Type material

Holotype: MBCN 26604, female; Spain, Cádiz, Puerto Real, 36.659672 -5.672454; 28.02.2021; leg. Daniel Rojas. – Paratypes: MBCN 26605, female; same collection data as the holotype. – MBCN 26606, female; Spain, same locality as the holotype, 15.11.2020; leg. Daniel Rojas. – MBCN 26607, male; Spain, same locality as the holotype, 3.03.2021; leg. Daniel Rojas. – MBCN 26608, male; same locality and collection date; leg. Daniel Rojas. – MBCN 26609, female; same locality and collection date; leg. Daniel Rojas. – MBCN 26610, female; same locality and collection date; leg. Daniel Rojas. – CLLG 1057, male; same collection data as the holotype, dissected and mounted on slides. – CLLG 1152, female; Spain, Cádiz, Rota, 36.643283, -6.373049, 13.11.2021; leg. Antonio Pizarro, dissected and mounted on slides. All geographic coordinates correspond to World Geodetic System 1984 (WGS84).

Diagnosis: A species of Porcellio characterized by the following combination of morphological features, not found in any other species within the ‘laevis-hoffmannseggii’ complex: color pattern with longitudinal stripes on the pleon; glandular fields of tergites 2-7 located in the center of the epimera; antennal flagellum with articles of same length; first male pleopod exopod with elongated posterior lobe and rounded tip; endopod apically bilobed with fine long setae; carpal bulge of seventh male pereopod with its widest part on the distal third; uropodal exopods slightly longer in males than in females.

Etymology: Named after the andalusian naturalist Daniel Rojas Pichardo, the first collector of the new species.

Description: Body convex, with outline as in Fig. 2A. Colour (see Fig. 1): purplish brown with white spots; pereon and pleon with a darker central stripe that may be (or not) crossed by another narrower and lighter central stripe; cephalon pigmented except for the anterior parts of lateral lobes; pereonites 1-7 with unpigmented stripe delimiting epimera; epimera with anterior and lateral parts unpigmented and a posterior white spot containing the noduli laterales; pleon purplish brown with two para-median white stripes; pleon epimera pigmented, except for lateral white spots; antennae, pleotelson and uropods pigmented.

Dorsal surface of cephalon and pereon with granulations; a row of granules on the posterior half and the posterior edge of pereon tergites 1-7; pleon smooth, except for a row of small granulations on the posterior margin. Glandular fields circular or ovoid, with 25-30 pores each, separated from lateral margin, on a pigmented area. Integument with shingle-like scales (length 0.015 mm) and triangular scale-setae (length 0.02 mm), each composed by a triffid seta and a hyaline scale with blunt apex (Fig. 2B). Noduli laterales (length 0.07 mm) located in a circular unpigmented area; noduli 1 to 4 approximately equidistant from lateral margins (Fig. 2C). Cephalon (Figs. 2D-E) without suprantennal line; lateral lobes obliquely protruding frontwards and directed downwards, with rounded anterior edge; median lobe less protruding than lateral ones, widely rounded or subtriangular; eyes composed by about 35 ommatidia; front swollen, forming a differentiated blunt tubercle. Hind margin of pereonites 1-2 with marked concavity on each side; pereonites 3-4, with straight hind margin; posterior margin of pereonites 5-6, regularly concave; hind margin of pereonite 7, slightly sinuous; posterior tips of epimera 1-4 blunt, 5-7 more acute, directed backwards. Pleon (Fig. 2F) slightly staggered relative to outline of pereon; pleon epimera with posterior tips directed backwards.

Pleotelson (Fig. 2F) with differentiated basal and distal parts; basal part with slightly convex sides; tip extends slightly beyond posterior margin of uropodal protopods. Pleopods 1-2 with monospiracular internal lungs. Uropod protopods (Fig. 2F) with slightly oblique posterior edge, not reaching further than telson tip; exopods acuminate, approximately 1.7 to twice as long as protopods; endopods, visible dorsally, about 1.4 times longer than the protopods. First antenna (Fig. 2G) with 3 articles of similar length; distal article with 10-12 aesthetascs and triangular lobe.

Second antenna (Fig. 3A), when extended backwards reaches the second pereon-tergite hind margin; peduncular articles with short teeth; flagellar articles of equal length; distal article bearing three rows of aesthetascs (Fig. 3B) and dense setae field on the opposite side (Fig. 3C). Right mandible (Fig. 3D) with dichotomized molar penicil and 2 free penicils. Left mandible (Fig. 3E) with dichotomized molar penicil, 2 penicils on pilose lobe and 5 free penicils. Maxillule, external branch (Fig. 3F) with 4 + 6 teeth (2 slightly toothed); internal branch (Fig. 3G) with 2 long penicils and sharp posterior corner. Maxilla (Fig. 3H) inner lobe twice as wide as the outer; 2 setae between the two lobes. Maxilliped (Fig. 3I), first palp article with 2 setae; second palp article with 1 medial strong seta; distal article with tuft of short setae; endite sub-quadrangular, with 2+1 triangular teeth on anterior edge and 1 strong seta on caudal face.

Male: Pereopod 1 (Fig. 4A), merus and carpus with brush of setae on sternal margin (Fig. 4B); dactylar ornamentation as in Fig. 4C; pereopods 2-4 with brushes similar to pereopod 1, progressively less dense. Pereopod 7 (Fig. 4D), carpus with elongate lobe on tergal margin, with widest part on distal third. Pleopod 1 (Fig. 5A-C) exopod with elongated posterior lobe, about twice long as wide, distally rounded; endopod with distal part sinuous, apically bilobed with a tuft of fine setae on inner apex. Pleopod 2 (Fig. 5D-E) exopod triangular with straight inner margin; endopod slightly longer than exopod. Uropodal exopods slightly longer than in female.

