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Pagurus pectinidactylus n. sp. (Figs. 2–4) lsid:zoobank.org:act: E87A7D6A-9982-43A6-B2D3-AE1D821153FF Type material. Holotype: ♂ sl. 3.2 mm (IEOCD-BR/2979), Mauritania, off Lemhaisrat, 19°00′09.7″N 16°23′38.0″W, sand, 21 m depth, 03 June 2012. Paratypes: 2 ovigerous ♀, sl. 3.9 and 2.9 mm (IEOCD-BR/2978), Mauritania, north of Nouakchott, 18°36′55.4″N 16°18′32.0″W, sand, 30 m depth, 03 June 2012. Description. Shield (Fig. 2A) flattened dorso-ventrally, broader than long, with tufts of short setae along shallow paragastric grooves and numerous low blister-like tubercles only visible when stained with methylene blue. Eleven pairs of phyllobranchia (Fig. 2B) with arthrobranchia on Mxp3, on cheliped and pereiopods 2–4, with one well-developed pleurobranchia on pereiopod 4. Rostrum broadly triangular, tip slightly sloping downwards, with small tubercle distally and produced to or slightly beyond level of obtusely triangular lateral projections; frontal margin between lateral projections thickened; lateral projections each armed with minute spine. Posterior carapace (Fig. 2D) membranous except for weakly calcified median and submedian parts, both with low blister-like tubercles and tufts of short setae; branchial region with tufts of simple setae. Ocular peduncles 0.7–0.8 length of shield, slightly swollen medially, with row of short setae medially; conspicuous oblique calcified scale covering the latero-proximal part; corneas slightly dilated; ocular acicles ovate, separated basally by 0.5 width of 1 acicle, terminating subacutely, with small submarginal spine and some setae on distal margin; dorsal surface shallowly concave medially; lateral extensions subquadrate, well-developed. Antennular peduncles (Fig. 2A, C) overreaching ocular peduncles by 0.3–0.4 length of ultimate segment; ultimate segment almost twice as long as penultimate segment, slightly widened distally, with tufts of long setae, one near distal margin, second medially; additional simple setae on proximal 1/3, one dorsal and one lateral; penultimate segment with few short setae; basal segment with lateral margin produced into blunt projection with narrow, blunt distal end; ventromesial distal margin produced as rounded projection; statocyst lobe unarmed; ventromesial distal angle unarmed. Antennal peduncles (Fig. 2A) overreaching ocular peduncles by approximately 0.3–0.4 length of fifth segment, and slightly longer than antennular peduncles. Second segment with dorsolateral distal angle strongly produced, reaching to 3/4 of fourth peduncular segment, terminating in strong spine, mesial margin with at least 10 additional small spines; dorsomesial distal angle with acute spine. Antennal acicles arcuate, reaching to proximal 0.4–0.5 of ultimate peduncular segment, with small terminal spine; mesial margin with row of rather stiff setae. Third maxiliped (Fig. 2E) moderately stout; dactylus about 0.8 length of propodus; carpus and merus unarmed; ischium with crista dentata with about 8 corneous teeth increasing in size and more widely spaced proximally, with 1 strong accessory tooth; basis with 2 minute denticles on mesial margin; exopod reaching distal margin of carpus. Chelipeds unequal and dissimilar, right appreciably larger than left. Male right cheliped (Fig. 3A–C) moderately stout, appreciably elongated. Chela subquadrate in dorsal view, about twice as long as wide. Dactylus approximately same length as palm, not overlapped by fixed finger when closed; dorsal surface not elevated, smooth except for short row of small spines proximally; longitudinal rows of tufts of setae on dorsal surface and longitudinal row of stiff setae along cutting edge; dorsomesial margin defined by single row of small, obtuse tubercles, with associated tufts of moderately long stout setae; ventral surface smooth, with 3 rows of tufts of long stout setae; cutting edge with row of 7 rounded calcareous teeth, smallest medially, terminating in small corneous claw. Palm including fixed finger 1.7 times longer than carpus; with row of blunt spines on dorso-mesial margin; dorsal surface flat, smooth, with clumps of stout short setae forming circular tufts arranged in longitudinal rows (Fig. 3F); dorsolateral margin elevated, clearly delimited by row of closely-spaced acute to subacute spines with associated short setae; space between spines widening distally, disappearing near tip; dorsomesial margin elevated, delimited by single row of spines with associated tufts of long setae; ventral surface slightly convex, smooth, with sparse tufts of long setae arranged in longitudinal rows. Cutting edge of fixed finger with 8 or 9 calcareous teeth, smallest proximally, terminating in small corneous claw. Carpus elongated, widened distally, about twice as long as maximum width and similar in length to merus; dorsal surface smooth, with tufts of stout short setae arranged in rows more dense on lateral and mesial margins; dorsomesial margin slightly elevated and clearly delimited with row of oblique moderately strong blunt or acute spines; dorsodistal margin unarmed, with rows of setae of different size; dorsolateral margin not well-delimited, with several rows of stout short setae; lateral surface slightly convex, perpendicular, smooth and almost glabrous; mesial face slightly concave, with scattered tufts of long simple setae, ventromesial margin forming a wind-like protuberance with spines continuing the dorsomesial row; ventral surface convex, with scattered tufts of moderately long setae. Merus subtriangular; dorsal surface with low protuberances with associated short setae; dorsodistal margin with 2 large and 1 smaller acute spines with associated short setae; lateral surface slightly convex, smooth, with few very short scattered setae; mesial surface slightly concave with scattered long setae; ventromesial margin smooth except for small spinulose area distally; ventrolateral margin not well-defined, rounded; ventral surface smooth, with scattered long setae. Ischium unarmed. Female right cheliped differs from males in length, being less elongated but similar in shape and armature to male. Male left cheliped (Fig. 3D, E) generally similar to right. Dactylus approximately 1.5 length of palm, dorsomesial margin with row of closely-spaced acute tubercles; dorsal surface smooth, with rows of tufts of moderately long setae; masticatory border toothless, except some very small proximally; rest of masticatory border bearing closelyspaced stout setae, forming a well-developed brush-like structure (Fig. 3D inset). Palm with row of subacute spines on dorsomesial margin, ending distally in two conspicuous spines; dorsal surface almost flat, unarmed; dorsolateral margin slightly raised and with row of closely-spaced subacute tubercles not extending to tip of fixed finger. Carpus with row of subacute oblique spines on dorsomesial margin; dorsal surface smooth, with longitudinal ridges of tufts of stiff setae; dorsodistal margin with several small spines. Merus with 2 spines on dorsodistal margin; ventromesial and ventrolateral margins each with row, or only few, very small blunt spinules or tubercles. Female left cheliped similar in shape to male, with more tubercles especially evident at dorsolateral surface of carpus, which is slightly elevated bearing some low tridentate tubercles with associated stiff setae, and merus, with dorsal surface crossed by rows of spinose sulcus, continuing along lateral and mesial surfaces, especially distal ones. Ambulatory legs (Fig. 4A, B) short and stout. Dactylus about same length (second) to 1.2 times longer than propodus (third) and about 5 times longer than broad, flattened laterally and gently curved ventrally terminating in strong corneous claw; lateral surface smooth, with very shallow longitudinal sulcus and few tufts of short setae on lower half; dorsal margin with irregular rows of tufts of short, thick setae, ending in row of long setae occupying distal 1/5; ventro-inner margins each with row of 5 or 6 (second) to 7 or 8 (third) strong corneous spines increasing in size distally, evenly distributed except most proximal of second pereiopod, which is more separated, and associated tufts of long setae. Propodus approximately 1.4 times longer than carpus (second and third); dorsal surfaces unarmed, each with multiple rows of short thick setae; lateral surface smooth, with longitudinal rows of short setae; ventral margins each with 3 corneous spines and tufts of short setae, denser near distal margin. Carpus with dorsal margin defined by row of small spines and associate tufts of short setae, with strong spine at dorsodistal margin; several rows of tufts of setae on lateral faces, short at upper half and near lower margin, longer at midline; ventral margin unarmed and glabrous. Merus with dentate dorsal margin, with associated rows of setae; lateral surface smooth and glabrous; ventral margins unarmed or with low protuberances or spinules, particularly on second pereiopods, and row of moderately long setae. Fourth pereiopods (Fig 4E) semichelate, subequal in length from left to right; dactylus with subterminal tuft of short setae on dorsal margin; carpus ending in conspicuous spine at distodorsal margin; stiff long setae and tufts of setae on dorsal margin of propodus to carpus and distodorsal and ventromesial margin of merus. Dactylus nearly straight, with convex dorsal margin, terminating in small, strongly-curved corneous claw; ventral margin with closely-set, microscopic corneous teeth; propodal rasp consisting of single row of corneous scales. Fifth pereiopods chelate, propodal rasp occupying distal half of segment. Sternite of somite XII in males (thoracomere 6, third pereiopods) with roundly subrectangular anterior lobe (Fig. 2F), bearing setae of different sizes on anterior margin. Sternite of somite XIV (thoracomere 8, fifth pereiopods) with widely separated lobes in males, each with moderately long setae (Fig. 4C); coxae of male each with gonopore partially masked by tuft of setae. Sternite of somite XII of females, with subquadrate single central lobe bearing moderately long simple setae (Fig. 4D); coxa of females each with gonopore, and associated tufts of setae above, without masking them. Males with 3 uniramous unpaired left pleopods; females with slightly unequally biramous 2-4, fifth uniramous.. Telson (Fig. 4F) with lateral incision indicating anterior and posterior portions; posterior lobes slightly asymmetrical, subquadrate, left slightly larger than right, separated by very small median cleft; terminal margins each with row of calcareous large teeth alternating with small ones, extending into lateral