in Portugal, with description of six new species

. In this work we present a revision of the genus Ommatoiulus Latzel, 1884 in Portugal. Based on recently collected material and older museum samples, including type specimens, we describe six new species to science, viz . Ommatoiulus alacygni sp. nov., O. camurus sp. nov., O. denticulatus sp. nov., O. litoralis sp. nov., O. staglae sp. nov. and O. stellaris sp. nov. The species O. alacygni sp. nov., O. denticulatus sp. nov. and O. staglae sp. nov. described from the Algarve are outstanding by their extremely reduced mesomerital process. The species O. porathi (Verhoeff, 1893) and O. andalusius (Attems, 1927) are recorded and redescribed for the first time after their original description. The finding of O. andalusius – originally described from Andalusia in Spain – constitutes a new record for Portugal together with two species, viz . O. fuentei (Brolemann, 1920) and O. martensi Mauriès, 1969. The taxonomic status of several species is revised. Thus Archiulus ( Schistocoxitus ) cingulatus Attems, 1927 is here considered as a junior synonym of Ommatoiulus lusitanus (Verhoeff, 1895) while Schizophyllum cervinum Verhoeff, 1910 is synonymized with Ommatoiulus moreleti (Lucas, 1860). An identification key to all hitherto known Portuguese species of Ommatoiulus is presented as well as a distribution map illustrating the various species occurrences in the country.


Introduction
With 76 described species from Europe and North Africa, the genus Ommatoiulus Latzel, 1884 is the most diverse of the tribe Schizophyllini, which also includes the monotypic genus Tachypodoiulus Verhoeff, 1893, andthe 'genus' Rossiulus Attems, 1926 -the latter has never been formally elevated to a full genus though treated as such in recent literature (see Akkari & Enghoff 2012). There are several subgeneric names in Ommatoiulus but these have long been considered as superfluous due to major discrepancies in their definition (see Akkari & Enghoff 2012 and references therein). The species of Ommatoiulus in the Spanish region Andalusia have recently been a subject of a revision by Akkari & Enghoff (2012) who recorded 19 species, ten of which new to science. The high proportion of new species -ca half of the total species known -in southern Spain is in line with the high species diversity in the Iberian Peninsula, one of the main Mediterranean biodiversity hotspots, in combination with the fact that the genus has not been subject to recent revisions in this area.
In the present work, we revise the genus Ommatoiulus in the southwest Iberian sector, Portugal; describe six new species to science and record further three species as new to Portugal. On the other hand, we revise the taxonomic status of several nominal species and propose new synonymies. Furthermore, we present an identification key based on gonopod structures to all hitherto known Ommatoiulus species of the area.

Material and methods
Specimens are preserved in 70% alcohol. Measurements and photographs were made with a Nikon DS-F2.5 camera mounted on a Nikon SMZ25 stereo microscope, using NIS-Elements Microscope Imaging Software with an Extended Depth of Focus (EDF) patch. Scanning electron microscopy (SEM) images are presented to illustrate the gonopods of some species. Samples prepared for SEM were dehydrated using 96% ethanol, followed by acetone, air-dried, mounted on aluminium stubs and coated with platinum. Images were taken using a JEOL JSM-6335F scanning electron microscope. All images were processed with Adobe Photoshop CS6 and assembled in Adobe InDesign CS6. The map was produced using Qgis 2.14. with GPS coordinates based on WGS 84.
A genus of Julidae characterized by: Lack of frontal setae. Lack of a flagellum on the anterior gonopod. Posterior gonopods consisting of a free mesomerite, a solenomerite, a strongly developed coxa with an often elaborate paracoxal process. Solenomerite at base with a large, roundish, open fovea for storage of spermatozoa.

Diagnosis
Most similar to O. denticulatus sp. nov. and O. staglae sp. nov., agreeing with these in having a very reduced mesomerite and a complex multi-branched solenomerite; differing from both in the shape of the solenomerite and the jagged coxa resembling, in posterior view, a swan wing.

