction, present, just behind the head, posterior attenuation present but not strong, largely just the apodous rings. Head: Antennae, dark brown, eye rows: clear and readable. Stadia: ♂ 9 rows of ommatidia = X; ♀ up to 12 rows = XIII. Body rings: Metazonite vaulting present and clearly noticeable, limbus not crenulate. Legs: Pale in colour. Despite the preservation of the holotype being of good quality, almost all the legs had broken accessory claws. Of the few that appeared unbroken, the accessory claws seemed to be slightly shorter than the claw (Fig. 6). Females also had a large number of broken accessory claws, with those intact appearing to be slightly shorter than the claw. Telson: Preanal ring with no projection, perhaps bearing setae but probably worn off in the male. Subanal scale not projecting, with one pair of setae. Anal valves with three pairs of setae.

Mature male secondary sexual characters: Mandibular stipites of mature males strongly expanded and projecting anteriorly, First pair of legs in mature male tight hooks with prominent ‘knees’ on the angle of the hook.

Gonopods (Figs. 8-‍15): Promerite narrow and slender in anterior view, relatively bulky in mesal view, with hooked apex. Mesomerite narrow and slender and almost completely enclosed by promerite apart from the distal part. Flat topped and with different textured apex. Opisthomerite broad and bulky in mesal view. Brachite large, projecting beyond the solenomerite, with angular ventral margin and clear cleft between it and the solenomerite with membranes across it. Phylacum large, curvaceous. Solenomerite with three small teeth at the apex and membranous area at tip. Paracoxal rim very high, paracoxal process lacking.

Etymology

Named for its superficial resemblance to a mint humbug (a stripy child’s sweet); “caramelos” are sweets in Spanish.

Relationships with other species

This species clearly falls into the problematic londinensis group, for full discussion see under C. sanctimichaelis. It can be distinguished by details of the gonopods and has more body rings in total and more body rings for the body height than other similar species (Figs. 16-‍17). The short accessory claw might also be diagnostic but most appeared to be broken so the reliability of this character is uncertain. It is also not well enough studied in all the similar species to be confident of its value.

Habitat

The type locality is situated in a limestone mountainous region range close to Tarragona with a Mediterranean climate and occasional snow. The site is situated in a humid area (Fig. 7) which is the source of many small water courses in the shadow of the mountain. The mountains are considered a transition zone between the Sistema Ibérico Mountains and the Catalan pre coastal mountain range but are separated from the Pyrenees.

Life history

Adults were found in April. There are not enough specimens to analyse for life history.

Distribution

Only known from the type locality (Fig. 18). When the manuscript was in the proof stage, communication with Hans Reip revealed that this species may be quite common west and south of the Ebro delta, a little south of the type location.

Cylindroiulus elosegiorum sp. nov.

urn:lsid:zoobank.org:act:AAED2DE2-B26D-45DB-B883-D43FB4A2EC39

Figs. 19-‍39

Holotype: ♂, Spain, Navarra: Leitza, Ariz Mendiak, at area “Kornieta”, 43.0698, -1.8823, 910 m, 4.02.2010, Fagus L. woodland, poor understorey, rocky, siliceous soils, mosses, Helleborus L. under bark & in leaf litter. Helen Read & Mark Frater leg. (MNCN 20.07/2072).

Paratypes:

5♂♂, 1 j♂, 14♀♀/j, details as above. (MNCN 20.07/2073-2092).

(7 & 8) Navarra: Leitza, Ariz Mendiak, at area “Kornieta”, 43.0698, -1.8823, 910 m, 20.04.2009, Fagus woodland, poor understorey, rocky, siliceus soils, mosses, under bark & in leaf litter. 3♂♂, 16♀♀. Desmond Kime (NHMD) & Steve Gregory (NHM) leg.

(9) Navarra: Leitza, town area, 43.0788, -1.9161, 470 m, 20.04.2009, house & garden around casa rural Aztieta under stones and logs. 2♂♂. Steve Gregory leg. (GEE).

(12b) Navarra: Sierra de Aralar, south of Errazkin, north of area “Axkarateko Malkorra”, 42.9972, -1.9703, 630 m, 21.04.2009, Fagus woodland. 7♂♂, 12♀♀, Karin Voigtländer leg. (SMNG VNR 014759-‍8).

(22) Navarra: Sierra de Aralar, south Baraibar, on road NA-7510 to Santuario de San Miguel, at area “Izáin”, 42.9714, -1.9384, 790 m, 22.04.2009, Karst area with deep grykes and bare limestone rocks, Fagus woodland & some Crataegus Tourn. Ex L. bushes in leaf litter, partly sieved out. 1♂, 4♀♀. Helen Read leg. (NHMD)

(23) Navarra: Sierra de Aralar, south Baraibar, on road NA-7510 to Santuario de San Miguel, at area “Izáin”, 42.9724, -1.9361, 730 m, 22.04.2009, Karst area with deep grykes and bare limestone rocks, Fagus woodland & some Crataegus bushes ground of deep and collapsed gryke, sieved from Fagus leaf litter. 2♂♂, 18 ♀♀, Karin Voigtländer leg. (SMNG VNR 014771-‍13).

(24B) Navarra: Sierra de Aralar, Sierra de San Miguel, mountain point Artxueta at radio mast, 42.9525, -1.9668, 1300 m, 22.04.2009, Karst area with bare limestone rocks, grove of low growing Fagus in litter and under bark. 1♂, 1j♂, 2♀♀, Helen Read leg. (GEE)

(17-‍1) Guipuzkoa: Sierra de Aralar, Beasain, road from Lazkao to Etxarri-Aranaz, westerly Pass, 42.9572, -2.1122, 550 m, 21.04.2009, forest of Fagus in leaf litter. 5♂♂, 1j♂, 11♀♀, Karin Voigtländer leg. (SMNG VNR 014831-‍2).

(29) Navarra: Between Sierra de Urbasa and Sierra de Andía, road (NA-120) 2,5 km SSW of Puerto de Lizarraga, “Fagadierer” (300 m E of Bentaberri), 42.8372, -2.0233, 860 m, 22.04.2009, Mediterranean Buxus L.-Fagus-Forest within leaf litter of Fagus. 1♂, 1j♂, 3♀♀. Steve Gregory leg. (ZMBN).

(34) Navarra: Sierra de Urbasa, on top, near road NA-7182, 42.8375, -2.1815, 930 m, 23.04.2009, grazed Fagus woodland, in parts with bushes of Crataegus, otherwise few bushes, partly covered with herbs: Corydalis DC., Urtica L., Rubus L. etc., very little dead wood on the ground, in leaf litter, mainly Crataegus. 1♂, 2♀♀, Karin Voigtländer leg. (SMNG VNR 014820-‍4).

(35) Navarra: Sierra de Urbasa, on top at southern cliff border, east of road NA-7182, 42.7989, -2.1417, 930 m, 23.04.2009, pasture on stony ground, some thorny bushes, occasional trees or groups of Fagus. 1♀, Karin Voigtländer leg. (SMNG VNR 014826-‍6).

(35) As above, in leaf litter. 1♂, 3♀♀/J, Helen Read leg. (NHMW).

(36) Navarra: Sierra de Urbasa, at southeastern cliff border, eastern zigzag of road NA-7182, under cliffs, 42.7953, -2.1397, 810 m, 23.04.2009, steep slope with forest of Fagus, thick leaf layer, thick humus layer (no more raw humus) – in the herb layer: Hepatica nobilis Schreb., Scilla L. sp., at the cliff: Saxifraga Tourn. Ex L. sp., Asplenium cf. alduterinum Milde, in thick leaf litter. 3j♂, 11♀♀, Karin Voigtländer leg. (SMNG VNR 014781-‍25).

(38) Navarra: Sierra de Urbasa, on top, near road NA-7182, 42.8411, -2.18, 920 m, 23.04.2009, Fagus woodland on limestone karst under stones and in leaf litter. J. Paul Richards (3♂♂, 6♀♀/J, SGMT: accession number SHEFM:2021.2) & Helen Read (1♀, 1j♂, NHMW) leg.

(45) Guipuzkoa: Montes de Altzania, Sierra de Urquilla, south of mountain chain Aikorriko Mendikatea, southwest of Mountain Aitzgorri - north of end of road from Salvatierra to Ordonana, Zalduondo, Zumarraundi - north slope of Aldasbarreneta valley, 42.9411, -2.3339, 1170 m, 24.04.2009, forest of Fagus, very mossy, almost no herbs, isolated fern and Daphne L. in leaf litter. 2♂♂, 2j♂, 10♀♀, K. Voigtländer leg. (SMNG VNR 014787-‍13). (SMNG).

Other specimens:

(7) Navarra: Leitza, Ariz Mendiak, at area “Kornieta”, 43.0698, -1.8823, 910 m, 20.04.2009, Fagus woodland, poor understorey, rocky, siliceous soils, mosses, under bark & in leaf litter. 3♂♂, 16♀♀, Helen Read leg. (specimens dried in tube).

(11) Navarra: Sierra de Aralar, 1 km ESE Betelu: at the road junction N-130 & NA-7514 (and at the confluence of the streams “Errazkinko Erreka” & “Ezpeleta Erreka”), 43.0214, -1.9656, 270 m, 21.04.2009, steep slope above flood area, below road – river bank wood, Platanus L., Fraxinus excelsior L., Alnus Mill., Salix L. 8♀♀/J, Hans Reip. leg. & det.

(29) Navarra: Between Sierra de Urbasa and Sierra de Andía, road (NA-120) 2,5 km SSW of Puerto de Lizarraga, “Fagadierer” (300 m E of Bentaberri), 42.8372, -2.0233, 860 m, 22.04.2009, Mediterranean Buxus-Fagus-Forest within leaf litter of Fagus. 2♀♀ Norman Lindner leg. Hans Reip det.

(30) Navarra: Sierra de Andía, at road NA-7330 (from Lezáun to Arizaleta), at area “Los Ristros”, ca. 500 m WNW Arizaleta, 42.7589, -1.9833, 700 m, 22.04.2009, Mediterranean Quercus L. forest (evergreen), used as wood pasture, incl. some small areas with dry meadows (with small thorny bushes, some bigger stones, some small Buxus bushes, Quercus ilex L., Genista L., shrubby Thymus L., shrubby Lonicera L., Sedum L., grasses). 2♀♀. J. Paul Richards leg.

(35) Navarra: Sierra de Urbasa, on top at southern cliff border, east of road NA-7182, 42.7989, -2.1417, 930 m, 23.04.2009, pasture on stony ground, some thorny bushes, occasional trees or groups of Fagus. 2♂♂, 2♀♀, 1j. Hans Reip leg. & det.