margin on left lobe and to posterior half of right; few short setae associated with spines and some tufts of longer setae on lateral margins. Distribution. So far only know from two off-shore stations, 21–30 m depth, within Mauritanian waters, north of Nouakchott. Etymology. Derived from Latin pecten (= comb) and Greek δάχτμλος (dáctilos = finger), referring to the characteristic row of closely-spaced stout setae on the masticatory border of left cheliped dactylus, forming a welldeveloped brush-like structure. Taxonomic remarks. The new species is assignable to the informal Pagurus anachoretus group (cf. Forest 1978) as it concurs with the general definition of the group (Forest & Ngoc-Ho 1992; McLaughlin & Forest 1999; Komai & Rahayu 2004). This group includes 12 species at present: Pagurus anachoretus from the Mediterranean, P. anachoretoides, P.gordonae, P.laurentae and P.souriei from the tropical eastern Atlantic, P.emmersoni and P.liochele from South Africa, and the Indo-West Pacific P. decimbranchiae Komai & Osawa, 2001, P. fungiformis Komai & Rahayu, 2004, P. hedleyi (Grant & McCulloch, 1906), P. kulkarnii Sankolli, 1961 and P. moluccensis Haig & Ball, 1998 (see Forest 1978; Forest & Ngoc-Ho 1992; McLaughlin & Forest 1999). Four other species from the tropical eastern Atlantic, P. alcocki, P. dartevellei (Forest, 1958), P. fimbriatus and P. triangularis, also fit the definition of the P. anachoretus group except for the cheliped form (Forest 1978; Forest & Ngoc-Ho 1992; McLaughlin & Forest 1999; Komai & Rahayu 2004). Pagurus fraserorum described by Landschoff (2018) also shares the main characters of the group except for the shape of the right cheliped, and based on molecular results, it has been suggested that it may constitute a separate group, more related with P. boriaustraliensis and P. pitagsaleei. Notwithstanding, the new species shows enough morphological differences to avoid any confusion with the species of the group or any other congeners known so far. The oblique calcified scale present in the base of the peduncles, is not known to occur in any other species of Pagurus. The shape and armature of the chelipeds are diagnostic, with a subquadrate shape in dorsal view, clearly elongate. The palm surface is flat and smooth, but with numerous distinctive clumps of stout short setae forming circular tufts arranged in longitudinal rows. The masticatory border of the left cheliped dactylus is also very conspicuous, lacking teeth, but bearing instead a row of closely-spaced stout setae, forming a well-developed brush-like structure, which has not been described for any other species of Pagurus in the study area. Although those characters are sufficiently distinctive to avoid any confusions, the new species shows a closer morphological resemblance with some species of the anachoretus group, specifically with Pagurus liochele, P. emmersoni, P. triangularis, P. fimbriatus and P. similimanus. The first two are considered endemic to South African waters. The new species differs from the two South African congeners in having broader carapace and rostrum, longer ocular peduncles, and more ovate ocular acicles. The antennules are shorter in P. emmersoni, overreaching distal margin of corneas by approximately 0.2 length of antennular segment 3, while they are similar in P. pectinidactylus n. sp. and P. liochele, overreaching the ocular peduncles by approximately 0.3–0.4 length of antennular segment 3. Another distinctive character can be found in the antennal segment 2 dorsolateral distal angle, which is strongly produced in both P. liochele and P. pectinidactylus n. sp., with mesial margin more densely dentate in the new species with at least 10 small spines, while in P. liochele there are 1–5. Moreover, the new species has a longer palm, with the characteristic clumps of setae arranged in circles on dorsal surface instead of any tubercles or spines, character that has not been described in any of the other species. The pereiopods are stout but more elongated in the new species, with the dactylus longer than propodus, bearing 5 (second) to 7 (third) strong corneous spines on ventral margin. The sternite of third pereiopods anterior lobe is roundly rectangular in P. pectinidactylus n. sp. and P. emmersoni, although with differences in setation, while in P. liochele the shape of the sternite is subovate. The other three species share also some morphological characters, the last two even with a distribution relatively close to that of the new species (see introduction). P. fimbriatus differs in the rostrum, which is obtuse, not pointed and not thickened. The shield is as long as broad while in the new species is broader than long. The antennal scales are shorter in P. fimbriatus, reaching only the base of the cornea, while in the new species reach distal end of corneas. Another important character can be found at dorsolateral distal angle of the second antennal segment, which is strongly produced in P. pectinidactylus n. sp., reaching to 3/4 of fourth peduncular segment, terminating in strong spine and with mesial margin bearing at least 10 additional small spines; based on the descriptions and figures provided by Forest (1966), this character is not present in P. fimbriatus. Pagurus triangularis and the new species share a pointed rostrum, but the former (like P. fimbriatus) has a narrower shield and shorter antennal acicles, which does not reach the distal margin of the cornea (at most it reaches its base), and also has a shorter dorsolateral distal process at the second antennal segment. P. similimanus also has a narrower shield, and a rostrum that is less projected than in the new species. The ocular peduncles are also shorter in P. similimanus, reaching to midlength of antennular segment 3, and proximal 1/3 of antennal segment 5, while in the new species reach the distal 1/3 of the antennular segment 3 and antennal segment 5. The antennal acicle is much longer, clearly exceeding the cornea in P. similimanus while in the new species reaches the distal margin of the cornea. The dorsolateral distal process of the second antennal segment has 6–8 little teeth on mesial margin, while in the new species the number is higher (up to 10). The shape of the chelipeds is also different in all these species, with a variable degree of elongation that never reaches that of the new species. The dorsal surface of the chelipeds is flat and unarmed in P. pectinidactylus n. sp., and the characteristic pattern formed by the clumps of setae is not mentioned in any of the other Eastern Atlantic species, neither the diagnostic masticatory border of the dactyl of the left cheliped formed by strong, densely arranged setae instead of teeth., Published as part of Almón, Bruno, Cuesta, Jose A. & García-Raso, J. Enrique, 2022, A new species of hermit crab of the genus Pagurus Fabricius, 1775 (Crustacea: Anomura: Paguridae) from north-western Africa, pp. 426-440 in Zootaxa 5209 (4) on pages 430-436, DOI: 10.11646/zootaxa.5209.4.2, http://zenodo.org/record/7333828, {"references":["Forest J. (1978) Sur deux pagurides nouveaux de l'Atlantique tropical africain: Pagurus laurentae et Paguristes cyanops spp. nov. Bulletin du Museium national d'histoire naturelle, Paris, 3 e serie Zoologie, 356, 525 - 538.","Forest, J. & Ngoc-Ho, N. (1992) Desription de Pagurus dartevellei (Forest, 1958) (Crustacea, Decapoda, Paguridae). Bulletin du Museum national d'histoire naturelle Section A Zoologie, biologie et ecologie animales, 14, 217 - 227.","McLaughlin, P. A. & Forest, J. (1999) Hermit crabs of the genus Pagurus Fabricius (Crustacea, Decapoda, Paguridae) from south-eastern South Africa. Annals of the South African Museum, 105, 297 - 344.","Komai, T. & Rahayu, D. L. (2004) Redescription of Pagurus moluccensis Haig & Ball, 1988, with description of a new species of Pagurus from Indonesia, and taxonomic notes on the Pagurus anachoretus group (Crustacea: Decapoda: Anomura: Paguridae). Raffles Bulletin of Zoology, 52, 183 - 200.","Komai, T. & Osawa, M. (2001) A new distinctive species of pagurid hermit crab (Crustacea: Decapoda: Anomura) from Japan. Zoological Science, 18, 1291 - 1301. https: // doi. org / 10.2108 / zsj. 18.1291","Grant, F. E. & McCulloch, A. R. (1906) On a collection of Crustacea from the Port Curtis district, Queensland. Proceedings of the Linnean Society of New South Wales, 31 (1), 2 - 53.","Forest, J. (1958) Les Crustaces Anomoures du Musee royal du Congo belge. Revue de Zoologie et de Botanique Africaines, 58, 144 - 168.","Landschoff, J., Komai, T., du Plessis, A., Gouws, G. & Griffiths, C. L. (2018) MicroCT imaging applied to description of a new species of Pagurus Fabricius, 1775 (Crustacea: Decapoda: Anomura: Paguridae), with selection of three-dimensional type data. PLoS ONE, 13 (9), e 0203107. https: // doi. org / 10.1371 / journal. pone. 0203107","Forest, J., 1966. Crustaces Decapodes: Pagurides. In: Campagne de la Calypso dans le Golfe de Guinee et aux Iles Principe, Sao Tome ' et Annobon (1956). Annales de l'Institut Oceanographique de Monaco, 44, pp. 125 - 172."]}

Integrative taxonomy studies have allowed us to clarify some taxonomic problems in cryptic species within species of InachusWeber, 1795 with two protogastric tubercles found in European waters. Several morphotypes of Inachus phalangium (Fabricius, 1775) are recognized, and a new species is described from the Atlantic coast of the Iberian Peninsula, for which distribution data are provided. Furthermore, two Inachus cf. thoracicus specimens with a unique sternal morphology were collected from Málaga Spain, but their preservation in formaldehyde prevented molecular analyses. Specimens of I. guentheri (Miers, 1879) have been re-examined and the validity of previous reports from tropical and subtropical Atlantic waters is discussed. These results allow us to separate and clarify the status of species within this group. An identification key for Inachus species with two protogastric tubercles is also provided., Financial support to EGO was obtained through a postdoctoral grant from the Spanish Ministry of Education (I-D+i 2008-2011). FP acknowledges the projects “CIDEGENT/2019/028 - BIOdiversity PAtterns of Crustacea from Karstic Systems (BIOPACKS): molecular, morphological, and functional adaptations” funded by the Conselleria d’Innovació, Universitats, Ciència i Societat Digital and “PRO2021-S02-PALERO - Fauna aquàtica en coves anquihalines del País Valencià: un món encara per descriure” funded by the Institut d’Estudis Catalans. We would like to thank the Junta de Andalucía PAIDI (Plan Andaluz de Investigación, Desarrollo e Innovación) for the funding of the groups RNM 141.