Etymology
The species epithet is a composite Latin noun meaning "swan wing" and refers to the shape of the gonopod coxa.
Colour. Faded after 35 years in alcohol (Fig. 1). Prozonites dark, greyish (perhaps originally deep brown); metazonites pale, almost whitish; dorsum with a thin black mid-dorsal line; legs pale brown to yellowish; head dark brown on the frontal part, paler towards the labral zone, mouthparts yellowish; preanal ring pale, anal valves blackish. Prozonites with oblique striae; metazonites with regular striation; suture complete, rectilinear but sometimes with a sinus at ozopore level. Anal valves with a marginal row of short setae, a submarginal row of longer ones, no setae (? broken) on disc. Subanal scale triangular, blunt and setose. Preanal ring with a protruding caudal projection bearing 2-3 setae on tip, no hyaline process (?broken).
Gonopods (Figs 2-3). Promerite (P) stout, basally parallel-sided in posterior view, twice as long as broad with a rounded lateral margin, distally gradually narrowing and ending in a narrow apical tip; mesal ridge (Mr) fairly broad, ending in a blunt process at mid-length of the promerite; posterior surface with a small concavity near the mesal ridge (lodging the mesomerite); rudimentary telopodite (T) small and located proximally near Mr (Fig. 3A). Posterior gonopod (Figs 2B-C, 3B-C): Mesomerite (Ms) reduced and very simple, less than half as long as the promerite; solenomerite (S) large, with an anterior long process (Sa) narrowing distad, apically tapering and bent posteriad, subapically bearing a triangular median hyaline lamella (Hl); seminal groove (g) running posteriorly from the fovea (F) located at the base of the solenomerite along a posterior acuminate process (Sp) and opening in 'og'; the solenomerite shows another thin and long process pointing laterad (Sl). Paracoxite (Px) large, twisted antero-laterad and extending beyond the rest of processes; distally abruptly expanding in a rounded bulge (Fig. 2 C, E) then apically narrowing in a curved thin apex pointing anteriad; coxa (Cx) very prominent, angular and characterised by the presence of a jagged margin with three protruding blunt processes, in lateral view reminding of the wing of a swan (Figs 2C, 3C).

Comments
The species was hitherto only known from a slide preparation of the gonopods of the holotype (NHMW) collected at an unspecified locality in Andalusia. The specimens from Portugal represent the first record of the species after its initial description.

Distribution
Northwest Portugal, North Spain, Southwest France.

Comments
Ommatoiulus bipartitus was described from Corunna (now: A Coruña) in NW Spain. All the Portuguese bipartitus specimens agree perfectly with Verhoeff's (1925) description of Schizophyllum involutum described from Portugal (Braga) and later synonymised with O. bipartitus (see Mauriès & Vicente 1977, Vicente 1985. The similarities between the two nominal species are unquestionable; however, the consistency in the shape of the paracoxite in the Portuguese/Galician population is rather striking. It differs from the 'real' bipartitus in having more square than rounded and curved margins, exhibiting an apical fold and bearing additional processes/teeth (see Fig. 5B). The promerite of the 'involutus' morphotype is a little more voluminous with a notably more prominent lateral margin, a larger mesoapical process and without the mid-apical tooth. The 'involutus' morph seems to occur in western Iberian sector, being also identified from Galicia. Another nominal species of relevance here is Ommatoiulus lienharti described from the Arcachon in southwestern France (see Brolemann 1921) and later recorded in northern Spain. O. lienharti shows important similarities with both morphotypes mentioned above (Fig. 5C). Mauriès (1971) included O. lienharti in his list of millipedes from Cantabria and Spanish Pyrenees, being certain that the species must occur in the area. A few years later, Mauriès & Vicente (1977) (Verhoeff, 1910) and O. dorsovittatus (Verhoeff, 1893), agreeing with these in the presence of a lateral notch on the promerite, a curved bifurcate mesomerite and a reduced paracoxite. Differing from both in the modified apex of the promerite, the deep curvature of the mesomerite and the shape of the solenomerite.