(36) Navarra: Sierra de Urbasa, at southeastern cliff border, eastern zigzag of road NA-7182, under cliffs, 42.7953, -2.1397, 810 m, 23.04.2009, steep slope with forest of Fagus, thick leaf layer, thick humus layer (no more raw humus) - in the herb layer: Hepatica nobilis, Scilla sp., at the cliff: Saxifraga sp., Asplenium cf. adulterinum Milde, Saxifraga in thick leaf litter. 1♂, 1♀, Norman Lindner leg. & 3♂♂, 5♀♀, 3j, Hans Reip leg. Hans Reip det.

(41) Navarra: Sierra de Urbasa, on top, 1km east of road junction to road NA-7182 at site “Bentakaita”, 42.854, -2.1595, 890 m, 23.04.2009, Fagus forest, Prunus, Corydalis, Mercurialis L., Anemone L., Erythronium L. in leaf litter. 1♀. Desmond Kime leg.

(45) Guipuzkoa: Montes de Altzania, Sierra de Urquilla, south of mountain chain Aikorriko Mendikatea, southwest of Mountain Aitzgorri - north of end of road from Salvatierra to Ordonana, Zalduondo, Zumarraundi - north slope of Aldasbarreneta valley, 42.9411, -2.3339, 1170 m, 24.04.2009, forest of Fagus, very mossy, almost no herbs, isolated fern and Daphne in leaf litter. 1j Hans Reip leg. & det.

(52) Alava: Sierra de Peña Gorbea [Gorbeia], Valley of the River “Bayas Ibaia”, north of Sarria, at (bridge) Arkarai, at area “Eczaray”, 43.017, -2.83, 720 m, 24.04.2009, slope with coniferous forest, mainly without herb layer, barely rotten wood, some dead wood pieces, on the lower edge some medium sized stones on the ground under bark of (freshly) dead wood or stumps. 2♂, 1j♂, 1♀. Desmond Kime & Steve Gregory leg.

Material identified by Hans Reip, no mature males but highly likely to be this species:

(48) Guipuzkoa: Montes de Altzania, Sierra de Urquilla, south of mountain chain Aikorriko Mendikatea, southwest of Mountain Aitzgorri – north of end of road from Salvatierra to Ordonana, Zalduondo, Zumarraundi – Portua Zarra, west of Eskaratza, 42.9334, -2.3283, 1200 m, 24.04.2009, small sharp summit of limestone rock/cliff, Fagus forest with small accumulations of leaves in leaves between rocks. 1J, Hans Reip leg.

(49) Alava: Montes de Altzania, Sierra de Urquilla, south of mountain chain Aikorriko Mendikatea, southwest of Mountain Aitzgorri – north of end of road from Salvatierra to Ordonana, Zalduondo, Zumarraundi – north of Zumarraundi, deep gryke with stream discharge, in the lower third, 42.9241, -2.3224, 980 m, 24.04.2009, forest of Fagus, thick leaf layer in leaf litter. 1♀, Hans Reip leg.

Total material studied: males: 39; juvenile males: 11; females: 120

Diagnosis

A small pale Cylindroiulus with exceptionally strongly down pointed telson projection and large upward pointing subanal scale. Reduced number of ommatidia, in jumbled rows, often variable in size. Gonopods relatively simple and conforming to the normal Cylindroiulus plan.

Description

Length: ♂ 8.9-16.6 mm; ♀ up to 19.2 mm. Body height: ♂ 0.7-0.9 mm; ♀ up to 1.4 mm. Rings: ♂ with 27-‍43 podous rings. ♀ up to 50. Body colour: Generally pale straw coloured with little pigmentation (Figs. 19-‍20). Some specimens darker posteriorly, probably the result of gut contents showing through the paler integument. Repugnatorial glands, when visible, pale brown/orange. Overall body shape: Anterior constriction not obvious, posterior part of trunk wider than anterior. Head: Antennae: slender and pale in colour. Eye rows: Eyes not in clear rows except in younger juveniles, difficult to ‘read’. Ommatidia reduced in number, often jumbled and sometimes varying in size in the same ocular field (Fig. 21). Ommatidia black and contrasting in comparison with pale body colour. ♂ with 8-‍14 ommatidia; F up to 15-‍16 ommatidia. Body rings: Striae on metazonites fairly strong and close together, limbus crenulate (Fig. 22). Legs: Generally pale, similar to body colour. With strong and crooked setae on more basal leg segments. Claw of normal shape; accessory claw slightly longer than claw, with distinct change of direction after surpassing the claw tip (Fig. 23). Telson: Preanal ring with strong projection, curved ventrally. With 1 pair of setae at end and another pair ventrally. Less strongly projecting in juvenile individuals and tending to be straighter/less down turned. Subanal scale projecting well past the anal valves (Figs. 20, 24). More-or-less parallel withventralmarginof trunk but sometimes appearing dorsally directed giving the impression of a pincer shape at the posterior end of the animal. With one pair of setae. Smaller relative to body size in juveniles (Fig. 25). Anal valves: With three pairs of setae (Fig. 24).

Mature male secondary sexual characters: Mandibular stipites of mature males barely expanded (Fig. 21). First pair of legs in mature males forming tight hooks with prominent ‘knee’ (Fig. 26). Keel of body ring 7 in mature males open, but not very wide, no obvious keel (Fig. 27)

Gonopods (Figs. 28-‍33): Promerite: Short and squat with no ‘window’. Flagellum of normal length. Mesomerite: Short and squat, similar in length to promerite. Opisthomerite: Simple in shape with membranous tip. Long thin phylacum and small brachite. Large paracoxal process which seems a slightly strange shape from some angles and in posterior view.

Etymology

Named after the Elosegi family from Leitza, in particular José Miguel, Migel Mari and Arturo, all of whom have helped me explore the Basque area and given freely of their time and knowledge. Note that members of the family spell their name in two different ways, Elósegui and Elosegi. The latter is the ‘original’ Basque spelling and the former the Spanish version. The family have expressed their preference for the Basque spelling.

Relationships with other species

This species is one of a small group that have strongly down pointed dorsal projections on the preanal ring and strongly pointed subanal scale (although it is perhaps the most extreme in this species), others being C. apeninnorum (‍Brölemann, 1897) (Italy, also presumably introduced to the Netherlands and UK), C. gestri (Silvestri, 1898) (Sardinia), C. propinquus (Portugal) and C. pyrenaicus (Spain). The gonopods however do not appear similar to any of these species. It is unknown what the function of a strongly pointed telson is, and this is also true of the subanal scale. Pointed telsons occur in a variety of Cylindroiulus species and it seems likely that this does not relate to any particular evolutionary relationships.

Life history

Cylindroiulus species add the apodous body rings to the number of podous body rings at each moult, and, if they have ommatidia they also add eye rows each time so it is possible to infer details of their life history by looking at the distribution of these characteristics across the collected specimens (‍Read, 1988). The eye rows for this species are very jumbled and are also reduced in terms of numbers of ommatidia (in comparison to the total, assuming that an extra row of ommatidia is added at each moult). As part of the characterisation of this species the numbers of ommatidia were counted and a plot showing the number of individuals for each ommatidia number, broken down by sex is shown in Figs. 35-‍36. Sometimes the number of ommatidia was not easy to determine and for a few specimens both sides of the head were examined. When the number of ommatidia varied on each side the head the lowest number was used in the graphs. Similar graphs illustrate the variation in the number of podous rings broken down by sex (Figs. 37-‍38). These figures show that the mature males have a relatively tight cluster of possible podous ring numbers. Allowing for the relatively high number of apodous rings in this species, between 3 and 5 for individuals with 39 or 40 podous rings, this probably reflects just one or two stadia. Aside from one very small mature male (perhaps misdesignated?) the distribution seems to showanormalpattern of juvenile males developing into matures for both number of ommatidia and podous ring number.

The graphs for the females show a more extended ‘tail’, i.e. more individuals with larger numbers of ommatidia and podous rings than the males. There is therefore a suggestion that the females undergo further moults once maturity has been reached. This situation is not uncommon in the genus Cylindroiulus. It therefore appears that the males are probably semelparous, but the females may be iteroparous. Having eight apodous rings was not unusual amongst juveniles of this species, hence the gaps in the graphs reflects this development pattern.

Distribution

This species was found quite widely and frequently over the area sampled in 2009, in the provinces of Navarra and Guipuzkoa and just into Alava. A species of the hills, it was found in several mountain ranges from Leitza in the east, across Sierra de Aralar to Gorbeia in the west, and south to Sierra de Urbasa and Sierra de Andia (Fig. 39). While many locations were deciduous woodland, frequently with Fagus sylvatica L. (Fig. 34), it was also found in stony wood pasture with scattered trees and in the garden of a house in the centre of the town of Leitza.

Cylindroiulus franzi Attems, 1952

Figs. 40-‍46

Material studied:

(9) West side of the Sierra de Grazalema; banks of the river Río del Bosque, hiking trail between El Bosque and Benamahoma; around 36.772, 5.481; 370 m; narrow and deeply incised valley, gallery forest; at humid sites in leaf litter. Sieving. 05.02.2008. 4♂♂, 24 ♀♀/J, Hans Reip & K. Vöigtlander leg. (SMNG).

Total material collected: male (4), females (24)

Diagnosis

A small and pale species with no telson projection but a characteristic tented shape to the top of the telson. Simple shaped gonopods with a prominent paracoxal process which is jagged on the posterior side.

Description

Length: ♂ 17 mm; ♀ up to 20 mm. Body height: ♂ 1 mm; ♀ up to 1.3 mm. Rings: ♂ 48-‍51, ♀♀ up to 54. Body colour: Overall pale in colour, telson a little darker, some individuals with small amount of darker mottling. Antennae and legs pale in colour. Overall body shape: Anterior constriction behind head; posterior attenuation only of apodous rings. Legs with accessory claw slightly longer than claw. Head: Eye rows rather jumbled with what look to be extra ommatidia (Fig. 40). First two rows often seemingly a single ocellus. Stadia: ♂ VII to VIII, ♀ up to IX. Body rings: Striae on metazonites regular. Limbus not crenulate. Telson: Preanal ring without clear dorsal projection but with a roof-like shape which gives the appearance of a prominent point dorsally (Fig. 41). Bearing 1 or 2 pairs of setae ventrally and one pair dorsally on the projection. Subanal scale not projecting, bearing 1 pair of setae. Anal valves generally with 3 pairs of setae.