DIOGENES CURVIMANUS (CLÉMENT, 1874) (FIGS 5A–E, 6A–G, 7C, F, I, L) Pagurus curvimanus Clément, 1874: pl. III, fig. 1. Diogenes pugilator - Forest & Guinot, 1956: 32, fig. 3. Type material: Neotype: 1 ♂ 3.4 mm, Spain: Guadalquivir, Doñana National Park, Huelva, (MNHN-IU-2019-3214), 36°47 ′ 59.5 ′′ N, 6°23 ′ 27.3 ′′ W, sand, shallow sublitoral, 17 July 2018. To p o t y p e s: 1 ♂, 1 ♀, s a m e d a t a a s n e o t y p e, (IEOCD-BR/2581-2582); 5 ♂ (IEOCD-BR/2585). Other studied material: Belgium: Nieuwpoort-bad, 5 ♂ and 5♀ (IEOCD-BR/2611-2620), 51°09 ′ 08 ′′ N, 2°43 ′ 00 ′′ E, sand, intertidal, 29 September 2018; French Atlantic: Arcachon, 1 ♂ (IEOCD-BR/2621), 44°40 ′ 0.04 ′′ N, 1°10 ′ 34.5 ′′ E, sand, shallow sublitoral, 02 October 2018; Spain: Ribeira, Galicia, 1 ♂, 1 ♀ (ZSMA2019 0398-0399); 1 ♀ (IEOCD-BR/2603), 42°33 ′ 46.36 ′′ N, 8°59 ′ 15.07 ′′ W, sand, 4–5 m depth, 07 October 2017; Doñana N.P., Huelva, 2 ♂ (IEOCD-BR/2586), 36°48 ′ 13.6 ′′ N, 6°23 ′ 46.2 ′′ W, sand, intertidal, 09 September 2018; 2 ♂ (IEOCD-BR/2589,2595), 37°12 ′ 06.5 ′′ N, 7°01 ′ 37.0 ′′ W, sand, intertidal, 01 October 2020; 1 ♂, 1 ♀ (IEOCD-BR/2593-2594), 37°10 ′ 44.6 ′′ N, 7°20 ′ 37.3 ′′ W, sand, intertidal, 13 January 2020; 1 ♂ (IEOCD-BR/2592), 36°58 ′ 02.6 ′′ N, 6°30 ′ 32.2 ′′ W, sand, intertidal, 14 January 2020; 4 ♂ (IEOCD-BR/2587-88, 2591-91), 36°48 ′ 11.6 ′′ N, 6°23 ′ 39.3 ′′ W, sand, intertidal, 11 February 2020; Isla Canela, Huelva, 2 ♂, 1 ♀ (IEOCD-BR/2598-2600), 37°10 ′ 51.8 ′′ N, 7°20 ′ 15.3 ′′ W, sand, intertidal, 25 July 2014; Santibáñez, Cádiz, 3 ♂ (IEOCD-BR/2609-2610), 36°27 ′ 52.3 ′′ N, 6°15 ′ 21.4 ′′ W, sand, intertidal, 5 October 2015; 2 ♂ (IEOCD-BR/2605-2606), 15 October 2015; 2 ♂ (IEOCD-BR/2607-2608), 30 May 2018; Fuengirola, Málaga, 15 ♂, 7 ♀ (IEOCD-BR/2583-2584), 36°32 ′ 57.92 ′′ N, 4°36 ′ 30.89 ′′ W, sand, 4 m, 04 March 2014; La Carihuela beach, Torremolinos, Málaga, 3 ♂ (IEOCD-BR/2601-2602), 36°36 ′ 28.2 ′′ N, 4°30 ′ 13.6 ′′ W, sand, 2–4 m, 11 December 2019; Guadarranque, Algeciras, 1 ♂, 1 ♀ (IEOCD-BR/2696-2597), 36°10 ′ 49.7 ′′ N, 5°24 ′ 42.1 ′′ W, sand, 4m depth, 27 October 1995; San García, Algeciras, 1 ♂ (IEOCD-BR/2604), 36°06 ′ 17.5 ′′ N, 5°25 ′ 56.7 ′′ W, sand, 4m depth, 25 July 1996; Cabo Pino, Marbella, 1 ♂ (IEOCD-BR/2622), 36°29 ′ 05.6 ′′ N 4°45 ′ 00.4 ′′ W, intertidal, 28/04/2005. Redescription: Shield (Figs 5A, 7F) slightly broader than long; rostral lobe broadly rounded, exceeded by lateral projections that are triangular, acutely pointed, with single spine at apex; anterior margins of shield between rostral lobe and lateral projections slightly concave; anterolateral margins sloping, slightly concave, smooth; anterolateral angles rounded, with a small spine and usually another smaller below; lateral margin straight or slightly convex; posterior margin truncate; dorsal surface not vaulted, smooth, with groups of paired setae, and lateral margins with few faint transverse tuberculate ridges extending on to lateral surface of shield. Branchiostegites with dorsal margin bearing row of small spines (up to 12). Posterolateral plates not well calcified, unarmed. Ocular peduncles (including corneas) about 0.6 times as long as shield, moderately stout, slightly inflated in distal half; corneas not dilated, corneal diameter about 0.3 peduncular length; row of short, plumose setae over inner surface of the peduncles. Ocular acicles (Figs 5A, C) broad, subtriangular, concealing basal part of ocular peduncles, with slightly concave mesial margin; anterior margin sloping outwards, bearing 14–16 acute spines, decreasing in size towards outer margin, covering the entire length of the anterior margin; innermost distal spine distinctly larger. Intercalary rostriform process simple, shorter than ocular acicles (including spines), tapering acutely. Antennular peduncles (Figs 5A, B) overreaching distal corneal margin by about 0.4–0.6 length of ultimate segment, not reaching distal margin of antennal peduncle; third segment short, unarmed, broadened distally, maximum length slightly more than three times distal width, subequal in length to penultimate segment, with a tuft of setae on distodorsal margin; second segment unarmed, with tufts of plumose setae on dorsodistal and ventrodistal margins; basal segment moderately broadened distally, unarmed, except for tiny spinules on rounded ventrodistal border. Antennal peduncles (Fig. 5A) overreaching distal corneal margin by almost the entire fifth segment length (0.8–0.9); fifth segment with row of moderately short, stiff setae on ventral surface, and tuft of short setae dorsodistally, unarmed; fourth segment unarmed, with tuft of long setae on distomesial margin; third segment unarmed; second segment with distolateral outer process stout and acute, with subdistal smaller spine and sparse, short setae on lateral outer margin; distomesial spine usually small and obtuse but evident with associated tuft of long setae; mesial margin bearing some sparse, short setae; first segment unarmed. Antennal acicle short and broad, subtriangular, slightly overreaching the proximal half of fourth peduncular segment, not reaching distal margin, bearing a simple strong terminal spine plus usually six to seven spines almost equal in size along mesial margin, and tufts of setae on both mesial and lateral margins. Antennal flagellum short and robust, noticeably setose, with rows of paired setae in dorsolateral (short), ventral (medium) and ventrolateral (long) surfaces. Third maxilliped (Fig. 5D) basis unarmed; ischium with crista dentata bearing one prominent distal spine and two smaller on distal half, with additional one to two spinules on proximal half; ischium and merus with rows of scarce setae on lateral margins; carpus, propodus and dactylus with dense tufts of thick, long, simple and plumose setae at distolateral and distoventral margins, with less numerous long setae at distodorsal margin, concealing part of the segments; exopod peduncle reaching half carpal length of endopod. Male left cheliped (Figs 6A, B, 7C, F, I) much larger than right (Fig. 5E); length and shape variable in males. Dactylus shorter than palm dorsal surface, slightly arched, ending in small but strong calcareous claw, crossing tip of fixed finger; upper, inner margins defined rows of small, obtuse tubercles, upper, outer by larger acute tubercles with associated sparse setae; dorsal surface between upper, outer and inner rows slightly concave; outer surface convex, not flattened, smooth; lower margin with tufts of stout, short setae; cutting edge almost straight, with row of teeth of similar sizes; inner surface with two rows of tubercles running parallel to upper margin on upper half, reaching distal part of dactylus; proximal surface with small subacute to obtuse tubercles. Fixed finger triangular, not proximally broadened and almost smooth, except for some sparse, small tubercles on outer surface and irregular row of low tubercles on lower margin; tufts of stout, short setae on upper margin; inner surface smooth, with tufts of short setae near cutting edge and lower, inner margin. Palm narrow, about 1.5 times as long as high (max. medial length – max. height); upper margin shorter than carpus; dorsal surface of palm slightly convex, covered with evenly spaced obtuse tubercles and rows of short setae; upper, inner margin not well defined; upper, outer delineated by obtuse tubercles; outer surface glabrous, with upper part slightly concave, medially strongly and evenly convex, with numerous small, obtuse tubercles arranged as discernible longitudinal rows; lower palmar margin slightly concave in distal half, defined by row of large obtuse tubercles, and tufts of short setae; inner surface covered with low, rounded tubercles and sparse, short setae. Carpus of males long, about as long as merus and twice as long as high (Fig. 6A, B); usually longer and higher than palm; dorsal surface broadened, with upper, inner margin indistinctly delineated by row of tubercles; upper, outer well defined by curved row of subacute to obtuse tubercles, increasing in size distally, with associated medium-sized setae; outer surface with upper part slightly concave, medially strongly and evenly convex with small, obtuse evenly distributed tubercles, largest near distomesial margin; distal margin serrated; lower margin defined by row of small, obtuse tubercles; lower margin straight in distal part, short and slightly concave proximally, with tufts of short setae; inner surface covered with low, rounded tubercles and tufts of short simple setae. Merus longer than high (Fig. 6A, B, G); subovate in dorsal view; distal margin spinose throughout, with short setae; dorsal surface with small subacute tubercles of similar sizes and tufts of long, simple and plumose setae; lateral surface with evenly distributed rows of rounded tubercles, except on distal lower area; small and shallow transversal furrow subdistally on lower half, with sparse, short setae; ventrolateral margin denticulate accompanied by sparse mediumsized simple and plumose setae, slightly concave in proximal half, with larger subacute tubercle on proximal margin; mesial face with weakly calcified, u-shaped patch; subdistal transversal furrow short and faint, with small tubercles and short setae (Fig. 6G); dorsodistal margin with spines of similar size; ventrodistal margin defined by slightly larger spines with associated plumose setae. Ischium with row of small tubercles on distal margin (Fig. 6A) and row of slightly bigger tubercles on ventromesial margin (Fig. 6G). Variability. Propodus of male varies from form described to one where lower margin is strongly concave, giving a sinuous appearance. Carpus can also show on occasion a row of well-developed rounded tubercles at distomedial part. This is especially common in females and young males with short chelipeds (the reduction in size of the tubercles seems to be quicker because of the faster grow rate of male cheliped). Female left cheliped differs from male in the following features (Fig. 6C). Dactyl higher and shorter in proportion, with upper margin delineated by row of acute to subacute tubercles. Palm short and broad, slightly longer than high; outer surface covered