Etymology
The species epithet is a Latin adjective meaning curved and refers to the strongly curved shape of the posterior gonopods. Colour. After 35 years in alcohol (Fig. 6). Prozonites uniformly light grey perhaps with faded marbling, metazonites anteriorly pale, dominated by a strong reddish brown colour with dense black sputter posteriorly and showing a striking yellow band on the posterior margin, a thin black mid-dorsal line; legs brownish; head dark brown, mouthparts lighter, antenna dark brown; telson and anal valves brownish. Prozonites with oblique striae; metazonites with regular striation; suture complete, rectilinear but sometimes with a sinus at ozopore level. Anal valves with a marginal row of short setae, a submarginal row of longer ones, disc glabrous (setae broken?); subanal scale triangular, clearly protruding and setose; preanal process broken on all specimens. Gonopods . Promerite (P) in posterior view broad at the base, gradually narrowing distad, lateral margin expanded and rounded, distally incised in a rounded notch; mesal ridge (Mr) broad and fusing with the mesal margin; promerite apex marked with a fold with a strongly sclerotized and serrated process on the mesal margin, and a subapical notch on the lateral one; rudimentary telopodite (T) small, located basally, close to the mesal ridge. Posterior gonopod: mesomerite (Ms) broad, hookshaped and strongly curved, distally bifurcating into two thin asymmetrical and downturned processes pointing meso-basad; solenomerite (S) much shorter than Ms, bearing strong serrations on the anterior and apical margin and fringed with a large number of spikes, and a thin acuminate mesal process (Sm) pointing distad; posterior process of the solenomerite (Sp) showing an acute tip lodging the opening of  Ommatoiulus SP.6 - Bailey & De Mendonça (1990).

Diagnosis
Most similar to O. alacygni sp. nov. and O. staglae sp. nov., but differs by the three-digitate shape of the anterior process of the solenomerite and the strong serrations on the lateral margin of the paracoxite.

Etymology
The species epithet is a Latin adjective meaning "with small teeth" and refers to the serrations on the lateral margin of the paracoxite. Colour. Brownish, probably faded after more than 30 years in alcohol. Prozonites pale grey with black sputter, posteriorly forming dorsal triangular spots; metazonites yellowish-brown, with a black sputter posteriorly and scattered setae near the posterior margin (Fig. 9A); dorsum with a thin black mid-dorsal line; legs pale brown to yellowish; head dark brown with dense black sputter, lighter towards the labral zone, mouthparts yellowish; antennae and telson dark, with a dense black sputter. Prozonites with narrow oblique striae; metazonites with regular striation; suture complete, rectilinear but sometimes with a sinus at ozopore level. Anal valves with a marginal row of short setae, a submarginal row of longer ones and ca 5 setae on the disc. Subanal scale triangular, pointed and setose. Preanal ring ( Fig. 9B) with 2 setae on the sides, a protruding caudal projection bearing ca 8 setae and a small hyaline straight projection.

Holotype
Gonopods (FiGs 9C-F, 10). Promerite (P) in posterior view broad at the base, gradually narrowing distad, lateral margin rounded; mesal ridge (Mr) broad and fusing with the mesal margin; rudimentary telopodite (T) not conspicuous. Posterior gonopod: mesomerite (Ms) simple and very reduced; solenomerite (S) complex, broadest at the base, showing an large anterior process (Sa) apically ramified in 3 asymmetrical tips and bearing a subapical median triangular hyaline lamella (Hl); posterior process of the solenomerite (Sp) long with an acuminate tip lodging the opening of the seminal groove 'og'; seminal groove (g) running from the fovea (F) located at the base of the solenomerite up to og; paracoxite (Px) large, twisted meso-posteriad, distally leaf-shaped with a pointed apex and showing strong serrations on the lateral margin; coxa (Cx) large and rounded.

Distribution
South Spain (Ciudad Real, Andalusia, Estremadura) and south Portugal.

Comments
This is the first record of O. fuentei from Portugal. The species was initially described from Ciudad Real and was subsequently recorded from Estremadura and Andalusia (Akkari & Enghoff 2012).