Mature male secondary sexual characters: Mandibular stipites of mature males expanded to medium extent, with a vertical anterior margin (Fig. 40). First pair of legs in mature male: tight hooks. Keel of body ring 7 in mature male very small, barely projecting but body appearing slightly wider at this point.

Gonopods (Figs. 42-‍45): Promerite and mesomerite relatively low and squat. Promerite slightly longer and much broader than mesomerite which appears narrowly projecting up almost through the promerite. Flagellum of normal length. Paracoxal rim barely present. Opisthomerite simple in shape, with hyaline region apically. Paracoxal process large and with jagged posterior margin, although rather less stout than illustrated by Mauriès (‍1978). Despite the size of the paracoxal process it can be difficult to see in situ being situated on the mesal side of the gonopods.

Relationships with other species

Attems considered that this species was in the subgenus Plagionicus, together with C. parisiorum and C. bouvieri (‍Brölemann, 1896) (now = C. parisorum Geoffroy et al. in press), but the simple gonopods look more similar to the Caucasian species C. placidus (Lignau, 1903) and C. besucheti Strasser, 1975 both considered in the C. placidus-group (‍Read, 1992). The original illustrations are rather poor, but a more complete description has been made by Mauriès (‍1978) including a drawing of the type. The current specimens accord well with that of Mauriès loc cit.

Ecology & life history

With so few samples further analyses are not possible.

Distribution

Previously known from Sierra de las Nieves close to Ronda and Mt Arastepa in the Sierra de Ronda. The current larger collection is from the same area. If the collection from Granada can be confirmed as the same species (see below) this would be a considerable expansion to the range, although still within Andalucía. (Fig. 46).

Cylindroiulus cf. franzi

No mature males were in these collections. The telson and overall appearance looks similar but location of one is somewhat distant from those recorded so far.

Material studied

Andalucía: P.N. Sierra de las Nieves, road to Quejigales. 23.3.2008. 1♀, H. Read & M. Frater leg.

Andalucía: Granada, Sierra Nevada, Güéjar Sierra, Barranco de las Ánimas, 1400 m, 14.8.2014. 1♀, 1j♂, J.D. Gilgado & A. Tinaut leg (NHMD).

Cylindroiulus karinhansorum sp. nov.

urn:lsid:zoobank.org:act:222887A7-67E9-418B-9569-68E736AE1F58

Figs. 47-‍59

Holotype: ♂, Spain, (21) Andalucía: Northern part of Los Alcornocales; south of the road from Cortes de la Frontera to Alcalá de los Gazules, forest track south of the mountain Peñón del Berrueco. 36.593, -5.423, altitude ca. 680 m, 8.02.2008. Hans Reip & K. Vöigtlander leg. (SMNG).

Paratypes: 1♂, 6j♂♂, 26♀♀/J, details as above. (SMNG).

1♂, 2♀♀, details as above (NHMD).

(11) Northern part of Los Alcornocales; south of the road from Cortes de la Frontera to Alcalá de los Gazules, east of the turn-off to Ubrique, south of the mountain Peñón del Berrueco, Casa de los Llanos, wayside; 36.614, -5.421, 5.02.2008. 740 m; Quercus suber L. forest, loose soil, slightly humid; under dead wood and stones, hand sampling. 1♂, 1j♂, 5♀♀. Hans Reip & K. Vöigtlander leg. (SMNG).

(13) Zahara de la Sierra and Grazalema, near the mountain área Los Cambroneros/Los Pilones, above Garganta Verde. 36.814, -5.403, 6.02.2008, 560 m, Nerium L., Ceratonia siliqua L, Quercus. Cool and humid, in leaf litter. Sieving. 1♀. Hans Reip & K. Vöigtlander leg. (SMNG)

(19) Sierra de Grazalema, hiking trail between Ubrique and Benaocaz (Calzada Romana), 36.689, -5.439, 8.02.2008, 470 m, meadow with diverse shrubs, under stones. Hand sampling. 2♂♂, 1♀, Hans Reip & K. Vöigtlander leg. (SMNG).

(20) Northern part of Alcornocales, south of the road from Cortes de la Frontera to Alcalá de los Gazules, forest way south of the mountain Peñón del Berrueco. Between 36.606, -5.419 and 36.605, -5.423, 8.02.2008, 780 m, open oak woodland (cork and other oaks), under bark, hand sampling. 1j♂, 6♀♀, Hans Reip & K. Vöigtlander leg. (SMNG).

Total material collected: males (6), juvenile males (8), females (41).

Diagnosis

A medium sized species, generally pale in colour. With slightly, but definitely, projecting telson and relatively simple gonopods.

Description:

Length: ♂ 13 mm; ♀ up to 24 mm. Body height: ♂ 0.8-1.0 mm; ♀ up to 1.5 mm. Rings: ♂ 37-‍44+3-4 apodous (one male apparently has 49 rings but is broken into three pieces so it is not entirely clear if they all belong to the same individual); ♀ to 59. Body colour: Generally pale in colour but variable with some individuals darker brown in colour, adults with typical Cylindroiulus colouring though rather faint. Some darker dorsally. Juveniles with almost pink sheen. Overall body shape: Anterior constriction present but not strong. Head: Antennae pale in colour. Eye rows rather mixed up and difficult to read the lines, some individuals easier than others and some showing substantial variation between the two sides of the same individual. Several appear to have extra ommatidia. Stadia: ♂ VII–VIII; ♀ to IX. Body rings: Metazonites with strong and even striae, vaulting almost absent, posterior attenuation only slight, of apodous rings. Limbus not crenulate. Legs: Pale in colour, with accessory claw substantially longer than claw (Fig. 55). Telson: Preanal ring clearly projecting but not long and not extending beyond the anal valves (distinct from that of franzi which is very short), chestnut brown in some specimens (Fig. 47). With 2-‍3 pairs of setae, 1-‍2 ventrally and one on the telson projection but not at the tip and set back a short distance. Subanal scale not projecting, with 1 pair of setae. Anal valveswith 1-‍3 pair of setae.

Mature male sexual characters: Mandibular stipites of mature males expanded but not greatly, anterior margin more or less vertical (Fig. 48). First pair of legs in mature males tight hooks. Keel of body ring 7 in mature males slightly projecting around a large gap in the body wall.

Gonopods (Figs. 49-‍54): Promerite slightly shorter than mesomerite, flagellum of normal length. Opisthomerite simple in shape, parallel sided and lacking brachite but with possible slight phylacum. Apex with hyaline structure. Paracoxal process sharp, clearly defined but not extensive. Paracoxal rim not particularly large.

Etymology

Named in honour of Karin Voigtländer and Hans Reip who collected the type material, contributed many other specimens to this study and who have both contributed greatly to our knowledge of millipedes.

Relationships with other species

Cylindroiulus karinhansorum sp. nov. does not easily fit into a Cylindroiulus species group. The relatively simple gonopods, similar to the Cylindroiulus ground plan make it difficult to comment on its taxonomic position; they are not dissimilar to C. franzi in overall appearance.

Ecology & life history

Six females from the type locality had fungi on their first two pairs of legs (Fig. 56) which was identified as Rickia laboulbenioides De Kesel by Henrik Enghoff. The species seems to have been found in wooded areas in the mountains, often with Quercus suber.

The stadial spectra of this species is given in Figs. 57-‍58. The eye rows were not clear enough to consider reliable, so the graphs have been drawn using the number of podous rings. Generally mature males had up to four apodous rings, the graph thus suggests that the mature males probably represent two different stadia, suggesting that they are semelparous. As expected, the females had a wider range of numbers of podous rings, with one having 59, substantially more than the others. There are hints that some females, at least might be iteroparous.

Distribution

Discovered from several locations in the Grazalema mountains and nearby in P.N. de los Alcornocales (Fig. 59).

Cylindroiulus londinensis (Leach, 1814)

Material studied

(3A) Navarra: Leitza, Ariz Mendiak, between area “Ustarleku” and “Karobieta”, above side stream to Gorriztaran, 43.0778, -1.8775, 615 m, 20.04.2009, grove of Castanea Mill., pollard trees on the slope – Ranunculus ficaria L., Daphne, Helleborus, Salvia, Rubus, Lathrea L. - soil loamy and calcareous. 2♂♂, 3♀♀, K. Voigtländer (2♂♂, 1♀ SMNG VNR 014744-‍13 & SMNG VNR 014746-‍12), J. Paul Richards, Helen Read (2♀♀, MNCN 20.07/2096) & Hans Reip (1♂, 2j) Reip-E517-F2449 leg.

(7 & 8) Navarra: Leitza, Ariz Mendiak, at area “Kornieta”, 43.0698, -1.8823, 910 m, 20.04.2009, Fagus woodland, poor understorey, rocky, siliceus soils, mosses, Helleborus, under bark & in leaf litter. 2♂♂, 2♀♀, Steve Gregory, Desmond Kime (1♂, MNCN 20.07/2095) & Helen Read (1♂) leg.

(11 & 12) Navarra: Sierra de Aralar, south of Errazkin, north of area “Axkarateko Malkorra”, 42.9972, -1.9703, 630 m, 21.04.2009, wood of Fagus trunk, dead wood. 2♂♂, 3♀♀, Helen Read (1♀, GEE) , Per Djursvoll & Karin Voigtländer leg. (1♂, SMNG VNR 014758-‍13).

(15) Guipuzkoa: Sierra de Aralar, Tolosa, 500 m south of Bedaio/Goikoa, 43.0494, -2.04, 420 m, 21.04.2009, farm buildings & stone wall, logs, wood, stones and rubble under stones and logs. 3♂♂, 10♀♀J. Helen Read leg.

(17) Guipuzkoa: Sierra de Aralar, Beasain, road from Lazkao to Etxarri-Aranaz, western Pass, 42.9572, -2.1122, 550 m, 21.04.2009, forest of Fagus. 1♂, 1♀ (Reip-E534-F2506). Hans Reip leg.

(18) Guipuzkoa: Sierra de Aralar, Beasain, road from Lazkao to Etxarri-Aranaz, Pass Puerto de Lizarrusti, 42.9614, -2.0983, 690 m, 21.04.2009, forest of Fagus, on slope under stones & in leaf litter. 3♂♂, 5♀♀/J. Karin Voigtländer (1♂, SMNG VNR 014764-‍11), Hans Reip (1♂, Reip-E535-F2513-AS183) & Norman Lindner leg.