in small, acute tubercles, with two prominent, acute tubercles distodorsally. Carpus relatively much shorter; upper, outer margin with two to three strong spines distally; outer medial surface with large, rounded tubercles increasing in size distally. Merus dorsal margin with larger, acute tubercles; lower margin defined by row of acute tubercles; lateral surface with subdistal transversal furrow more evident; mesial face with acute tubercles on ventral area and subdistal transversal furrow well defined. Right cheliped (Fig. 5E). Much shorter than left, robust (reaching one-third of carpus in neotype); dactylus and fixed finger with prominent hiatus, both terminating in small but strong calcareous claws. Dactylus (measured along mesial margins) about 2.5 times as long as palm upper margin, noticeably arched; upper, inner and outer margins defined by row of subacute tubercles with associated long setae; dorsal surface convex, broad and tuberculate; outer surface strongly convex with row of subacute tubercles parallel to the upper, outer margin; rest of outer surface with sparse, small tubercles and short, simple setae; inner surface with row of tubercles below upper, inner margin, and associated long, plumose setae; lower, inner margin with sparse setae. Palm with dorsal surface broad, short and convex; upper, outer and inner margins defined by row of subacute tubercles; outer surface broad, convex medially, with evenly spaced, subacute tubercles grading distally to acute tubercles; lower palmar margin defined by flat, setosed tubercles with long setae; fixed finger proximally broadened, outer surface slightly concave, with flat, setosed tubercles grading to acute near cutting edge; lower margin defined by flat, setosed tubercles; inner surface with low, rounded tubercles and long setae especially dense on fixed finger inner surface. Carpus dorsal surface narrow throughout; upper, outer margin defined by longitudinal row of spines and short setae; upper, inner margin defined by row of subacute tubercles with long, simple and plumose setae; additional row of small tubercles below the upper, outer margin, delimiting a shallow concave area; rest of outer surface slightly convex with low tubercles and numerous tufts of long setae; lower surface nearly smooth with sparse, short setae; inner surface weakly tuberculated, with sparse, long, plumose setae. Merus distodorsal margin with spinules and long setae; dorsal margin defined by row of small, obtuse spines and tufts of long, plumose setae; lateral surface covered with low, rounded tubercles and sparse, short setae, giving an almost smooth appearance; shallow tuberculate furrow subdistally, bearing short setae; ventrolateral margin delimited by row of small spines decreasing in size proximally, and tufts of long, plumose setae; mesial face with small, weakly calcified, u-shaped patch proximally, smooth, with ventromesial margin defined by row of minute spines and long, plumose setae. Ischium crenulated on distolateral and ventromesial margins. Second and third pereiopods slender (Figs 6D, 7L) subequal in length. Dactyl about 1.3 times as long as propodus, weakly curved; terminating in small corneous claw; upper and lower, outer margins unarmed, with rows of long thin setae, more numerous and longer on lower margin; outer surface with shallow, longitudinal sulcus medially, without setae in the proximal one-fifth, then continuing with sparse, short setae associated with sulcus; inner surface with two longitudinal rows of thin-spaced, long, stout setae adjacent to upper and lower margin, the proximal onethird with additional irregular rows of short, plumose setae placed over the sulcus. Propodus slightly longer than merus in second and third pereiopods, with upper margin defined by row of small acute tubercles (second), or faintly dentate (third), covered with long, simple setae; two rows of small tubercles running parallel to upper and lower margins, with tufts of short setae associated with tubercles; lower margin smooth, with sparse, short setae. Carpus upper margins defined by row of obtuse spines (second) or spinules (third), with dense, medium-sized, simple and plumose setae, ending in two (second) or one (third) conspicuous distal spines; rows of small, obtuse tubercles near outer upper, medial and lower surfaces with tufts of short setae in outer medial and lower surfaces; lower margin smooth with scarce short setae. Meral length about three times the maximum width; upper margin defined by small tubercles (second and third), with sparse, long, plumose setae; lower margin defined by row of low, small, subacute tubercles (second and third), with poorly developed spinules on lower distal margin (second) or without distal spines (third), and dense rows of long, plumose setae. Ischium unarmed, with long, plumose setae on distal margin. Fourth pereiopod (Fig. 6E). Dactyl with row of eight minute, spiniform setae on distal part of ventral margin. Propodus suboval, much longer than broad, with plumose, long setae along unarmed dorsal margin; propodal rasp consisting of five to six rows of corneous scales, covering distoventral part including fixed finger; rest of segments unarmed, with clumps of long, plumose setae. Fifth pereiopods. Propodus slightly shorter than merus, and about 1.7 times longer than carpus; group of subacute corneous scales in distodorsal surface of propodus and smaller ones in dactylus and fixed finger; long clumps of strong simple setae. Male unpaired left pleopods 2–5, uniramous, marginally setose. Female gonopores, paired; two to four unpaired pleopods well-developed, biramous; fifth pleopods without exopod, as in male. Telson (Fig. 6F) with shallow, median cleft, slightly asymmetrical; left posterior lobe slightly larger than right, with small spines on lateral margin, becoming blunt tubercles anteriorly; oblique terminal margin with larger spines; right posterior lobe with row of spines on less oblique terminal margin, extending on to posterior half of lateral margin. Coloration (Fig. 7C, F, I, L). Greyish white eye peduncles, with two diffuse bluish rings and a broader proximal brown triangular stain, projecting towards apex as a narrow line. Bluish ophthalmic scales. Whitish antennules, with triangular brown patch on second segment, orange in proximal two-thirds of third segment and flagellum. Antenna with greenish white or reddish peduncle, with scattered reddish brown streaks and conspicuous greenish blue spot on dorsal face of fourth segment. Carpus of the left cheliped with deep reddish orange background and an incomplete medial reddish brown ring. Orange coloured white hand with chestnut brown spot proximally, extending in two narrow lines towards middle outer surface; additional, more apparent, stain in proximal part of dactylus. Right cheliped with white background on merus and carpus, with some brown spots, and orange proximal carpal area; white palm with some brown spots in proximal upper area, which extends following central rows of tubercles. Second and third pereiopods with reddish white background colour; merus with reddish brown rings on middle (incomplete) and proximal areas. Carpus with an incomplete brown ring on medial area and small spot in the upper, proximal area. Propodus with large stain covering almost the entire proximal half, shorter in dorsal surface. Dactylus with a brown base that extends along the sulcus, sometimes faint and almost imperceptible. Habitat: Sandy beaches, in tide pools and subtidal areas up to 15 m depth, with maximum abundances in the shallow subtidal, between 4–, Published as part of Almón, Bruno, Cuesta, Jose A., Schubart, Christoph D., Armenia, Lisa & García-Raso, J. Enrique, 2022, Redescription of the hermit crab Diogenes pugilator (Decapoda: Anomura) reveals the existence of a species complex in the Atlanto-Mediterranean transition zone, resulting in the resurrection of D. curvimanus and the description of a new species, pp. 1116-1146 in Zoological Journal of the Linnean Society 195 on pages 1132-1142, DOI: 10.1093/zoolinnean/zlab093, http://zenodo.org/record/6993813, {"references":["Clement MC. 1874. Description d'un pagure nouveau. Bulletin de la Societe d'Etude des Sciences Naturelles de Nimes 2: 155 - 157.","Forest J, Guinot D. 1956. Sur une collection de Crustaces Decapodes et Stomatopodes des mers tunisiennes. Bulletin Station Oceanographique de Salammbo 53: 24 - 43.","Noel P. 2016. 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Review of the Pakistani species of Diogenes Dana 1851 (Decapoda Anomura Paguroidea Diogenidae). Tropical Zoology 17: 155 - 200.","El-Wakeil KFA, Ahmed ES, Obuid-Allah AH, El-Shimy NA. 2009. Hermit crabs (Crustacea: Decapoda: Anomura) inhabiting the intertidal and shallow subtidal region of Red Sea coast of Egypt. Zootaxa 2213: 57 - 63.","Siddiqui FA, McLaughlin PA. 2003. A new species of the hermit crab genus Diogenes (Decapoda: Anomura: Paguroidea: Diogenidae) from Pakistan, with a comparative diagnosis of D. guttatus Henderson, 1888. Proceedings of the Biological Society of Washington 116: 956 - 966.","Puillandre N, Lambert A, Brouillet S, Achaz G. 2012. ABGD automatic barcode gap discovery for primary species delimitation. Molecular Ecology 21: 1864 - 1877.","Lobo J, Costa PM, Teixeira MA, Ferreira MS, Costa MH, Costa FO. 2013. Enhanced primers for amplification of DNA barcodes from a broad range of marine metazoans. 