Etymology
The species epithet is a Latin adjective meaning "of the coast" and refers to the affinity of the specie with coastal habitats.  Colour. Light brown, mostly faded in alcohol (Fig. 12). Prozonites dark brownish, with black sputter denser on the lateral side at the line of ozopores (Fig. 12B) and forming triangular spots along the middorsal line, metazonites pale whitish and translucent; dorsum with a thin black mid-dorsal line; legs pale brown to yellowish; head light brownish (Fig. 12A), mouthparts bright yellowish; anal valves with obviously faded black sputter, preanal ring pale (Fig. 12C). Prozonites with oblique striae; metazonites with regular striation; suture complete, rectilinear but sometimes with a sinus at ozopore level. Anal valves setose with a marginal row of short setae, a submarginal row of longer ones and 1-2 (? perhaps more, broken) setae on the disc. Subanal scale triangular, pointed and setose. Preanal ring with a protruding caudal projection with (5+5) setae and bearing a straight, relatively thick hyaline process. Gonopods (Figs 13,14). Promerite (P) in posterior view slender, subrectangular with parallel margins and slightly curved laterad; mesal ridge (Mr) thin, merging with the mesal margin and distally ending in serrated subapical process; apically almost straight, showing a small hyaline process extending beyond the apical margin; rudimentary telopodite (T) oval, consipicuous on the posterior surface at mid-length of the promerite. Posterior gonopod (Figs 13,14): Mesomerite (Ms) slender and curved anteriad to nest against the promerite, apically narrowing into a pointed tip; solenomerite (S) large with 3 distinct processes: an anterior process (Sa) the longest, gently narrowing apically and curved anteriad, a median process distally bearing a hyaline lamella (Hl) and a posterior process (Sp) the thinnest and most acuminate apically lodging the opening of the seminal groove 'og'; the seminal groove (g) running from the fovea (F) located at the base of the solenomerite up to og; paracoxite (Px) broad, distally expanding in a hammer-shaped process pointing posteriad; coxa (Cx) large and rounded, protruding distad.

Descriptive notes
Promerite with a round apex and a deep lateral notch ( Fig. 15A-B). Posterior gonopods complex ( Fig. 15C-D) with a large, club-shaped mesomerite ( Fig. 15E-F), distally expanded mesolaterad, apically with a thin process pointing distad. Solenomerite complex with a main short bulgy process, deeply furrowed, posterior half strongly serrated and apically protruding in an acuminate stem lodging the opening of the seminal groove, anteriorly with a long slender process; paracoxite large and deeply divided in two asymmetrical branches.

Distribution
Portugal.

Habitat
Predominantly in grassland but occurring in Cistus litter (Bailey & De Mendonça 1990).

Comments
Verhoeff (1895) described Schizophyllum (Eleutheroiulus) lusitanum from Coimbra without providing any illustration of the gonopods, but his verbal description, although in part difficult to interpret, is consistent with the much better described Archiulus cingulatus Attems, 1927. For example, in Verhoeff's description of the mesomerites ("Mittelblätter") he wrote (translated from German): "the latter ends apically in a pointed tooth. A second such [tooth] projects at approximately right angles on the lateral side somewhat distal to the mid-length", cf. Fig. 15E, F. Verhoeff donated a few syntypes of the species to NHMW; these were probably overlooked by Attems who in 1927 described Archiulus cingulatus based on material from 'Peisha' (probably Baja now), and for which he even erected the subgenus Schistocoxitus based on the unique shape of the paracoxite deeply divided in two processes (now also noted for the Andalusian species Ommatoilus sabinarensis Akkari, Mauriès & Enghoff, 2012).
The comparison side by side of the syntypes of O. lusitanum and O. cingulatus (see Fig. 15) from ZMB and NHMW collections respectively leaves no doubt that both names are synonyms. Therefore, Archiulus cingulatus is here considered as a junior synonym of Ommatoiulus lusitanus (Verhoeff, 1895). Mauriès, 1969 Ommatoiulus martensi Mauriès, 1969: 332-334 (Mauriès 1969). Both species have subsequently been identified from several localities in Western Pyrenees in Spain (O. martensi) and from northern Spain and the Basque country (O. haackeri). These two species have nearly the same gonopod configuration with main differences lying in 1) shape of the apical process of the promerite; 2) length of the distal process of solenomerite and 3) shape of the lateral process of the paracoxite (Mauriès 1969). After examination of several specimens we have failed to assign some of them to either haackeri or martensi with certainty. However, we assign the Portuguese species to O. martensi though a revision of the taxonomy of both (nominal) species is planned for a revision of the genus Ommatoiulus of northern Spain.