Measurements: Maximum length of male, 11.1 mm; female, 14.2 mm.

Distribution: West of the province of Cádiz, Andalusia, in the south of the Iberian Peninsula. Probably more widespread.

Type locality: Puerto Real, Cádiz, Spain. Sandy area with sparse shrub vegetation.

Remarks: Although Porcellio danieli sp. nov. is closely related to some North African species of the genus, according to the available descriptions and illustrations it differs from all of them by a unique combination of morphological characteristics. Due to its somatic, integumentary and male sexual features, it can be classified in an intermediate position between the North African group (‘laevis’) and the Betic-Rifian group (‘hoffmannseggii ‘) as defined by Vandel (1951). With the species of the ‘laevis’ group, it shares the following morphological characters: median frontal lobe not very developed; rounded/oval glandular fields, containing numerous pores and located medially of pereon-tergites 2-7 (anterior position only on pereonite 1); weak dorsal granulations; exopodite of the first male pleopod triangular with elongated posterior lobe with rounded distal end. With the species of the ‘hoffmannseggii’ group, it shares the following characters: well-marked sinuosity of pereonites 1-2 and protuberance on the sternal edge of the carpus of the male seventh pereopod. However, according to Vandel (1951), these last two characteristics are also present in some species of the ‘laevis’ group (see discussion). Following Schmalfuss (1987b), both groups were considered herein a single systematic unit (see discussion).

The males of Porcellio danieli sp. nov. have the first pleopods with the posterior lobe of the exopodite ending in a rounded tip, and a bulge on the sternal border of the carpus of the seventh pereopod. Within the laevis/ hoffmannseggii group, only a few species present this combination of characters and, among them, only Porcellio magnificus Dollfus, 1892, has been recorded in the Iberian Peninsula. However, P. danieli sp. nov. differs greatly from this latter taxon by its somatic characteristics and dimensions (see Schmalfuss, 1987a).

The other species of the laevis/hoffmannseggii group recorded in the Iberian Peninsula, which characteristics are well-defined (see discussion), are the cosmopolitan Porcellio laevis Latreille, 1804, Porcellio hoffmannseggii hoffmannseggii Brandt, 1833, Porcellio flavocinctus Budde-Lund, 1885, Porcellio franzi Schmölzer, 1955, Porcellio ornatus Milne Edwards, 1840, Porcellio variabilis Lucas, 1849 and Porcellio wagneri Brandt, 1841.

Porcellio danieli sp. nov. differs from P. laevis in its habitus and dimensions, and in the shape of the first pleopod and the seventh pereopod of the male. It differs from all the other aforementioned species by having the posterior lobe of the exopodite of the first male pleopod triangular, with a rounded apex, instead of apically widened and notched (see Vandel, 1962; Schmalfuss, 1987a; Schmölzer, 1955; Medini & Charfi-Cheikhrouha, 1998; Medini-Bouaziz et al., 2006; Taiti & Rossano, 2015; Cifuentes, 2018).

The comparison with the male sexual characters of Porcellio ribauti Verhoeff, 1907, a North African species reported in Spain by Schmölzer (1955), is not possible because this species is only known from a single female specimen. According to its redescription (Caruso & Di Maio, 1990b), P. ribauti has a much broader body with expanded epimera and very large lateral cephalic lobes, whereas P. danieli sp. nov. has a more convex body, smaller dimensions and small lateral cephalic lobes.

In the Maghreb, several species belonging to the laevis/ hoffmannseggii group also have the exopodite of the first male pleopod with a triangular posterior lobe which is neither distally broadened and notched, but can be more or less sharp and possesses a rounded tip: Porcellio riffensis Caruso & Di Maio, 1990, Porcellio lepineyi Verhoeff, 1937, Porcellio atlanteus Verhoeff, 1937, Porcellio omodeoi Caruso & Di Maio, 1990, Porcellio villiersi Paulian de Félice, 1939, Porcellio gauthieri Paulian de Félice, 1939, Porcellio aghousi Paulian de Félice, 1939, Porcellio albinus Budde-Lund, 1885, Porcellio spinipes Dollfus, 1893 and Porcellio saharaiensis Di Maio & Dalens, 1991.

Porcellio danieli sp. nov. differs from P. riffensis in having a bulge on the carpus of the seventh male pereopod and by the morphology of the endopodite of the first pleopod. It differs from P. lepineyi mainly in the color pattern, the carpal lobe in pereopod 7 of the male (much more pronounced in adult specimens), the shape and structure of the endopodite of the first male pleopod, and the sexual dimorphism of the uropods. From P. atlanteus, it differs mainly in coloration, in the shape of the tergal edge lobe of the carpopodite of the seventh male pereopod, and in the shape of the endopodite of the first male pleopod. It differs from P. omodeoi mainly by the barely marked carpal lobe in this species, in the shape of the head lobes, and in the endopodite of the second male pleopod.

The remaining species have been ruled out for comparison because they are endemic to very remote mountainous areas or because they are clearly Saharan or steppe species with very different general somatic characteristics (dimensions, habitus, colour, appendages, etc.) (see Paulian de Félice, 1939; Caruso & Di Maio, 1990a, b; Di Maio & Caruso, 1991; Di Maio & Dalens, 1991).