(21) Navarra: Sierra de Aralar, south Baraibar, on road NA-7510 to Santuario de San Miguel, at area “Urkia”, 42.9762, -1.9318, 710 m, 22.04.2009, wet wood pasture, Corylus L. with some Quercus, Ilex, Ranunculus ficaria, Daphne, Oxalis L., Helleborus, Euphorbia L., Geranium L., Lathrea in leaf litter. 1♂, 11♀♀. Helen Read (GEE) & Hans Reip (1♂, 2j, Reip-E540-F2524 & Reip-E542-F2528) leg.

(22) Navarra: Sierra de Aralar, south Baraibar, on road NA-7510 to Santuario de San Miguel, at area “Izáin”, 42.9714, -1.9384, 790 m, 22.04.2009, Karst area with deep grykes and bare limestone rocks, Fagus woodland & some Crataegus bushes in leaf litter, partly sieved out. 1♀ (Reip-E543-F2533), Hans Reip leg.

(24) Navarra: Sierra de Aralar, Sierra de San Miguel, mountain point Artxueta at radio mast, 42.9525, -1.9668, 1300 m, 22.04.2009, Karst area with bare limestone rocks, pasture, under stones and dung. 3♀♀, Per Djursvoll leg.

(26) Navarra: Sierra de Aralar, Sierra de San Miguel, southern lower slope at Iraneta, 42.9258, -1.9597, 500 m, 22.04.2009, dry Quercus wood in leaf litter. 1♂ (Reip-E546-F2553), Hans Reip leg.

(36) Navarra: Sierra de Urbasa, at southeastern cliff border, eastern zigzag of road NA-7182, under cliffs, 42.7953, -2.1397, 810 m, 23.04.2009, steep slope with forest of Fagus, thick leaf layer, thick humus layer (no more raw humus) – in the herb layer: Hepatica nobilis, Scilla sp., at the cliff: Saxifraga sp., Asplenium cf. adulterinum, in thick leaf litter. 2 ♂♂, 2♀♀ (Reip-E557-F2586-AS181/2), Hans Reip leg.

(48) Guipuzkoa: Montes de Altzania, Sierra de Urquilla, south of mountain chain Aikorriko Mendikatea, southwest of Mountain Aitzgorri – north of end of road from Salvatierra road to Ordonana, Zalduondo, Zumarraundi – Portua Zarra, west of Eskaratza, 42.9334, -2.3283, 1200 m, 24.04.2009, small sharp summit of limestone rock/cliff, Fagus forest with small accumulations of leaves in leaves between rocks. 5♀♀, Karin Voigtländer (4♀♀, SMNG VNR 014791-‍9) & Hans Reip (1♀, Reip-E563-F2610) leg.

(52) Alava: Sierra de Peña Gorbea [Gorbeia], Valley of the River “Bayas Ibaia”, north of Sarria, at (bridge) Arkarai, 43.017, -2.830, 720 m, 24.04.2009, semi-open stream valley with old Fagus trees and bushes, with pasture beside river, Erica L., Ulex L., Ilex L., Prunus L. in leaf litter/humus. 1♂, 2♀♀, Steve Gregory leg.

(55) Alava: south Salvatierra, Campezo, near village Oteo, 42.726, -2.3628, 710 m, 25.04.2009, Quercus faginea Lam. Wood; Q. ilex, Arbutus unedo L., Viburnum lantana L., Juniperus L. under stones and leaf litter. 3♂♂, 5♀♀/J, Steve Gregory, Desmond Kime, J. Paul Richards & Helen Read (1J GEE).

Navarra: San Miguel de Aralar, Huarte-Araquil, 42.89, -2.02, 1250 m, 28.06.2007. 1♂, I. Zabalegui leg.

Gipuzkoa: Hernani, Aiako Harria P.N., Ugaldetxo, 43.22, -1.93, 50 m, 29.06.2007. 1♂, 1♀, 1J, I. Zabalegui leg.

From elsewhere in Spain: Zaragoza: Longás. 1♂, 1♀. 14.05.2015. J.D. Gilgado leg. (NHMD)

Total material studied: males: 23; females/juveniles: 54

Distribution

The additional records do not add greatly to the known distribution for C. londinensis (Fig. 60) which appears to be on the edge of its range on the Iberian Peninsula.

Notes

In a few locations only females were found but the shape of the telson coupled with the size of the specimens allowed placement within this species with reasonable certainty.

One Cylindroiulus londinensis had an expanded cheek plate and hook shaped first leg pair but one pair of normal walking legs on body ring 7. The combination of these characteristics is not suggestive of an intercalary male (these usually have a very short first pair of legs that are not hook shaped and a cheek plate intermediary between a mature male and a juvenile). Presumably this individual is just an aberrant example or a gynandromorph. It was not examined for vulvae as is now in the collection of S. Gregory.

Cylindroiulus cf. londinensis

Material studied

Navarra: Selva de Irati, 22.06.2013, Fagus sylvatica forest, 1♀, V. Martínez-Pillardo leg. (NHMD).

This female was exceptionally large in size with a body height of 5 mm. Probably 12 rows of ommatidia (= stadia XIII) and 45+1 body rings. The preanal process was distinct but not especially long. Another exceptionally large female was found at location (7) Leitza, area Kornieta. At this locality mature males of C. londinensis were also found, adding to the likelihood of these specimens being this species but just exceptionally large sized individuals.

For further discussion of this species see under Cylindroiulus sanctimichaelis below.

Cylindroiulus punctatus (Leach, 1815)

Material studied

(2) Navarra: Pamplona, south of Airport, near Salinas de Pamplona, at Río Elortz, 42.7539, -1.6431, 440 m, 19.04.2009, edge of deep stream valley, meadow slope. 1♂, 2♀♀, Hans Reip leg.

(3) Navarra: Leitza, Ariz Mendiak, between area “Ustarleku” and “Karobieta”, above side stream to Gorriztaran, 43.0778, -1.8775, 615 m, 20.04.2009, grove of Castanea, pollard trees on the slope – Ranunculus ficaria, Daphne, Helleborus, Salvia, Rubus, Lathrea – soil loamy and calcareous. 2♀♀/J, J. Paul Richards & Steve Gregory leg.

(9) Navarra: Leitza, town area, 43.0788, -1.9161, 470 m, 20.04.2009, house & garden around casa rurale Aztieta under stones and logs. 1♀, Steve Gregory leg.

(31) Navarra: Sierra de Andía, Valle de Goñi, 500 m NE Goñi at area “Mustillogorri”, 42.855, -1.8992, 930 m, 22.04.2009, grazed deciduous Quercus woodland, within agricultural areas (pastures) under stones. 1♂, 3♀♀, Steve Gregory & J. Paul Richards leg.

Although only females were found in Leitza the shape of the telson projection of this species is characteristic and it seems highly likely that the specimens found were this species.

Distribution

This widespread north-western European species just extends into the Iberian Peninsula south of the Pyrenees. The current collection confirms its presence in this area and adds slightly to previous records (Fig. 61), but it does not appear to be common.

Cylindroiulus pyrenaicus Brölemann, 1897

Figs. 62-‍74

Material studied:

(52) Alava: Sierra de Peña Gorbea [Gorbeia], Valley of the River “Bayas Ibaia”, north of Saria, at (bridge) Arkarai, 43.017, -2.830, 720 m, 24.04.2009, semi-open stream valley with old Fagus trees and bushes, with pasture beside river, Erica, Ulex, Ilex, Prunus in leaf litter/humus. 8 ♂♂, 14 ♀/J, Steve Gregory leg.

(42) Alava: Sierra de Urbasa, west side, at road A-2128 to Salvatierra/Agurain, south of Puerto de Opacua, south of next road junction at spring of Arroyo/Larrondoa, east of Alto de los Tiles, at site “Orguchi”, 42.7957, -2.3194, 940 m, 23.04.2009, Fagus wood in gorge, fairly open with Calluna Salisb., Anemone, Juniperus, Erythronium, Lathraea, Narcissus bulbocodium L., Helianthemum Mill., Pteridium Gled. Ex Scop. & grasses in leaf litter. 1♂, 14♀♀/J, Desmond Kime leg.

(21b) Navarra: Sierra de Aralar, south Baraibar, on road NA-7510 to Santuario de San Miguel, at area “Urkia”, 42.9762, -1.9318, 710 m, 22.04.2009, wet wood pasture of Fagus, Anemone nemorosa, Arum, Geranium, Helleborus, Euphorbia, Mercurialis, Ranunculus ficaria under stones and logs. 1♂, 3♀♀, Helen Read leg.

(5b) Navarra: Leitza, Ariz Mendiak, between area “Ustarleku” and “Karobieta”, Gorriztaran/Ibaia-stream valley, 43.0789, -1.8839, 550 m, 20.04.2009, slope to road verge, Corylus et al. 1♀, Karin Voigtländer leg. (SMNG VNR 014753-‍9).

(12) Navarra: Sierra de Aralar, south of Errazkin, north of area “Axkarateko Malkorra”, 42.9972, -1.9703, 630 m, 21.04.2009, Fagus woodland. 1♂, 10♀♀/J, 1j♂, Per Djursvoll & J. Paul Richards leg.

(50B) Alava: Montes de Altzania, Sierra de Urquilla, south of mountain chain Aikorriko Mendikatea, southwest of Mountain Aitzgorri – north of end of road from Salvatierra to Ordonana, Zalduondo, Zumarraundi – 1 km south of “Zumarraundi”, “Urbieta”, 42.9175, -2.3305, 830 m, 24.04.2009, eastern slope of open Quercus forest with some Fagus trees in leaf litter. 1♀. Per Djursvoll leg.

(27a) Navarra: Between Sierra de Urbasa and Sierra de Andía, road (NA-120) from Lizaraga to Puerto de Lizaraga, north of cliff, 42.8631, -2.0114, 930-‍960 m, 22.04.2009, steep slope below cliff, stony meadow. 1♀, 1J♂, J. Paul Richards leg.

Total material studied: males: 11; juvenile males: 2; females/juveniles: 49.

There are three samples where no males were found but the pale head, along with the overall size and telson/subanal scale shape indicated this was the correct species.

Diagnosis

Small to medium sized species, with strong and down pointing telson projection and projecting anal valve, although not as pincer-like as in C. elosegiorum sp. nov. Generally well pigmented but with obvious pale head, first few rings and anal valves.