Distribution
O. moreleti is widespread in Portugal and western Spain (see maps in Baker 1984). It has been introduced to the Macaronesian Islands, Bermuda, South Africa and notably Australia, where it is known as a pest under the name of the "Portuguese millipede".

Comments
This species, which has become widespread through commerce, and which exhibits considerable gonopodal variation (e.g., Baker 1984), has been the source of much taxonomic confusion. Verhoeff (1910) described Schizophyllum cervinum from Serra da Estrela in Portugal and provided illustrations of isolated gonopod processes: promerite, paracoxite and mesomerite (see Abb. XI, XII, XXIV). In his description for the species, Verhoeff (1910) underlined the important similarities between this species and Ommatoiulus moreleti. The examination of Verhoeff's slide preparations of a cervinum ZMB3692, which we herewith designate as a lectotype for Schizophyllum cervinum (see Fig. 16) leaves no doubt that Schizophyllum cervinum is actually a junior synonym of Ommatoiulus moreleti (Lucas, 1860) with which it agrees in the shape of all gonopod processes.

Diagnosis
Most similar to O. armatus and O. dorsovittatus in the voluminous solenomerite with lobes and a serrated furrow, the presence of a lateral notch on the promerite and the reduced paracoxite. Different in the distal part of the mesomerite, promerite and a more slender shape of the solenomerite and paracoxite.

Descriptive notes
Promerite simple, in posterior view gradually narrowing in its distal half, apex rounded and lateral margin shallowly incised (Fig. 17A, C); mesomerite (Ms) large, extending beyond the rest of the processes, curved laterad and distally expanding into two oppositely directed processes (Fig. 17B, D); solenomerite (S) with a main process reaching ⅔ rd of mesomerite length, marginally strongly serrated, apically with a rounded lobe covered with short spines and a small pointed accessory process (Fig. 17D), paracoxite (Px) slender, almost as long as solenomerite (Fig. 17B, D) and emerging from a broad coxa (Cx).

Diagnosis
Small species, most similar to O. sabulosus (Linnaeus, 1758) both having a promerite with no incision and gently narrowing distally, a simple and slender mesomerite, a solenomerite bearing a long mesal projection lodging the opening of the seminal groove and a distal lamella, and a simple and stout paracoxite. Different in the shape of the distal lamella of the solenomerite and also in the shape of the mesomerite and paracoxite.

Comments
Ommatoiulus porathi was described verbally by Verhoeff (1893b), based on two males and six females. One year later, Verhoeff (1894) provided drawings of the mesomerite and opisthomerite, and in 1921 he included O. porathi in the identification key for the subgenus Bothroiulus (Verhoeff, 1921). The species has, however, never been collected again until now and has been regarded as dubious. The newly collected specimen agrees entirely with Verhoeff's description, and its gonopods agree perfectly with those of the type specimen (Fig. 18)

Diagnosis
Most similar to O. alacygni sp. nov. and O. denticulatus sp. nov. sharing the above mentioned characters for the group (see diagnosis of O. alacygni); different by the bipartite solenomerite with a long sinuous branch bearing a pointed accessory branch and a shorter one ending in a curved 'fish-tail' process and the absence of conspicuous serrations on the coxa and paracoxite.
Gonopods (Figs 19B,(20)(21). Promerite (P) in posterior view subrectangular short and broad with an apical broad tooth pointing posteriad; posterior surface biconcave with a conspicuous rounded rudimentary telopodite (T) located close to the mesal ridge (Mr); the latter broad and protruding in a short blunt process ending at mid-length of the promerite and separated from the mesal margin by a big rounded notch (Fig. 19B); lateral margin more or less straight, gently curving distad, apical margin oblique. Mesomerite (Ms) very reduced with a curved, downturned apex, shorter than the promerite and half as long as the anteriormost solenomerite process, bi-sinuous, nesting against the promerite posterior concavities; solenomerite (S) large, with two long and oppositely directed processes: the anterior process (Sa) longest, slightly curved, bearing an accessory branch at ⅔ rd length, distally gradually tapering into a thin curved spine directed posteriad, and the posterior process (Sp) gradually narrowing distally, apically abruptly expanding and bifurcating into a 'fish-tail' with two slender short tines. Seminal groove (g) running posteriorly from the fovea (F) (Figs 19B,21A) located at the base of the solenomerite up to og at the tip of Sp. Paracoxite (Px) narrow, expanded distally and bearing two apical asymmetrical processes pointing posteriad (Figs 20B, 21B). Coxa (Cx) large with a prominent posterior margin protruding in a rounded bulge.