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DIOGENES ARMATUS ALMÓN ET AL. SP. NOV. (FIGS 3A–G, 4A–G, 7B, E, H, K) The description of the new species in this paper has been carried out by four of the authors, B. Almón, J. Cuesta, C. Schubart and J.E. García Raso. Z o o b a n k r e g i s t r a t i o n: l s i d: z o o b a n k. o r g: a c t: 10CBBC54-4576-463A-AFE5-E7BE0D3A179A Type material: Holotype: ♂ 3.4 mm, Spain: Torregorda, Cádiz, (MNHN-IU- 2019- 3213), 36°26 ′ 51.7 ′′ N 6°14 ′ 49.4 ′′ W, sandy beach, intertidal shallow pools, 16/03/2015; Allotype: ♀ 3.8 mm, Isla Canela, Huelva, (MNHN-IU-2014-5736), 37°10 ′ 51.8 ′′ N 7°20 ′ 15.3 ′′ W, sand intertidal, 25 July 2014. Paratypes: Spain: Torregorda, Cádiz, 1 ♀ (IEOCD-BR/2645), 36°26 ′ 51.7 ′′ N 6°14 ′ 49.4 ′′ W, intertidal sandy beach, 27 July 2014; 1 ♂ (ZSMA2019 0402), 30 May 2018; 1 ♀ (IEOCD-BR/2643), 23 October 2014; 1 ♂ (IEOCD-BR/2644), 27 July 2014; 1 ♂ (IEOCD-BR/2642), 30/05/2018; 1 ♂ (IEOCD-BR/2646), 16/03/2015. Other material: Spain: Bajo la Cabezuela, Cádiz, 5 ♂ and 1 ♀ (IEOCD-BR/2623-2626), 36°31 ′ 43.6 ′′ N 6°15 ′ 01.0 ′′ W, intertidal, 30 August 2019; Es Torrent, Ibiza, 8♀ and 4 ♂ (IEOCD-BR/2629-2638), 38°58 ′ 02.2 ′′ N, 1°16 ′ 07.2 ′′ E, shallow subtidal, 26 May 2018; Port D’Es Torrent, 1 ♂ (IEOCD-BR/2639), 38º58.036 ′ N, 1º16.12 ′ E, shallow subtidal, 22 May 2018; Portugal: Lagoa da Albufeira, 2 ♂ (IEOCD-BR/2647-2648), 38°30 ′ 34.9 ′′ N, 9°10 ′ 28.6 ′′ W, sand, intertidal, 12 June 2018; French Mediterranean: Corsica, 1 ♂ (IEOCD-BR/2628), 42°42 ′ 49.8 ′′ N, 9°18 ′ 0.06 ′′ E, sand, shallow subtidal, 12 June 2003; French Atlantic: Arcachon, 1 ♂ (IEOCD-BR/2627), 44°40 ′ 0.04 ′′ N, 1°10 ′ 34.5 ′′ E, sand, intertidal, 02 October 2018; Tunisia: La Goulette, Tunis, 7 ♂ (IEOCD-BR/2640-2641), 36°49 ′ 10.80 ′′ N, 10°18 ′ 45.11 ′′ E, sand, 3–4 m, 24 November 2009. Etymology: The specific name armatus, masculine Latin adjective for armed, is given to highlight the spiny appearance of the chelipeds of this species compared to other Atlantic congeners, with more developed ornamentation composed mainly of spines and spiny tubercles instead of more rounded ones. Description: Shield (Figs 3A, 7E) slightly longer than broad; rostral lobe acutely rounded, exceeded by lateral projections that are triangular, acutely pointed, with single spine at apex; anterior margins between rostral lobe and lateral projections markedly concave and thickened; anterolateral margins sloping, slightly concave, mostly smooth, sometimes with some isolated minute spine; anterolateral angles rounded, with one spine on each side and another smaller submarginal spine sometimes inconspicuous; lateral margins slightly convex; posterior margin roundly truncate; dorsal surface slightly vaulted, with lateral margins usually cut by few transverse tuberculate ridges, extending on to lateral surface of shield; dorsal surface with additional short, transverse rows of small tubercles and tufts of short, stiff setae. Branchiostegites with dorsal margin bearing row of 10–11 strong spines, proximal two to three smaller. Posterolateral plates not well calcified, unarmed. Ocular peduncles (including corneas) about 0.6 times as long as shield, moderately stout, slightly inflated in upper half; corneas not dilated, corneal diameter about 0.25 peduncular length; row of short, plumose setae over inner surface of peduncles. Ocular acicles (Fig. 3A, C) subtriangular, with slightly concave mesial margin; anterior margin slightly convex, bearing 11–12 acute spines, decreasing in size towards outer margin, covering entire length of anterior margin; innermost spine distinctly larger and often slightly curved laterally. Intercalary rostriform process simple, shorter than ocular acicles, tapering acutely. Antennular peduncles (Fig.3A,B) overreaching distal corneal margin by about 0.4 length of ultimate segment, extending slightly beyond distal margin of antennal peduncle; third segment unarmed, not broadened distally, slightly less than five times longer than distal width, subequal in length to penultimate segment, with some isolated setae on dorsal surface and distal tuft; second segment unarmed, with tuft of plumose, short setae on dorsodistal margin; basal segment unarmed, except for the spinose, rounded, ventrodistal border. Antennal peduncles (Fig. 3A) overreaching distal corneal margin by 0.5 length of fifth segment; fifth segment unarmed, with row of moderately long, stiff setae on ventrolateral surfaces, and tuft of short setae on distodorsal margin; fourth segment unarmed, with tufts of plumose setae on disto-inner and some scarce simple setae on disto-outer margins; additional tuft of plumose setae on ventrolateral inner margin; third segment unarmed; second segment distolateral outer process stout and acute, with subdistal smaller spine; smaller but strong spine on distomesial angle; plumose setae near both spines and ventral surfaces; first segment with distal margin crenulated; plumose, short setae present in lateral and ventral surfaces. Antennal acicle short, subtriangular, reaching slightly below distal margin of fourth peduncular segment, bearing a simple strong terminal spine plus three to four strong spines on mesial margin and tufts of plumose setae on both mesial and lateral margins. Antennal flagellum short and robust, noticeably setose; articles with paired, long, ventrolateral and shorter ventral paired setae; short paired ones also on dorsal and outer articles surface. Third maxilliped (Fig. 3D) basis unarmed; ischium with crista dentata bearing three prominent (distal) and two smaller (proximal) spines; ischium and merus with rows of scarce setae on lateral margins; carpus, propodus and dactylus with dense tufts of thick, long setae in dorsodistal and dorsomedian margin, concealing part of the segments; exopod peduncle reaching one-third of endopod carpal length. Male left cheliped (Figs 4A–D, G, 7B, H) much larger than right (Fig. 3E). Dactylus about 1.2 times as long as upper margin of palm, gently arched, ending in large calcareous claw, crossing tip of fixed finger; upper, outer margin defined by rows of well-developed spines, inner margin by row of acute tubercles; two additional rows of subacute tubercles below upper, outer margin, first one incomplete; conspicuous strong spine at beginning of first row, near joint of dactyl with palm; dorsal surface between rows concave; rest of outer surface flattened, covered of small subacute tubercles; cutting edge sinuous, with row of calcareous teeth of various size, largest at distal half of dactyl; lower margin with irregular row of blunt sunken tubercles and tufts of setae associated with most of tubercles; inner surface with an additional complete row of rounded tubercles, separated by a depression from tubercles of upper, inner margin; inner, lower surface with two irregular rows of smaller sunken tubercles parallel to cutting edge, with tufts of dense setae associated with each tubercle. Fixed finger delimited proximoventrally by a shallow concavity separating the slightly convex outer surface of fixed finger and beginning of inflated palm surface; lower margin with two to three rows of small, rounded tubercles extending on to palm; a row of larger blunt sunken tubercles above this, another row parallel to cutting edge and a third medially, each with tufts of short, stout setae; rest of outer surface with small, blunt or subacute tubercles; cutting edge sinuous, with single row of various sized teeth, largest on proximal one-third; inner surface almost smooth except for three to four irregular rows of sunken tubercles, each with tufts of short dense setae. Palm robust, about 1.2 as long as high (max. medial length – maximum height); upper margin shorter than carpus; dorsal surface with four to six irregular rows of spines, space between rows widening distally with less produced spines; spines on upper, outer margin stronger and more produced, decreasing in size towards upper, inner margin; outer surface strongly inflated in proximal half, with proximal border defined by a crest of subacute tubercles; distal, lower surface depressed, near fixed finger; upper, outer part (below upper, outer row of spines), markedly concave; distinct bispinose protuberance present at distal angle just below upper margin; lower margin sinuous, convex proximally and slightly concave at distal half, defined by closely spaced rows of obtuse, spinose tubercles; rest of palm outer surface covered with spines of different sizes, biggest forming four longitudinal rows, usually evident especially when some colour remains; marginal series of subacute tubercles on upper, proximal half, as extension of proximal crest; transverse row of small spines at distal part of palm; inner surface covered with scattered rounded tubercles, with several rows of bigger tubercles below upper, inner margin; two additional well-defined rows, first on upper half, running from proximal medial margin to upper, distal margin, ending in a blunt protuberance, and second on midline zone, extending to half length of palm; sometimes a third faint row on lower half, composed by scattered tubercles; lower, inner surface, with irregular rows of low tubercles; most of largest tubercles with tufts of short thick setae. Carpus longer than high (Fig. 4A, B); dorsal margin with two to three irregular rows of spines; outermost row strongest; outer surface convex, covered with small spines becoming rounded tubercles near lower proximal area; row of stronger spines present on middle outer surface, largest near distal margin; broad almost smooth, concave area just below upper, outer and inner margins, becoming deeper and widening proximally; lower margin denticulate, gently protruded, forming a sinus proximally; disto-outer border dentate; pilosity abundant, with simple setae, except in lower, outer margin, where they are less abundant and plumose; inner surface covered with closely spaced, thick tubercles, and dense tufts of stiff, plumose setae; distal inner margin crenulated. Merus longer than high (Fig. 4A, G), subtriangular in dorsal view; distal margin with spines of different sizes, largest on dorsal area, with row of short setae; dorsal surface with rows of acute tubercles decreasing in size proximally and associated tufts of long, plumose setae; lateral surface with acute tubercles adjacent to dorsolateral margin and small spines near ventrolateral margin, with sparse setae associated with tubercles and spines; spinose transversal furrow subdistally, with short setae; rest of lateral surface smooth; ventrolateral margin denticulate, slightly concave in proximal half, with a row of strong acute tubercles increasing in size distally, accompanied by rows of plumose, long setae; mesial face with weakly calcified u-shaped patch, distally divided by shallow, transverse, unarmed furrow, with tufts of long setae; distal mesial part divided into dorsal and ventral lobes by median cleft; dorsal lobe with distal margin