Description

Length: ♂ 16-‍18 mm; ♀ up to 24 mm. Body height: ♂ 1.2-1.5 mm; ♀ up to 2 mm. Body rings: ♂ 36-‍40 podous rings; ♀ up to 42. Body colour: Generally glaucous and dull, dorsal generally darker than ventral. Head and first 1, 2 or 3 rings usually noticeably paler (Fig. 62), becoming yellow in preserved specimens. Despite the overall pale colour there is often a darker band covering and linking the eyes (Fig. 63). Telson, mostly dark, with pale posterior margin and dorsal projection gradually fading in colour to straw/yellow (Figs. 62, 65). Colour in the juveniles sometimes appearing less contrasting. Overall body shape: Anterior constriction behind head but no posterior attenuation. Head: Antennae appearing relatively long and slender. Eye rows generally clear but final row often with reduced number of ommatidia and rather jumbled (Fig. 63). Stadia: ♂ 5RO = VI to 8RO = IX; ♀ up to 9RO = X. Body rings: Metazonites rugged and almost fluted, with close and strong striae sometimes giving a banded appearance, at least anteriorly (Figs. 64). Limbus not crenulate under light microscope. Legs: Legs with accessory claw surpassing claw (Fig. 66). Telson: Preanal ring with long and strong projection, strongly bent ventrally towards the tip. One pair of setae on the tip of the projection, another pair ventrally (Fig. 65). In smaller juveniles of stadium VI the telson projection appears relatively shorter and the down turn less pronounced (Fig. 67). Subanal scaleprojectingbeyond theanal valves a relatively long distance but slightly ventrally, so less ‘pincer-like’ in appearance than C. elosegiorum sp. nov., with 1 pair of setae towards the tip. In smaller juveniles the projection is still present but relatively less well developed. Anal valves with 3-‍4 pairs of setae (Fig. 65).

Mature male secondary sexual characters: Mandibular stipites of mature males distinctly expanded and roughly square shaped. First pair of legs in mature male: small tight hooks. Keel of body ring 7 in mature male only very slightly enlarged.

Gonopods (Figs. 68-‍72): Promerite very rounded with rather weak attachment to the gonopod block. Flagellum of normal length. Mesomerite short and squat, slightly shorter than C. sagittarius, rounded in posterior view. Opisthomerite chunky but posterior margin very membranous and therefore hard to see when glanced at quickly, so the opisthomerite appears at first sight more slender and simple in outline, thickened distally. (Compare Fig. 70 where the membranous area appears faint with Figs. 68-‍69 where it is drawn in). Membranous area on posterior margin of opisthomerite runs adjacent to the paracoxal process so they appear part of the same structure. Paracoxal process more obvious in posterior view and triangular in shape. Female first leg pair thickened and second also perhaps slightly enlarged.

Relationships with other species

One of a relatively small number of Cylindroiulus species with strongly projecting sub anal scale as well as a strongly down curved and pointed telson. Others with this feature are C. apenninorum, C. elosegiorum sp. nov. (see above), C. gestri and C. propinquus all of which have different shaped gonopods and, as mentioned above under C. elosegiorum sp. nov., this does not appear to have any evolutionary implications.

Ecology

The species was found in the leaf litter but, while all locations had trees present, many were described as being more open and/or a wood pasture and almost all were associated with a stream or, in one case, wet wood pasture.

Life history

The stadial spectrum (Fig. 73) indicates that males are semelparous, being mature just in two stadia (VIII and IX). The pattern for the females indicates a similar life cycle (Fig. 74).

Distribution

Kime & Enghoff (‍2017) shows the range for this species along the whole of the Pyrenees and extending a little away from the mountains in southeast France and west in Spain along the coast. The current collection falls within the known range and added nothing new to the atlas maps. Recent experience suggests that it is fairly widespread, from Pau area in France, across the Basque country at least as far as Gorbeia in Alava although it does not appear to be particularly common. C. pyrenaicus has recently been found in the UK where it is likely to have been introduced, and also in several sites in northwest France (‍Gregory et al., 2018).

The current collection of C. pyrenaicus was compared carefully with the original description made by Brölemann (‍1897) and the specimens from the current collection accord well with the original description. The size, general appearance and shape of the telson and subanal scale look very similar. It was noticeable that live specimens have an obvious pale/yellow head which is not mentioned by Brölemann loc. cit. This perhaps becomes less obvious in alcohol but the combination of head colour contrasting with the body and the telson/anal valve shape makes this species quite characteristic. The gonopods at first sight seem a little different to those illustrated by Brölemann (‍1897) but when drawn they are similar. The difference seems to be due to the very membranous appearance of most of the opisthomerite which when drawn give the impression of very chunky posterior gonopods.

Cylindroiulus sagittarius Brölemann, 1897

Figs. 75–83

Material studied

(3) Navarra: Leitza, Ariz Mendiak, between area “Ustarleku” and “Karobieta”, above side stream to Gorriztaran, 43.0778, -1.8775, 615 m, 20.04.2009, grove of Castanea, pollard trees on the slope, Ranunculus ficaria, Daphne, Helleborus, Salvia, Rubus, Lathrea – soil loamy and calcareous. 19♂♂, 3j♂, 25♀♀, Karin Voigtländer (8♂♂, 3♀♀, 5J SMNG VNR 014824-‍1 & SMNG VNR 014744-‍12), Steve Gregory, Helen Read (Hans Reip (7♂♂, 2♀♀, 7j, Reip-E516-F2446-AS175) leg.

(5) Navarra: Leitza, Ariz Mendiak, between area “Ustarleku” and “Karobieta”, Gorriztaran/Ibaia-stream valley, 43.0789, -1.8839, 550 m, 20.04.2009, Alnus wood along stream in the inundation area, in the topsoil layer or under leaves. 2♂♂, 14♂♂/J (Reip-E522-F2460, Reip-E523-F2466 & Reip-E523-F2461), Hans Reip leg.

(8) Navarra: Leitza, Ariz Mendiak, road to area “Kornieta”, between area “Ustarleku” and “Karobieta”, above spring Gorriztaran/Ibaia-stream, 43.0794, -1.8699, 750 m, 20.04.2009, Fagus pollards, calcareous soils under logs and & leaf litter. 1♂, 1j♂, 4♀♀, Steve Gregory leg.

(12) Navarra: Sierra de Aralar, south of Errazkin, north of area “Axkarateko Malkorra”, 42.9972, -1.9703, 630 m, 21.04.2009, young Fagus and old big Quercus, Ilex, Ruscus L., Hedera helix L., Astragalus L.? Euphorbia L.?, ferns in leaf litter. 3♂♂, 4+♀♀/J, Karin Voigtländer (2♂♂, 4♀♀ SMNG VNR 014758-‍14), Per Djursvoll & J. Paul Richards leg.

(20) Navarra: Sierra de Aralar, south Baraibar, on road NA-7510 to Santuario de San Miguel, 42.9851, -1.9203, 670 m, 22.04.2009, open pasture with some Fraxinus and Quercus, stony, calcareous under stones, in Corylus litter. 1♂, 1♀, Per Djursvoll leg.

(22) Navarra: Sierra de Aralar, south Baraibar, on road NA-7510 to Santuario de San Miguel, at area “Izáin”, 42.9714, -1.9384, 790 m, 22.04.2009, Karst area with deep grykes and bare limestone rocks, Fagus woodland & some Crataegus bushes in leaf litter, partly sieved out. 1♀, Helen Read leg.

Guipuzkoa: Aiako Harria P.N., Oiartzun, Oieleku, 43.248, -1.83, 600 m, 09.2005, pollarded Fagus trees, under & in Fagus logs. 8♂♂, 8j♂♂, 16♀♀, Helen Read leg.

Gipuzkoa: Errenteria, (Aiako Harria P.N.) Goikoarkaitzeta (Añarbe), 43.29, -1.89, 350 m, 6.04.2007. 1♂, S. Pagola-Carte leg.

Guipuzkoa: Aiako Harria P.N., Oiartzun, Oieleku, 43.248, -1.83, 600 m, 6-‍15.05.2007, Fagus forest, Kaila trap on Fomes. 1♂, S. Pagola-Carte leg. (NHMD).

Navarra: Leitza, Ariz Mendiak, at area “Kornieta”, 43.0698, -1.8823, 910 m, 4.2.2010, Fagus woodland, poor understorey, rocky, siliceus soils, mosses, Helleborus under bark & in leaf litter. 1♀, Helen Read & Mark Frater leg.

Material collected in 2009 and identified by Hans Reip:

(3) Navarra: Leitza, Ariz Mendiak, between area “Ustarleku” and “Karobieta”, above side stream to Gorriztaran, 43.0778, -1.8775, 615 m, 20.04.2009, grove of Castanea, pollard trees on the slope – Ranunculus ficaria, Daphne, Helleborus, Salvia, Rubus, Lathrea – soil loamy and calcareous. 7♂♂, 2♀♀, 7J, Hans Reip leg.

(5) Navarra: Leitza, Ariz Mendiak, between area “Ustarleku” and “Karobieta”, Gorriztaran/Ibaia-stream valley, 43.0789, -1.8839, 550 m, 20.04.2009, Alnus wood along stream under mosses on dead wood. 2♂♂, 5♀♀, 9J, Hans Reip leg.

Total material studied: males: 45; juvenile males: 14; females/juveniles: more than 94.

Diagnosis

A medium sized species, chestnut brown, usually darker dorsally and often with paler head, but with dark band across the ommatidia patches. Large number of body rings. Strong, prominent and straight telson projection. Opisthomerite of gonopods unique in shape and very elongated with the flagellum projecting from the end.

Description

Length: ♂ 18-‍22 mm; ♀ up to 19 mm. Body height: ♂ 1.0-1.4 mm; ♀ up to 1.9 mm. Body rings: ♂ 44-‍54 podous rings; ♀ up to 57. Notable for the large number of body rings and the large number of apodous rings, even in mature individuals. Mature males had between 2 and 6 apodous rings. Body colour: Mid/chestnut brown, generally paler ventrally, with darker mid dorsal pattern (Fig. 75). Head often paler but with darker band extending across the ocular patches and the ‘forehead’. Sometimes collum and anal valves paler too. Overall body shape: Anterior constriction not obvious in dorsal view, although strangely narrowed ventro-dorsally. Body more or less parallel sided but apodous rings (usually high in number) reduced in height relative to rest of body. Head: Antennae generally pale with darker marks. Eye rows mostly clearly readable, ommatidia barely reduced in number. Stadia: ♂ VIII or IX; ♀ up to XI. Body rings: Metazonites with striae strong and regularly placed, quite wide apart. Limbus not appearing crenulate. Legs: Generally paler in colour. Accessory claw extending beyond claw to roughly 25% of the claw length. Telson: Preanal ring with very strong, level and pointed projection (Fig. 76). One pair of setae at end of projection and one pair ventrally. Subanal scale not enlarged and bearing 1 pair of setae. Anal valves with 3 pairs of setae.