Distribution
South Portugal, Algarve.

Diagnosis
One of the smallest species of the genus Ommatoiulus, different from all congeners by the combination of two distinct meso-apical teeth on the promerite, a complex solenomerite with subapical ramifications and a distally expanding mesomerite.

Etymology
The name is a Latin noun in the genitive, meaning "of the star" and refers to the type locality ('estrela' is Portuguese for 'star').  Colour. Brown, lighter on the lateral sides below the ozopore line and with two pale bands on the dorsum. Prozonites (Fig. 22) with scattered black sputter, becoming dense and discontinued with a round-oval pale spots sub-laterally, dorsum paler with median black spots; metazonites pale and translucent; dorsum with a thin black mid-dorsal line; legs light brown to yellowish; head brown with dense black sputter on the frontal part between the ocelli fields, lighter towards the labral zone, mouthparts yellow, antennae purple-brown, telson light brown to yellowish. Prozonites with oblique striae; metazonites with regular striation; suture complete, rectilinear but sometimes with a sinus at ozopore level. Anal valves with a marginal row of short setae, a submarginal row of ca 10 longer ones and ca 2 setae on the disc. Subanal scale triangular, blunt and setose. Preanal ring with a short triangular caudal projection, bearing a small hyaline process and 2+2 setae on the lateral sides.
Gonopods (Figs 22B-C, 23). Promerite (P) in posterior view subrectangular with parallel margins until ⅔ rd length where the lateral margin abruptly expands, bends anteriad, then gently curves towards the apex, the latter with two apical triangular teeth pointing posteriad; mesal ridge (Mr) fairly broad, ending in a blunt process at mid-length of the promerite; rudimentary telopodite (T) conspicuous, located proximally on the posterior surface (Fig. 23A), broad and rounded. Posterior gonopod (Figs 22-23): Mesomerite (Ms) shorter than promerite, with a broad base, gradually narrowing and bent laterad at mid-length (Fig. 23B), distally expanding meso-laterad (Fig. 23D) showing a smooth rounded apex, a serrated subapical margin, with a strong mesal tooth and a lateral downturned triangular process (Fig. 23D); solenomerite (S) large, with one main lamella marked with deep serrated notches and divided in dissimilar processes separating at different level and with a posterior serrated margin (Fig. 23B) and an anterior reduced process (Sa) short and pointing distad; the lamella distally shows a thin process curved antero-laterad and gently tapering into an acuminate tip pointing distad; a thin process with an acuminate tip and pointing posteriad and a slightly broader posterior process (Sp) lodging the opening of the seminal groove 'og', the groove (g) running posteriorly from the fovea (F) located at the base of the solenomerite up to og (Fig. 23B). Paracoxite (Px) stout, uniformly broad, mesal margin with a rounded projection at mid-length, apically rounded with a downturned serrated margin.

Distribution
Only know from high altitudes, 1798-1867 m in the Serra da Estrela Mountain massive, Portugal.