bearing spines of different sizes and tufts of medium-sized setae; ventral lobe with distoventral margin defined by large, acute tubercles, and dense tufts of long, plumose setae. Ischium with transversal row of small spines on distolateral margin and row of minute tubercles on proximal margin; mesial surface with longitudinal row of strong spines on ventral margin (Fig. 4G). Female left cheliped (Fig. 4E) differs from male in the following features. Palm oval, globose, almost as long as high, with lower margin convex throughout, without sinuous, concave area of males. Carpus lower margin with pronounced smooth sinus at proximal half; distal half protruding in a rounded projection defined by large spines of similar sizes. Merus dorsal and ventrolateral margins defined by well-developed spines. Right cheliped (Fig. 3E) appreciably shorter than left, robust, usually reaching proximal one-third of palm of left cheliped; dactylus and fixed finger with prominent hiatus, both terminating in small but strong calcareous claws. Dactylus (measured along mesial margin) slightly more than 2.0 times longer than palm, noticeably arched; upper, inner and outer margins defined by rows of small spines decreasing in size distally, with tufts of long setae arising from tubercles; outer surface with additional row of spines parallel to upper, outer margin with tufts of long setae; inner surface with two rows of tufts of long, stout setae, first on upper half and second near lower margin. Palm with outer surface not broadened, slightly flattened, with irregular rows of strong spines obscured by tufts of long setae, especially evident at midline and distal margin of propodus and fixed finger; upper, outer margin defined by row of strong spinose tubercles, inner margin not well defined; lower margin defined by row of small low acute tubercles; fixed finger not broadened proximally, cutting edge with row of small subacute calcareous teeth decreasing in size distally, and two rows of tufts of setae parallel to cutting edge. Carpus with row of large spines on upper, outer margin, increasing in size distally, and an additional row of small acute tubercles below it defining a concave area between them; rest of outer surface with low tubercles arranged regularly and numerous long, simple and plumose setae; lower surface unarmed; disto-outer margin spinose; inner surface smooth and glabrous. Merus distal margin with spines of different size, largest on dorsal area, with long, simple setae; dorsal margin with row of small, obtuse spines of similar size and tufts of long, plumose setae; lateral surface covered with low, rounded tubercles, with long setae; shallow furrow subdistally, with sparse setae; ventrolateral margin delimited by row of subacute tubercles, with subdistal spine and tufts of long, plumose setae; mesial face with small, weakly calcified, u-shaped patch proximally, smooth, with ventromesial margin defined by row of acute tubercles increasing in size proximally. Ischium with row of spinules on distolateral and ventromesial margins. Second and third pereiopods stout (Figs 3F, 7K) subequal in length. Dactylus 1.4 times as long as propodus, weakly curved; terminating in small corneous claws; upper, outer surfaces unarmed, with sparse simple setae of different length; lower with row of more numerous, long setae; outer surface with shallow, longitudinal sulcus medially, with row of setae below sulcus in proximal one-fifth, then continuing along upper border of sulcus; inner surface with two rows of long, stout setae running along upper and lower margins, and a short row occupying proximal one-fifth of inner sulcus. Propodus about same length as merus (second) or shorter (third), each with upper margin defined by rows of small spines (second), or faintly dentate (third), and with row of short to long stiff, simple setae combined with some plumose; lateral surfaces each with longitudinal row of setae arising from tiny, low, spinose tubercles near upper margin; lower margins smooth, with two rows of short setae; inner surface smooth and almost glabrous, with few sparse, plumose setae. Carpus upper margin defined by rows of strong spines (second) or smaller spines (third) increasing in size distally, with rows of simple and plumose setae; lateral surfaces with two rows of low tubercles in the upper half, with moderately long setae; lower surface smooth except for three to four tufts of stout, plumose setae on distal half; inner surface smooth and almost glabrous. Merus upper margin of second and third pereiopods defined by row of low, small, spinose tubercles decreasing in size distally, with tufts of long, plumose setae; lower margin of both pereiopods with row of low, spinose, small tubercles and long, simple and plumose setae; lower distal margin with a single spine (second) or without spines (third); rest of outer and inner surfaces smooth and glabrous. Ischium with upper and lower distal margin serrated, with long, plumose setae. Fourth pereiopods (Fig. 4F). Dactyl with row of 11 outer subacute submarginal spiniform setae. Propodus suboval, with a conspicuous spine dorsodistally and abundant plumose setae; propodal rasp consisting of seven to eight rows of corneous scales increasing in size distally, covering one-third of propodus distoventral surface. Carpus with a row of small tubercles dorsally, ending in a broad spine; with clumps of long, simple and plumose setae. Fifth pereiopods. Propodus slightly shorter than merus, carpus 1.5 times shorter than propodus; groups of subacute corneous scales in distodorsal surface of propodus and smaller ones in dactylus and fixed finger; long clumps of strong simple setae. Male unpaired left pleopods 2–5 uniramous, marginally setose. Female gonopores paired; two to four unpaired pleopods, well-developed, biramous; fifth pleopods without exopod, as in male. Telson (Fig. 3G). Asymmetrical, with small, median cleft; left posterior lobe distinctly larger than right, with strong terminal spine and row of smaller spines on lateral margin, becoming blunt anteriorly; oblique terminal margin with strong spines mixed with some small ones; right posterior lobe with row of strong spines mixed with small ones on less oblique terminal margin, becoming blunt anteriorly and extending on to posterior half of lateral margin. Coloration (Fig. 7B, E, H, K). Ocular peduncles reddish white, with brownish red, triangular stain covering around two-thirds of peduncle; eyeballs velvet black with honey yellow stippling. Ocular acicles orange proximally, turning greenish medially and white apically. Antennular background bluish; third segment and flagellum with diffuse orange patch mingled with background; second segment with dark brown dorsal stripe. Antennas translucent blue, going greenish at basal segments level; characteristic brown spot on antennal scale proximally. Merus and carpus of left cheliped greenish in background, with some diffuse tile red areas; brown transversal stripes on medial zone of merus, and base, medial and disto-inner areas of carpus. Palm greenish white, lighter than merus and carpus, with orange path in upper, proximal surface of dactylus, and some narrow ones following main spinose ridges; base of largest tubercles with orange taints; small, deep reddishbrown taints at mid-dorsal and apical area of palm. Right cheliped overall colour oil green, with white ve

DIOGENES PUGILATOR (ROUX, 1829) S.S. (FIGS 1A–F, 2A–F, 7A, D, G, J) Pagurus pugilator P. Roux, 1829 (in Roux, 1828 –30): part 3, pl. XIV, figs 3, 4. Pagurus varians O.G. Costa, 1838 (in Costa & Costa, 1838 –71). Type material: Neotype: ♂ 3.0 mm (MNHN-IU-2019- 3215), France, Mediterranean Sea: Frontignan, near Sète, 43°27 ′ 11.5 ′′ N, 3°48 ′ 49 ′′ E, sand, shallow subtidal, 29 August 2019. Topotypes: 2♂, same data as neotype, (ZSMA20190 400- 0401), 1 ♂ (IEOCD-BR/2664); 1 ♂ (IEOCD-BR/2676), 09 June 2018; 6 ♂ (IEOCD-BR/2665-2669), 29 August 2019. Other revised material: Spain: off Ebro Delta, 2 ♂ (ICMD 143 /1998), 40°35′N, 0°43 ′ E, 3–6 m, 4 May 1982; La Carihuela, Torremolinos, Málaga, 13 ♂ and 4 ♀ (IEOCD-BR/2660-2663), 36°36 ′ 28.2 ′′ N, 4°30 ′ 1 3.6 ′′ W, sand, 2–4 m, 11 December 2019; Fuengirola, Málaga, 1 ♂ and 1 ♀ (IEOCD-BR/2670), 36°32 ′ 57.92 ′′ N, 4°36 ′ 30.89 ′′ W, sand, 4 m, 4 March 2014; Pozuelo, Granada, 5 ♂ (IEOCD-BR/2677-2678 - 2679), 36 °44 ′ 37. 95 ′′ N, 3°39 ′ 30.47 ′′ W, sand, subtidal, 10 January 2020; Guadarranque, Algeciras, 1 ♀ (IEOCD-BR/2675), 36°10 ′ 49.7 ′′ N, 5°24 ′ 42.1 ′′ W, sand, 4 m depth, 11 December 2019; San García, Algeciras, 1 ♂ (IEOCD-BR/2674), 36°06 ′ 17.5 ′′ N, 5°25 ′ 56.7 ′′ W, sand, 4 m depth, 25 July 1996; Tunisia: La Goulette, Tunis, 1 ♂ (IEOCD-BR/2659), 4 ♀ (IEOCD-BR/2671-2673), 36°49 ′ 10.80 ′′ N, 10°18 ′ 45.11 ′′ E, sand, 3–4 m, 24 November 2009. Redescription: Shield (Figs 1A, 7D) subquadrate, nearly as long as broad; rostral lobe broadly rounded, exceeded by lateral projections that are triangular, acutely pointed, with single spine at apex; anterior margins of shield between rostral lobe and lateral projections slightly concave; anterolateral margins sloping, smooth; anterolateral angles rounded, usually with two to three small spines (two left and three smaller right in holotype); lateral margins slightly convex; posterior margin truncate; dorsal surface slightly vaulted, with lateral margins each usually cut by few transverse spinulose ridges extending on to lateral surface of shield; dorsal surface with additional faint short, transverse rows of small tubercles and tufts of short, stiff setae. Branchiostegites with dorsal margin bearing a row of eight to nine strong spines. Posterolateral plates not well calcified, unarmed. Ocular peduncles (including corneas) about 0.6 times as long as shield, moderately stout, slightly narrowed in middle; corneas not dilated, corneal diameter about 0.3–0.4 peduncular length; row of short, plumose setae on mesial margin of peduncles. Ocular acicles (Fig. 1A, B) subtriangular, with sinuous mesial margin; anterior margin slightly convex, bearing three to four (sometimes a small fifth) distal spines decreasing in size towards outer margin, innermost distal spine distinctly larger and often slightly