Mature male secondary sexual characters: Mandibular stipites of mature males smoothly expanded and curved ventrally, not extensively expanded (Fig. 77). First pair of legs in mature males small hooks. Keel of body ring 7 in mature males opening very wide.

Gonopods (Figs. 78-‍81): Promerite: Medium length and fairly wide; no ‘window’. Mesomerite: Considerably shorter than promerite, apex rounded. Opisthomerite: Very long and thin, expanded at proximal part to a small but distinct brachite and a membranous area between brachite tip and solenomerite. Smaller phylacum. Flagellum usually seen poking out of the tip of the opisthomerite. No obvious paracoxal process, just a slight broadening at posterior and base of opisthomerite.

Relationships with other species

Gonopod shape is in this species is relatively unusual and it does not easily fall into a group with any other species. It has some resemblance to C. punctatus (in habitat as well as appearance) although lacks the long paracoxal process of that species.

Associated fungi

The species is notable for the fungus Rickia laboulbenioides found on the legs of some individuals (identified by Henrik Enghoff). In the current collection fungi were seen in 7 out of 21 mature males examined but only on two mature females. The fungus was found on the anterior legs, extending just beyond the gonopods. Some individuals appeared to have just a small quantity of fungi, but others had high densities making the legs appear ‘furry’.

Ecology

This species always seems to be found in and around decaying wood, fulfilling a role that in the UK is occupied by Cylindroiulus punctatus, although it perhaps has a preference for wood in the later stages of decay. Many of the sites where it has been found are wood pasture with old trees, especially beech (Fagus sylvatica L.). Cylindroiulus punctatus is also found in the Pyrenees/Basque area but seems less common.

Life history

Collections were made in two different times of the year (April and September). Mature males were only found in stadia VIII and IX and the breakdown in number was almost identical for each season (9 in stadia VIII and 3 or 4 in stadia IX) so the two collections were combined to produce the stadial spectrum which is shown in Figs. 82-‍83. This shows that the males are clearly semelparous, in the females however, while there is a peak in stadia VIII and IX, a small number were found in stadia X and XI hinting that either some females mature later or that a small number may moult following reproduction and therefore may have the capacity to be iteroparous.

Distribution

Cylindroiulus sagittarius was confined to the Pyrenees and north-west Spain (‍Kime & Enghoff, 2017) and the current collection does not extend the range in this area. It has recently been found in the UK (‍Gregory & Owen, 2019) where presumably it has been introduced.

Cylindroiulus cf. sagittarius

The following collections did not include mature males, but as this species has been recorded from the same location on different dates, and they appear identical, it is highly likely they are this species.

Guipuzkoa: Aiako Harria P.N., Oiartzun, Oieleku, 43.248, -1.83, 600 m, 13.07.2007, Fagus forest, under bark of Fagus log. 2j♂, 4♀♀, S. Pagola-Carte leg. No mature males but look very much like C. sagittarius.

Guipuzkoa: Aiako Harria P.N., Oiartzun, Oieleku, 43.248,-1.83, 600 m, 15.06.2007, Fagus forest, under bark of Fagus log. 1♀, S. Pagola-Carte leg. No mature males but look very much like C. sagittarius.

Cylindroiulus sanctimichaelis Attems, 1927

Figs. 84-‍88, 91, 96-97

Material studied:

(1) Navarra: Pamplona, Airport, 42.7669 -1.6431, 430m, 19.04.2009. 1♂, 4♀♀, Karin Voigtländer (1♂, 3♀♀, 1j, SMNG VNR 014741-‍4) & Hans Reip (Reip-E511-F2426-AS176) leg.

(2) Navarra: Pamplona, south of Airport, near Salinas de Pamplona, at Río Elortz, 42.7539, -1.6431, 440 m, 19.04.2009, edge of deep stream valley, meadow slope, dry area. 3♂♂, 5♀♀ (Reip-E512-F2430). Hans Reip leg.

(8) Navarra: Leitza, Ariz Mendiak, road to area “Kornieta”, between area “Ustarleku” and “Karobieta”, above spring Gorriztaran/Ibaia-stream, 43.0794, -1.8699, 750 m, 20.04.2009, Fagus pollards, calcareous soils under logs and in leaf litter. 1♀, Steve Gregory leg.

(15) Guipuzkoa: Sierra de Aralar, Tolosa, 500 m south of Bedaio/Goikoa, 43.0494, -2.04, 420 m, 21.04.2009, farm buildings & stone wall, logs, wood, stones and rubble under stones and logs. 3♂♂, 2♀♀, Helen Read leg.

(16) Guipuzkoa: Sierra de Aralar, 10 km SSE Beasain: Kaxeta (at junction of road from Lazkao to Etxarri-Aranaz [GI-120] and road to Aia [GI-4153], Arrandoa, Kaxeta (stream “Arroyo Marumendi”), 42.9581, -2.1564, 310 m, 21.04.2009, Stream bank. 2♂♂, 8♀♀, Steve Gregory, Hans Reip (Reip-E533-F2497-AS177) & Norman Lindner leg.

(18) Guipuzkoa: Sierra de Aralar, Beasain, road from Lazkao to Etxarri-Aranaz, Pass Puerto de Lizarrusti, 42.9614, -2.0983, 690 m, 21.04.2009, forest of Fagus, on slope under stones & in leaf litter. 1♂, 1♀, Per Djursvoll leg.

(20) Navarra: Sierra de Aralar, south Baraibar, on road NA-7510 to Santuario de San Miguel, 42.9851, -1.9203, 670 m, 22.04.2009, open pasture with some Fraxinus and Quercus, stony, calcareous under stones, in Corylus litter. 12 ♂♂, 22♀♀/J, Per Djursvoll, Helen Read (1♂), Karin Voigtländer (2♂♂, 3♀♀, SMNG VNR 014765-‍19) & Hans Reip (5♂♂, 1♀ Reip-E538-F2519) leg.

(24) Navarra: Sierra de Aralar, Sierra de San Miguel, mountain point Artxueta at radio mast, 42.9525, -1.9668, 1300 m, 22.04.2009, Karst area with bare limestone rocks, pasture under stones and dung & in leaf litter. 8♂♂, many females and juveniles but badly preserved so dried and broken and precise number difficult to determine. Helen Read, Per Djursvoll, Karin Voigtländer (1♂, 3♀♀, 1J, SMNG VNR 014817-‍1) & Hans Reip (4♂♂, 3♀♀, 11j, Reip-E544-F2543) leg.

(26) Navarra: Sierra de Aralar, Sierra de San Miguel, southern lower slope at Iraneta, 42.9258, -1.9597, 500 m, 22.04.2009, dry Quercus wood in leaf litter. 1♂ (Reip-E546-F2554), Hans Reip leg.

(27) Navarra: Between Sierra de Urbasa and Sierra de Andía, road (NA-120) from Lizaraga to Puerto de Lizaraga, north of cliff, 42.8631, -2.0114, 930-‍960 m, 22.04.2009, steep slope below cliff, stony meadow. 1♂, 1♀, J. Paul Richards leg.

(28) Navarra: Between Sierra de Urbasa and Sierra de Andía, road (NA-120), Puerto de Lizarraga (direction south), 42.8589, -2.0122, 1030 m, 22.04.2009, meadow with big stones & some bushes (Sambucus). 2♂♂, 6♀♀, Steve Gregory, J. Paul Richards, Hans Reip (1♀, 2j Reip-E548-F2560) & Norman Lindner leg.

(30) Navarra: Sierra de Andía, at road NA-7330 (from Lezáun to Arizaleta), at area “Los Ristros”, ca. 500 m WNW Arizaleta, 42.7589, -1.9833, 700 m, 22.04.2009, Mediterranean Quercus forest (evergreen), used as wood pasture, incl. some small areas with dry meadows (with small thorny bushes, some bigger stones, some small Buxus bushes, Quercus ilex, Genista, shrubby Thymus, shrubby Lonicera, Sedum, grasses) on the meadow under stones. 1♂, 8♀♀, Steve Gregory leg.

(31) Navarra: Sierra de Andía, Valle de Goñi, 500m NE Goñi at area “Mustillogorri”, 42.855, -1.8992, 930 m, 22.04.2009, grazed deciduous Quercus woodland, within agricultural areas (pastures) under stones. 4♂♂, 7♀♀, 2j♂, J. Paul Richards, Steve Gregory & Desmond Kime leg. (GEE).

(35) Navarra: Sierra de Urbasa, on top at southern cliff border, east of road NA-7182, 42.7989, -2.1417, 930 m, 23.04.2009, pasture on stony ground, some thorny bushes, occasional trees or groups of Fagus under stones. 1♂, 1♀, Per Djursvoll & Karin Voigtländer (1♀, SMNG VNR 014826-‍5) leg.

(37) Navarra: Sierra de Urbasa, under southern cliff border, Valle de Amescoa Alta, south of road NA-7130, between Larraona and Aranarache, at site “Karrerabide”, 42.7722, -2.2408, 770 m, 23.04.2009, small wood of Quercus, grazed by livestock, pronounced shrub layer: Sambucus nigra L., Viburnum L., Crataegus, Rubus, Rosa L., mainly without herb layer, neighbouring: pasture and arable field in the soil, upper layer. 1♂, 10♀/J, Hans Reip leg.

(39) Navarra: Sierra de Urbasa, under southern cliff border, Valle de Amescoa Alta, south of road NA-7130, south of Larraona, 42.7697 -2.2605, 780m, 23.04.2009, wood pasture with Quercus and scrub and grassy trackside verges under stones. 4♂♂, 4♀♀, 1j♂ (1♂, 1♀, MNCN 20.07/2093-2094) Helen Read, J. Paul Richards & Steve Gregory leg.

(44) Guipuzkoa: Montes de Altzania, Sierra de Urquilla, south of mountain chain Aikorriko Mendikatea, southwest of Mountain Aitzgorri – north of end of road from Salvatierra road to Ordonana, Zalduondo, Zumarraundi – high plain Alizania, north of Portua Zarra, 42.9354, -2.3298, 1160 m, 24.04.2009, karst plain with small growing Fagus trees and open grasslands under stones, in accumulations of leaves between the stones. 4♂♂, 15♀♀/J, Karin Voigtländer (1♂, 2♀♀ SMNG VNR 014828-‍7 & SMNG VNR 014786-‍6) & Hans Reip (3♂♂, 8♀♀, 5j, Reip-E559-F2600), leg.