Discussion
Fifteen Ommatoiulus species are here recorded for Portugal among which six are new to science. In all species we described, the males share the secondary sexual characters consisting in the presence of mandibular stipes vertically expanded into rounded lobes; hook-shaped first pair of legs (Fig. 25) and the presence of postfemoral and tibial pads on subsequent legs.  (Verhoeff, 1893), right gonopods. E. O. fuentei (Brolemann, 1920), left gonopods.  Machado (1946)] as the record was not accompanied by any descriptive notes or illustrations, and confusion with another species seems likely.  (Attems, 1927 Attems (1928), Baker (1984) and Schubart (1966) who shed light on a few morphotypes. Among the morphotypes noted by Baker (1984), we suspect that the type D from Braga (northern Portugal) actually belongs to the O. bipartitus complex, while the type C from Coimbra largely agrees with some Spanish specimens we examined from Madrid and Avilà (unpublished data). Morphological variation was not only observed in O. moreleti but also in the other species from the group. Whether Schizophyllum involutum, long considered as synonymous with O. bipartitus (Mauriès & Vicente 1977), should be resurrected remains an open question as specimens perfectly matching Verhoeff's (1925) description and showing the characteristic shape of promerite and paracoxite have been found again in Oporto and in Braga, type locality of S. involutum but have also been identified from A Coruña in Galicia (unpublished), type locality of O. bipartitus.
Ommatoiulus bipartitus has, on the other hand, also been identified from northern Spain in Asturias and Leon (unpublished records) (Fig. 5A), whereas O. lienharti, now also regarded as a synonym of O. bipartitus, and described from Arcachon (SW France) has further been identified from Álava, Cantabria and the Basque Country. The lienharti and involutus morphotypes agree in the shape of the promerite (devoid of the apical projection) and a less curvy aspect of the apex of the paracoxite, typical of O. bipartitus. The lienharti and involutus morphotypes both exhibit a larger number of distal processes and teeth on the paracoxite than typical O. bipartitus. Whereas the typical bipartitus is confined to Galicia, the involutus and lienharti morphotypes have a wider distribution stretching from the French Pyrenees to Portugal. During our study, we observed also other morphs, for example the one from the Basque Country (Fig. 5D) and which we could not assign with certainty to any of the already mentioned ones. Thus we believe that only a detailed combined molecular and morphological study will elucidate the taxonomic status of these morphotypes/species and their phylogenetic relationships.
In a previous treatment of the genus Ommatoiulus ( (Brolemann, 1920) -the smallest known species of genus Ommatoiulus (body diameter less than 1.7 mm in males). O. stellaris sp. nov. is also notable by the presence of two distinct and strong apical teeth on the promerite, a structure similar to that seen in some Pachyiulini including the genus Dolichoiulus Verhoeff, 1900(Enghoff 1992 although the rest of the features of the promerite are completely Ommatoiulus-like while Dolichoiulus traits of the promerite like spoon-shape, protruding lateral margin, and mesal ridge, are absent.
The Ommatoiulus fauna of the Algarve region (see Fig. 26) comprises 3 new species, viz. Ommatoiulus alacygni sp. nov., O. denticulatus sp. nov. and O. staglae sp. nov. all characterized by a remarkably reduced mesomerite, a complex solenomerite and a large paracoxite. They might form an additional species group E. In O. staglae sp. nov. and O. denticulatus sp. nov., the posteriormost gonopod process shows further spectacular modifications, being strongly contorted and bearing big serrations on the paracoxite margin as in denticulatus or a strongly prominent and serrated coxa as in O. alacygni sp. nov. The reduced mesomerites are particularly interesting from an evolutionary point of view. A mesomerite (= an anterior, more or less separate process originating from the posterior gonopod) is regarded as an important evolutionary innovation in the family Julidae (Read 1990, see also Enghoff et al. 2011). Together with the promerite, the mesomerite in most cases forms a pair of forceps which have been shown to grab the operculum of the female's vulva during copulation (Haacker & Fuchs 1970, Tadler 1996. In Ommatoiulus the mesomerite may take numerous highly different shapes, and it is not always obvious that the forceps function has been retained. In the closely related 'genus' Rossiulus, the mesomerite is strongly reduced, a condition paralleled in Ommatoiulus group E. A comparative study on the copulatory mechanisms of Ommatoiulus species with differently modified/reduced mesomerites would probably be quite rewarding.