curved; rest of anterolateral outer margin with 11–12 small tubercles of about the same size, occupying at least half of anterior margin. Intercalary rostriform process simple, slightly shorter than ocular acicles (including spines), tapering acutely. Antennular peduncles (Fig. 1A, C) overreaching distal corneal margin by about 0.8–0.9 length of ultimate segment, reaching distal margin of antennal peduncle; third segment unarmed, not broadened distally, about 3.5 times longer than distal width, subequal in length to penultimate segment, with three simple setae on dorsal margin and a distal tuft near distodorsal margin; second segment unarmed, with row of short setae on dorsodistal margin; basal segment unarmed, except for the spinulose rounded ventrodistal border. Antennal peduncles (Fig. 1A) overreaching distal corneal margin by 0.6–0.7 length of fifth segment; fifth segment with row of moderately long stiff setae on ventral surface and short setae dorsodistally, unarmed; fourth segment unarmed, with tufts of long setae on dorsodistal and ventrodistal border; third segment unarmed; second segment with distolateral outer process stout and acute, with subdistal smaller spine; smaller but strong spine on distomesial angle; plumose setae near both spines and ventral surfaces; ventrodistal border spinose; first segment with spinules of different sizes on distal border; plumose, short setae present on lateral and ventral surfaces. Antennal acicle short, subtriangular, reaching slightly below distal margin of fourth peduncular segment, bearing a simple strong terminal spine plus usually five spines along mesial margin, and tufts of setae on both mesial and lateral margins. Antennal flagellum short and robust, noticeably setose; articles with paired long ventrolateral setae and shorter setae on dorsal and ventral surfaces. Third maxilliped (Fig 1D) basis unarmed; ischium with crista dentata bearing two prominent and two small distal spines, with two additional variably sized spines on proximal half; ischium and merus with rows of scarce setae on lateral margins; carpus, propodus and dactylus with dense tufts of thick, long setae in dorsodistal and dorsomedian margin, concealing part of segments; exopod peduncle reaching one-third of endopod carpal length. Male left cheliped (Figs 2A, B, F, 7A, G) much larger than right (Fig. 1E). Dactylus about 0.8 times as long as palm measured along upper margin, slightly arched, ending in large calcareous claw, crossing tip of fixed finger; upper, inner margin well defined by a row of spinose tubercles, upper, outer margin by small spines; two more incomplete rows of small subacute tubercles running parallel to upper, outer margin; conspicuous one to two strong spinose tubercles at beginning of these additional rows, near joint of the dactyl with the palm; outer surface not flattened, covered with small subacute tubercles; lower margin with two rows of tufts of setae; some sparse setae covering outer surface and between dorsal rows; cutting edge sinuous, with row of calcareous teeth of various size, largest at distal half of dactyl; inner surface with irregular rows of tubercles in upper half running parallel to upper margin, reaching distal part of dactylus, defining a shallow concave area between the two rows; two irregular rows of tufts of dense plumose setae just above the cutting edge. Fixed finger delimited proximoventrally by a shallow concavity separating the slightly convex outer surface of fixed finger from markedly inflated palm surface; lower margin with two to three rows of small, rounded tubercles extending on to palm; outer surface covered with small subacute tubercles; cutting edge sinuous, with single row of various sized teeth, largest in proximal area, with a row of spaced tufts of setae below it; inner surface almost smooth with some tubercles on proximal surfaces and three rows (upper, medial and lower) of tufts of short, stout setae. Palm robust, about as long as high (max. medial length – maximum height); upper margin shorter than carpus; dorsal surface with irregular rows of spines, more produced on upper, outer margin, decreasing in size towards inner margin; space between rows widening distally and with less produced tubercles; outer surface strongly inflated in upper half, decreasing gently to lower region; distal lower part depressed near fixed finger; upper, outer part (below spinose row on upper margin) slightly concave, covered with small acute tubercles; distinct bispinose protuberance present at upper disto-outer angle; lower margin almost straight, defined by row of obtuse tubercles, with distal lower margin slightly concave; rest of palm outer surface with small spinose tubercles and some slightly larger subacute tubercles forming diffuse and not always well-defined rows; ventral margin of palm with blunt tubercles that continue along lower, inner face, defining a more or less flattened tuberculate area, delimited by sinuous crest-like row of large, rounded tubercles (Fig. 2B inset; see also Fig. 7G), defining an inflection change; rest of inner surface slightly flattened, covered with rounded, flat tubercles of different sizes and scarce simple setae associated with some of them; sometimes two rows of longitudinal slightly bigger tubercles outlined on upper half, and on medial zone, similar in extension reaching distal margin. Carpus longer than high (Figs 2A, B, 7A); upper margin with two to three irregular rows of strong spines; outer surface convex, covered with small spines; row of stronger spines present on middle outer surface, largest near distomedial margin; area just below upper, outer and inner margins slightly concave, becoming deeper and widening proximally, innermost shallower; lower margin denticulate, convex in distal two-thirds, forming a marked sinus proximally; distoouter margin spinulose submarginally; pilosity on outer surfaces abundant but not dense; inner surface covered with closely spaced thick tubercles and tufts of stiff, plumose setae. Merus longer than high (Fig. 2A, F), subtriangular in dorsal view; distal margin with spines of different sizes, largest on dorsal area, without setae; dorsal surface with rows of low acute tubercles, turning to obtuse spines distally, and tufts of long, plumose setae; lateral surface with small spines adjacent to dorsolateral and ventrolateral margins and spinose transversal furrow subdistally with sparse long setae, rest of lateral surface smooth and glabrous; ventrolateral margin denticulate, slightly concave in proximal half, accompanied by long, plumose setae and strong subdistal spines; mesial face with weakly calcified, u-shaped patch, distally divided by shallow transversal furrow dorsally bearing denticulate protuberances (Fig. 2F) and tufts of long thick setae; dorsodistal margin with spines of different sizes; ventrodistal margin defined by large acute tubercles with associated long, plumose setae. Ischium with short transverse row of small spines on distolateral surface (Fig. 2A) and longitudinal row of spines on ventromesial surface (Fig. 2F), with tufts of setae associated with spines and tubercles. Female left cheliped (Fig. 2C) similar to males; larger tubercles on outer surface of palm more numerous than in males, but arranged in similar way; lower margin of palm slightly more sinuous, faintly concave distally; lower margin of carpus only slightly convex, forming a less prominent sinus proximally; spines on dorsolateral and ventrolateral merus margins smaller than in males, rest of lateral surface with low, rounded tubercles. Right cheliped (Fig. 1E) appreciably shorter than left, robust, not reaching proximal margin of palm (usually midlength of carpus) of left cheliped; dactylus and fixed finger with relatively narrow hiatus, both terminatinginsmallcalcareousclaws. Dactylus slightly more than 2.0 times longer than palm (measured along mesial margin), gently arched; upper, inner and outer margins defined by rows of small tubercles with long setae; outer surface convex, covered with small spines arranged in rows; cutting edge with row of small calcareous teeth, regular in size, terminating in small calcareous claw and tufts of setae parallel to cutting edge; inner surface smooth except for tufts of setae parallel to cutting edge. Palm with upper, outer margin defined by a row of small spinose tubercles; outer surface broad, strongly convex, with irregular rows of small spinose tubercles, obscured by tufts of long setae; lower margin defined by small, obtuse tubercles; fixed finger broadened proximally; cutting edge with row of small, subacute calcareous teeth and two rows of tufts of long, stout setae parallel to the cutting edge; inner surface covered with evenly distributed low tubercles and associated long, plumose setae. Carpus widened distally, with row of strong spines on upper, outer margin increasing in size distally, and an additional row of rounded tubercles below it, delimiting a broad depression between them; outer surface with low tubercles and numerous tufts of long setae; lower surface nearly smooth, with some scattered low, rounded tubercles; inner surface with distal margin dentate, rest of inner surface covered with low, rounded tubercles and sparse tufts of short, plumose setae. Merus distal margin with spines of different size, largest on dorsal area, with long, simple setae; dorsal margin weakly delimited by a row of small, blunt tubercles, becoming obtuse small spines distally and tufts of long, plumose setae; lateral surface with small spines adjacent to dorsolateral margin and crenulate transversal furrow subdistally, with sparse, long setae, rest of lateral surface with small, rounded tubercles and sparse setae; ventrolateral margin delimited by a row of low, acute tubercles increasing in size distally, and tufts of long, plumose setae; mesial face with weakly calcified, u-shaped patch proximally, smooth, with ventromesial margin defined by row of acute tubercles of similar size. Ischium with short, transverse row of small spines on distolateral surface and longitudinal row of acute tubercles on ventromesial margin, with tufts of setae associated with spines and tubercles. Second and