(54) Navarra: Pamplona, south, Sierra de Izco, Sierra de Alaiz, north of Altos de la Cruz, south of village Guerendiain, 42.7047, -1.6011, 640 m, 25.04.2009, Northern slope, closed area of Buxus bushes with some Quercus trees, Crataegus, Hedera helix, Trifolium L, no humus layer in leaf litter. 1♂, 1♀, Karin Voigtländer (1♀, SMNG VNR 014822-‍5) & Hans Reip (1♂, SMNG VNR 014822-‍5) leg.

(55) Alava: south Salvatierra, Campezo, near village Oteo, 42.726, -2.3628, 710 m, 25.04.2009, Quercus faginea wood; Q. ilex, Arbutus unedo, Viburnum lantana, Juniperus under stones and leaf litter. 1♂, 2♀♀, Helen Read leg.

(56) Alava: south Salvatierra, Campezo, near village Oteo, Parque Santa Lucia, 42.701, -2.319, 610 m, 25.04.2009, Quercus ilex forest with Buxus sempervirens, in leaf litter. 6♂♂, 20+♀♀/J, Helen Read, Desmond Kime (GEE) & Per Djursvoll leg.

(59) Alava: Sierra Chiquita, northern side, Santa Cruz de Campezo, south of Ermita de Ivernalo, 42.6655, -2.3329, 680 m, 26.04.2009, evergreen forest of Quercus, with Buxus and Erica in leaf litter. 2J (Reip-E570-F2647), Hans Reip leg.

Bizkaya: Leioa, Jauregia Jardines del Palacio Arataza, 18.04.2015. 2♂♂, V. Martínez -Pillado leg. (NHMD).

Bizkaya: Leioa, Peruri, Auzoa, 31.10.2015. 2♂♂, 2♀♀, V. Martínez -Pillado leg. (NHMD).

Bizkaya: Leioa, road to Porto Zahorro (Algorta), 17.04.2015. 1♂, 1♀, V. Martínez -Pillado leg. (NHMD).

Bizkaya: Leioa, 30.10.2015, captured at night. 1♂, 3♀♀, V. Martínez-Pillado leg. (NHMD).

Teruel: Cretas, Túmulos. 40.952, 0.224. 4.04.2015. 1♂, 4♀♀, J.D. Gilgado & V. Martínez leg. (NHMD).

Gipuzkoa: Urkillaga a Lareo Ataun, 42.97, -2.14, 650 m, 16.03.2004. 2♂♂, I. Zabalegui leg.

Total material studied: males: 66; juvenile males: 3; females/juveniles: more than 129.

Diagnosis

A chunky, dark brown species, similar to C. caeruleocinctus but with differences in the shape of the opisthomerite of the gonopods. Larger specimens have a distinct ‘golden glow’.

Description

Length: ♂ 21-‍23.5 mm; ♀ up to 39 mm. Body height: ♂ 1.4-1.9 mm; ♀ up to 2.9 mm. Rings: ♂ 39-‍44 podous rings +2 apodous; ♀ up to 50+1 rings. Body colour: Chestnut brown metazonites, alternating with darker brown prozonites, slightly paler ventrally, some markedly so. Many specimens with a gold sheen to the metazonites, not as strongly metallic as C. caeruleocinctus but larger specimens with a ‘golden glow’ Fig. 84, collum generally dark in preserved specimens. Legs yellow brown. Overall body shape: Anterior constriction present, with distinct narrower ‘neck’. Posterior attenuation quite strong, posterior quarter to fifth of body length noticeably narrower. Head: Antennae brown. Eye rows apparently clear and easy to ‘read’ but, in reality rows rather muddled so difficult to be sure of exact number in most mature specimens. Stadia: ♂ probably mostly 9 rows = X but up to 11 rows = XII; ♀ up to 11 or 12 rows = XII-XIII. Body rings: Metazonites with striae fairly strong, even and relatively widely spaced. Limbus not crenulate. Legs: Brown, accessory claw slightly longer than claw. Telson: No projection beyond anal valves. One pair of setae dorsally and one pair ventrally (Fig. 85). Subanal scale normal and not projecting. With one pair of setae. Anal valves usually with three pairs of setae but ranges from 1 to 5 pairs.

Mature male secondary sexual characters: Mandibular stipites of mature males relatively strongly projecting both anteriorly and ventrally. First pair of legs in mature male forming a small tight hook. Body ring 7 in mature males expanded ventrally to a distinct keel with anterior gonopods of some males projecting through the gap in ring 7.

Gonopods (Figs. 96-‍97): Promerite: Long and slender. Mesomerite: Slender, shorter than promerite, blunt tipped. Anterior gonopods appearing identical to caeruleocinctus. Opisthomerite large, angular brachite, slightly convex in outline (C. caeruleocinctus is generally slightly concave). Opisthomerite more prominent and irregular in comparison to C. caeruleocinctus. There is an obvious division between the solenomerite and the phyllacum (as in C. caeruleocinctus and different to C. londinenis (‍Blower 1985)). High paracoxal rim. No paracoxal process, although perhaps a small ‘nick’ in the posterior line of the gonopods basally.

Ecology

Vicente (‍1981) noted that this species was very abundant in Quercus ilex forest, amongst the leaves. For Serra et al. (‍1997) it was the second most abundant Julid species in a sclerophyllous forest consisting of Q. ilex with an understorey of Arbutus unedo. Serra et al. (loc. cit.) recorded a mean annual density of 35.22 individuals per meter squared for this species with peaks in later summer and autumn. He also recorded a preference for the intermediate and deeper soil horizons and that it was notable for the high proportion of females relative to males. Sampling by Serra et al. (‍1997) was carried out by taking soil cores and extraction using Berlese-Tüllgren devices. In the present collection it was found in the leaf litter of various woodlands, but the most extensive collection was made in a Quercus ilex forest where it was very abundant.

Life history

The current collection had a good number of specimens, so opportunity was taken to count eye rows and podous segments in order to attempt to shed a light on the life history of this species. Unfortunately, the eye patches were composed of a large number of ommatidia and, while appearing from a distance to be regular and easy to count, they were actually more difficult to determine than expected, especially for older individuals. The graphs include data for some individuals in which a ‘best guess’ has been made.

The stadial spectrum for the males (Fig. 85) shows a relatively tight group of mature stadia and that of the number of podous rings (Fig. 86) seems to substantiate this. The females show a wider range of both stadia (Fig. 87) and podous rings (Fig. 88), not only through the inclusion of juveniles but also to the right of the graphs. That for the stadia is bimodal and this indicates that while it seems likely that the majority of males reach maturity and die following reproduction, some of the females may well survive to reproduce more than once.

Distribution

This species has a widespread but scattered distribution within Spain (Fig. 91), from the foothills of the Pyrenees to Andalucía and down the eastern side of the Iberian Peninsula. There seem more records in the north, but this may be the result of greater collecting effort in these regions. Despite it being found frequently in the current collections the known range has barely been increased.

Cylindroiulus cf. sanctimichaelis

Material studied:

Castilla-La Mancha: Guadalajara, Embalse de Tajera (cerca de Cifuentes). 14.3.2015. 2♂♂, J.D. Gilgado leg. (NHMD).

Zaragoza: Luesia, Casa Forestal, 2.42378940825893, -1.0053219496876111, 14.5.2015, 2♂♂, 7♀♀, J.D. Gilgado leg. (NHMD).

The specimens from these collections were very similar to each other and clearly fall in the C. londinensis group (see below), probably closest to C. sanctimichaelis. In terms of size and ring number they fit within C. sanctimichaelis range (Figs. 16-‍17), although some were towards the low end of the height range. The eye rows appeared rather more regular, and the males appeared to have slightly longer legs in relation to the body size (Fig. 89) although this was clearly an illusion. Body height divided by total mid-body leg length (using the method described in Enghoff (‍1982) gave figures of 1.38 and 1.35 for males from Zaragoza and Guadalajara respectively and 1.33 and 1.40 for males from Parque Santa Lucía and Oteo respectively. The brachite, solenomerite and phylacum of the gonopods are all in a line (Fig. 90) and the brachite and phylacum are both large, thus resembling sanctimichaelis. However, the phylacum is more pointed distally and more rounded in overall shape, with less distinction between it and the solenomerite. The margin of the brachite also has a flatter profile. It is not easy to determine if these specimens are sufficiently consistently different to consider them a separate species or if they are individuals within an overall variable species. Genetic studies would be useful to helpelucidate.

Relationships between C. sanctimichaelis and similar species

Cylindroiulus sanctimichaelis is one of a group of seven very similar species all found in the Iberian peninsular: C. sanctimichaelis, C. caeruleocinctus, C. caramelos sp. nov., C. chalandei, C. finitimus, C. ibericus, 1913 and C. londinensis. All have a relatively large body size for Cylindroiulus and similar gonopods with a large and obvious brachite, a phyllacum of varying size, a small or absent paracoxal process but a high paracoxal rim.

In the present collection, several people (including myself) initially misidentified C. sanctimichaelis as C. caeruleocinctus and it was only when looking more critically at the gonopods that it became evident that they were closer to C. sanctimichaelis. The specimens from the Basque region appear to be slightly smaller, and generally with slightly fewer podous rings, than previous descriptions. In addition, the number of pairs of setae on the anal valves was more variable and ranged from 1 to 5 (with a mean of 3.2). However, careful examination of the gonopods revealed them to be most like C. sanctimichaelis.

Cylindroiulus sanctimichaelis was described by Attems (‍1927) who distinguished it from the similar C. ibericus, a species described previously by Brölemann (‍1913). Several previous publications have also compared groups of 2-‍3 species such as Mauriès (‍1964) for C. finitimus, C. londinensis and C. caeruleocinctus and Vicente (‍1981) for C. chlandei, C. finitimus and C. sanctimichaelis. Older literature can be misleading because of the confusion with C. teutonicus (now regarded as C. caeruleocinctus) and that of C. londinensis/C. finitimus (C. finitimus was previous considered a subspecies of C. londinensis).

Attems (‍1927) compared C. sanctimichaelis, C. ibericus and C. teutonicus (i.e. C. caeruleocinctus) and considered them to be very similar. He noted that C. sanctimichaelis had a clear notch in the gonopod between the brachite and the solenomerite. In his key he separates C. caeruleocinctus from the other two on the basis of the presence of the teeth on the apex of the solenomerite and then C. sanctimichaelis as having 3 pairs of setae on anal valves (C. ibericus having 5-‍6 pairs), and shape of the posterior gonopods. However, in the current collection the number of setae on the anal valves of what is here considered to be C. sanctimichaelis is variable and there are often more than 3 pairs, however the gonopods do look more like those figured by Attems as C. sanctimichaelis. While the number of pairs of setae on the anal valves may be indicative of particular species it may not be a reliable character and sometimes the setae are worn off and the point of insertion difficult to see.

Mauriès (‍1964) looked in detail at C. caeruleocinctus, C. londinensis and C. finitimus and decided they were all good species, despite Ribaut (‍1905) originally describing C. finitimus as a variety of C. londinensis. Mauriès (loc. cit.) separated them by the telson shape (in C. londinensis) and the relative length of brachite to the solenomerite (and phylacum). He also noted the clear division or notch between the brachite and the solenomerite in C. caeruleocinctus and C. finitimus. A character also used by Blower (‍1985) to separate C. caerueocinctus from C. londinensis.

Mauriès (‍1964) concluded that C. finitimus is a distinct species with constant characters, a geographic division (higher altitudes and in the valleys of the central and eastern Pyrenees) and a well-defined habitat.

Vicente (‍1981) compared C. chalandei, C. finitimus and C. sanctimichaelis and was clearly able to distinguish between them, with C. sanctimichaelis seeming to have been quite common and C. finitimus rather rarer in her region. Vicente’s (‍1981) figures of gonopods show the brachite of C. sanctimichaelis much longer than the solenomerite and phyllacum, (like Mauriès’s drawings of C. londinensis) and a phyllacum more pronounced, both in size and definition from the solenomerite, than in C. chalandei and C. finitimus. The brachite was also flat topped and not pointed in C. finitimus, a feature shown in some of the illustrations in Mauriès (‍1964). Vicente (‍1981) also lists C. finitimus as being slightly larger than C. sanctimichaelis but with fewer body rings. Vicente & Ascaso (‍1990) subsequently made a detailed study of the ecology of C. ibericus suggesting that there were no difficulties distinguishing it as aspecies.

Table 1 shows the characteristics of these similar species with details taken from the literature, both the original descriptions and previously published comparisons. Some information was added from personal observation where specimens were available, for example the shape of the tip of the mesomerite in posterior view, something indicated in the literature as being potentially of help in distinguishing some species. The relative lengths of the three main structures of the opisthomerite (brachite, solenomerite and phylacum) have been used previously by authors to distinguish the species and this does seem helpful for some of them. See also Figs. 92-‍104 which illustrate the gonopods of the various species. Specimens of C. finitimus in NHMD were checked by Henrik Enghoff and a comparison made with photographs from the current collection. They were found to clearly differ and had a much smaller phylacum. No specimens of C. chalandei or C. ibericus were available for study.

In summary:

Other characters that might be worth considering in future include the colour. Larger preserved specimens of C. sanctimichaelis observed here appeared to have a ‘golden glow’ which is rather different to the metallic copper colour seen in C. caeruleocinctus, but the literature suggests that other species have a tendency to be golden too (such as C. finitimus and C. chalandei). C. caramelos sp. nov. (see above) is apparently black with white legs. The ventral margin of the brachite could also be useful as it can be more-or-less flat, or with a convex or concave curve.

The distributions of the species may also help. Kime & Enghoff (‍2017) show that C. ibericus has a very narrow distribution in Huesca. Cylindroiulus finitimus seems also only to have been found at high altitude in the Pyrenees and Mauriès (‍1964) referred to it as an altitudinal variant of C. londinensis. The locations of both of these species fall within the range of C. sanctimichaelis which is considerably more extensive. Cylindroiulus londinensis and C. caeruleocinctus are much more widely distributed across Europe but it seems likely that they may have originated in Iberia. Cylindroiulus caramelos sp. nov. has only been found from area of mountains near Tarragona connected to the Sistema Iberia but not part of the Pyrenees, however also within the range of C. sanctimichaelis, but it may also be common further south around the Ebro delta (H. Reip pers. comm.)

It is possible that some of these species, particularly C. caeruleocinctus and C. sanctimichaelis, form a complex in Iberia but that one morph has become more widespread in northern Europe and remained relatively consistent in form. This situation is perhaps similar to that of Ommatoiulus moreletii (Lucas, 1860) which seems to be variable in Spain and Portugal but those introduced to Australia are less so (‍Baker, 1984). Another Ommatoiulus species, O. diplurus (Attems, 1903), also seems to be polymorphic in Spain, with considerable variation in gonopod structure (‍Akkari & Enghoff, 2012).

In conclusion, it seems likely that all the six previous described species, plus that newly described here, are reliable and can be distinguished, but they are challenging to determine. Carefully scrutiny is needed to separate them and there is some variation within some of the apparent species as here understood. It may be that there are fewer, more variable species, with C. caerueocinctus and C. sanctimichaelis (for example) two ends of a spectrum, however they are here regarded as two separate species. The specimens from Zaragoza and Guadalajara seem a little different from the ‘typical’ C. sanctimichaelis as seen in other specimens in this collection, but not different enough to be confident in ascribing them to a new species. This approach was supported by the graphs showing rings vs body height, where they fall within the current variability of C. sanctimichaelis (Figs. 16-‍17).

It is clear that many previous workers have struggled to determine species within this group, and it would benefit from combined genetic and morphological study to explore the species concept within this group and to look at the variation within some of the more widespread species, both within the Iberian Peninsula and across other parts of northern Europe.

In light of the number of Iberian species of millipede that have been found in Britain in recent years (see under C. pyrenaicus and C. sagittarius above) it is potentially possible C. sanctimichaelis, which seems quite a widespread species in Iberia, may also be found, so specimens of C. caeruleocinctus should be scrutinised carefully.

Cylindroiulus unciger Attems, 1952

Figs. 105-112

Material studied:

Aragón: Zaragoza, Monegros, region ‘Retuerta da Pina, near Pina de Ebro, 1.10.1993, stones. 1♂, 1♀, J. Blasco-Zumeta leg. (NHMD).

Madrid: Guadalix de la Sierra beside Embalse de Pedrezuela, 14.04.2015, under stones in a meadow with Quercus ilex ssp. rotundifolia (Lam.) Tab. Morais and Fraxinus sp., calcareous soil. 1♂, 7♀♀, J.D. Gilgado leg. (NHMD).

Total material collected: males (2), females (8)

Diagnosis

A small species, lacking any telson projection or distinguishing external characteristics, but with posterior gonopods unlike any other species in the genus so far described.

Description

Length: ♂ 19-‍21.1 mm; ♀ up to 25 mm. Body height: ♂ 1.2-1.3 mm; ♀ up to 1.8 mm. Body rings: ♂ 49-‍50 podous + 2 apodous, ♀ up to 55 podous rings + 1 apodous. Body colour: Pale brown with darker brown head with usual Cylindroiulus colour pattern on the head, darker collum. Metazonites golden/straw brown and prozonites grey. Darker both dorsally and anteriorly. Photographs of a live animal provided by the collector (Fig. 105) show a pretty banding pattern at least dorsally. Overall body shape: With little variation, anterior constriction slight and no posterior attenuation. Head: Antennae brownish in colour. Eye rows making a full equilateral triangle and rows reasonably ‘readable’. Stadia: ♂ X-XI; ♀ up to XI or XII. Body rings: Striae regular and vaulting negligible. Limbus not crenulate. Legs: golden with brown markings, darker anteriorly. Accessory claw of similar length to claw. Telson: Pre anal ring lacking dorsal projection, not dark in colour but with darker margins. Bearing one pair of setae ventrally and another pair on the tip of the telson (Fig. 106). Subanal scale bearing one pair of setae. Anal valves darker basally, becoming more orange/brown posteriorly. With three pairs of setae.

Mature male secondary sexual characters: Mandibular stipites of mature males expanded ventrally, with almost vertical anterior margin, (Fig. 107). First pair of legs in mature male tight hooks with slight ‘knee’. Keel of body ring 7 in mature male projecting ventrally to a low ridge around a wide ventral opening.

Gonopods (Figs. 108-‍111): Promerite clearly longer than mesomerite, with no perforation or ‘window’. Flagellum of normal length. Mesomerite flat topped, parallel-sided and fitting into a slight ‘cavity’ in the promerite. Large and high paracoxal process. Opisthomerite distally divided into two parts with large gulf between them. Solenomerite expanded along the length by a phylacum. Brachite large and bird-like in shape with a ‘beak’ pointing towards the solenomerite opening. The shape of the opisthomerite is characteristic but much more complex than that illustrated in the original description. No obvious paracoxal process but slight ‘bump’ on posterior margin of opisthomerite.

Relationships with other species

Attems (‍1952) described C. unciger in the subgenus Alpicylindrus, together with C. festai Manfredi, 1937, C. limitaneus (Brölemann, 1905), C. tricuspis Verhoeff, 1932 and C. aostenus Verhoeff, 1932, which are all Italian and have gonopods that look nothing like the those illustrated by him for C. unciger. Attems (‍1952) also described differences between the gonopods of C. unciger and those of C. bouvieri which he considered a similar species. Again, the gonopods in the description of C. bouvieri by Brölemman (‍1896) (= C. parisorum) look nothing like those illustrated by Attems (‍1952) for C. unciger.

The present specimens are in accord with the original description in terms of size and telson shape although the type was black in colour and had 56 body rings. The illustrations of the gonopods in the original description are very simplistic but the overall shape, the relative length of the promerite to mesomerite and the high paracoxal process all accord with the current specimens. The location in the province of Madrid also seems to indicate that they are this species.

Ecology & Life history

There are insufficient specimens from the current collection or in the original description to make any comments about the ecology or life cycle of this species.

Distribution

The type specimen was found in El Pardo, Madrid Province, just north of the city. Guadalix de la Sierra is just a little further north. Kime & Enghoff (‍2017) show an additional location near Zaragoza collected in dry gypsum ground with planted Pinus halepensis Mill., probably not far from Monegros which is just east of Zaragoza, thus the two specimens add slightly to our knowledge of the distribution of this species (Fig. 112). Both referees commented that this species is locally common in many areas in and around Madrid Province.

Cylindroiulus indet.

Two collections of females and juveniles could not be identified to species and could be C. latestriatus or C. britannicus (‍Verhoeff, 1891).

(43) Alava: Salvatierra/Agurain, in the town (urban), 42.8539, -2.3923, 590 m, 23.04.2009. in Castanea sativa Mill. litter. 1♀, 1J, J. Paul Richards leg.

(14) Navarra: Sierra de Aralar, village Erratzin, village square, 43.0075, -1.9603, 400 m, 21.04.2009, open village habitat, meadow (synanthropic) under stones and in dead wood. Steve Gregory leg.

Key to species of Cylindroiulus from Spain and Portugal