third pereiopods (Figs 2D, 7J) moderately stout, subequal in length. Dactylus about 1.2 times as long as propodus, weakly curved; terminating in moderately small corneous claw; upper and lower outer surfaces unarmed, with rows of long, simple setae more numerous in lower outer surface; outer surface with shallow, longitudinal sulcus medially, with row of setae along lower margin of sulcus in one-fifth proximal border, then continuing along upper border; inner surface with longitudinal rows of long, stout setae adjacent to upper and lower margins and a short row of plumose setae occupying proximal one-fifth of inner sulcus. Propodus about same length as merus (second) or shorter (third), with upper margin defined by row of spinules (second) or tiny blunt tubercles (third), and withrowofsetae; lateralsurfaceseachwithlongitudinal row of setae arising from tiny, low protuberances near upper margin and second inconspicuous row below midline; lower margins smooth with scarce short setae. Carpus upper margin with row of strong spines, increasing in size distally and with row of sparse, plumose setae dorsally (second), or with small spines on upper margin, with only one dorsodistal spine (third); lateral surfaces with low tubercles arranged in two rows, and setae associated with them; ventral surfaces with sparse, short setae. Merus upper margin with small spines increasing in size distally (second), almost smooth (third); lower margin dentate, with well-developed distal spine (second), or almost smooth, without distal spine (third); tufts of long, plumose setae on upper and lower surfaces. Ischium unarmed, with long setae on distal margin. Fourth pereiopods (Fig. 2E). Dactyl with row of nine to ten minute spiniform setae on distal part of ventral margin. Propodus suboval, with numerous setae on unarmed dorsal margin; propodal rasp consisting of five to six rows of corneous scales, covering distoventral part, including fixed finger; rest of segments unarmed, with clumps of long, plumose setae. Fifth pereiopods. Propodus almost as long as merus and two times longer than carpus; group of subacute corneous scales in distodorsal surface of propodus and smaller ones in dactylus and fixed finger; long clumps of strong simple setae. Male unpaired left pleopods 2–5 uniramous, marginally setose. Female gonopores paired; two to four unpaired pleopods well-developed, biramous; fifth pleopods without exopod, as in male. Telson (Fig. 1F) with small, median cleft, markedly asymmetrical; left posterior lobe with a strong terminal spine and with row of spinules on lateral margin, becoming blunt anteriorly; oblique terminal margin with few small spines; right posterior lobe with row of small spines on less oblique terminal margin, extending on to posterior half of lateral margin. Coloration (Fig. 7A, D, G, J). Whitish ocular peduncles with orange stain at base, projecting towards apex in diffuse narrow line. Antennules with the central dorsal area covered by a brownish orange line; honey yellow flagellum. Antennal scales with a vermilion red stain at the base; antennal flagellum with vermilion red rings evenly spaced. Characteristic vermillion red paired stains on branchial zone of cephalothorax. Left cheliped with snow white background colour; merus and carpus with the upper half orange with greenish tints, and a narrow scarlet red stripe in the middle zone, complete in the merus, partial in the carpus. Hand with broad wood brown well defined spots in: upper half of the base of the dactylus, lower distal area of the palm, and proximal central area of the palm; faint oil green areas on the distal area of the propodus, and pale orange on the upper, proximal area. Right cheliped snow white background colour; arterial blood red spot on the upper, proximal carpal area; orange zones on the superior distal zone of the carpus and proximal of the propodus; buff orange ring near the proximal area of the dactylus. Second and third pereiopods of buff orange colour at the dorsal part (with the proximal and distal parts darker), whiter at the ventral half; arterial blood red dorsal lateral stripes at the mid merus, carpus (incomplete), propodus and base of the dactylus. Habitat: Sandy beaches of tidal lagoons along intertidal or shallow subtidal areas, up to 25 m, although more frequent around 5 m depth. Distribution: Known with certainty from the French Mediterranean (Frontignan, neotype locality), northeastern (Barcelona), and south-eastern Iberian Peninsula (Málaga, Granada, Algeciras) and Tunisian coasts (La Goulette). By the figure and coloration of the large cheliped in Costa (1839, T2, f2), its presence in Italy (Gulf of Naples and Taranto), seems also probable. Remarks: A number of subspecies or varieties of D. pugilator have been established based on morphological differences, e.g. Diogenes pugilator orientalis for the Black Sea and Adriatic populations (Codreanu & Balcesco, 1968). Currently, none of the European varieties have been accepted so far, because of the consideration of D. pugilator as a highly variable species (Ingle, 1993). The application of molecular techniques has shown that, even if this consideration is true to a certain extent, there is in fact a number of different well-defined species, previously lumped under D. pugilator and that in future studies, some of these synonymized taxa could be revalidated in the light of new and updated information. Since the type specimen, Published as part of Almón, Bruno, Cuesta, Jose A., Schubart, Christoph D., Armenia, Lisa & García-Raso, J. Enrique, 2022, Redescription of the hermit crab Diogenes pugilator (Decapoda: Anomura) reveals the existence of a species complex in the Atlanto-Mediterranean transition zone, resulting in the resurrection of D. curvimanus and the description of a new species, pp. 1116-1146 in Zoological Journal of the Linnean Society 195 on pages 1119-1127, DOI: 10.1093/zoolinnean/zlab093, http://zenodo.org/record/6993813, {"references":["Roux P. 1828 - 1830. Crustaces de la Mediterranee et de son littoral, decrits et lithographies. Paris and Marseille: Editions Levrault. [176 unnumbered pp., 45 pls; published in 9 parts: 1, 2 (pls 1 - 10), 1828; 3 (pls 11 - 15), 1829; 4 - 9 (pls 16 - 45) 1830.]","Costa OG. 1838 - 73. Fauna del Regno di Napoli ossia enumerazione di tutti gli animali che abitano le diverse regioni di questo regno e le acque che le bagnano contenente la descrizione de nuovi o poco esattamente conosciuti con figure ricavate da originali viventi e dipinte al naturale. Animali molli. Naples: Azzolino & Compagno [325 pp. with each chapter paginated separately, 42 pls also numbered separately according to chapters].","Codreanu R, Balcesco D. 1968. Etude biometrique comparee de certains caracteres dans deux populations du pagure Diogenes pugilator (Roux) de la Mer Noire et de l'Ocean Atlantique. Bulletin Biologique de la France et de la Belgique 102: 369 - 383.","Ingle R. 1993. Hermit crabs of the northeastern Atlantic Ocean and Mediterranean Sea. An Illustrated key. London: Natural History Museum Publications, Chapman and Hall."]}

An annotated checklist of the marine decapod crustaceans (excluding crabs) of the Iberian Peninsula has been compiled 50 years after the publication of “Crustáceos decápodos ibéricos” by Zariquiey Álvarez (1968). A total of 293 species belonging to 136 genera and 48 families has been recorded. This information increases by 116 species the total number reported by Zariquiey Álvarez in his posthumous work. The families with the greatest species richness are the Paguridae (28) and Palaemonidae (18). References by geographic sectors and for all species are given. The results show that 264 species are reported in the Atlantic sectors, while 178 have been found in the Mediterranean. The species richness and the differences between and within sectors are discussed; these are mainly due to the dimension of the areas, the depth ranges and the confluence of distinct water masses with a different origin and different physicochemical features. Consequently, the greatest richness of decapod species (excluding crabs) is found in the Gulf of Cádiz, with 194 species. The total number of decapods found in and around Iberian waters, including crabs, freshwater species and some new records not yet published, reaches 448., Se ha realizado un listado comentado de los crustáceos decápodos marinos (excluidos los cangrejos) de la península ibérica, después de 50 años de la publicación “Crustáceos decápodos ibéricos” por Zariquiey Álvarez (1968). En total se mencionan 293 especies de 136 géneros y 48 familias. Estos datos incrementan en 116 el número de especies citadas por Zariquiey Álvarez en su obra póstuma. Las familias con mayor riqueza específica son Paguridae (28) y Palaemonidae (18). Se citan referencias para cada sector geográfico y especie. Las aguas atlánticas (en su conjunto) muestran una mayor riqueza, con 264 especies, mientras que 178 han sido citadas en aguas mediterráneas ibéricas. Se da el número de especies por sectores y se comentan las diferencias, consecuencia, principalmente, de la dimensión y rango de profundidad de cada uno de ellos y de la confluencia de masas de aguas con orígenes y caracteríticas fisico-químicas diferentes. Así, la mayor riqueza específica (sin incluir los cangrejos) se encuentra en el Golfo de Cádiz con 194 especies. El número total de especies de crustáceos decápodos en aguas ibéricas, incluyendo cangrejos, especies de agua dulce y algunas nuevas citas de especies aún no publicadas, sería de 448.

Trabajo presentado en la The Crustacean Society Mid-Year Meeting (TCS), celebrado en Barcelona del 19 al 22 de junio. de 2017.-- Esta reunión de la Sociedad internacional de Crustáceos se incluyó el 11th Colloquium Crustacea Decapoda Mediterranea y la 5th Crustacean Larval Conference., Project CGL2009-11225 funded by MINECO, Plan R + D + I and FEDER; Carlos Sánchez, Marta Pascual, Asvin P. Torres, Enrique Macpherson, Paul F. Clark and Mº Grazzia Mazzocchi. FPI Fellowships BES- 2010-033297 supported by MINECO (BES-2010-033297).

Trabajo presentado en la The Crustacean Society Mid-Year Meeting (TCS), celebrado en Barcelona del 19 al 22 de junio. de 2017.-- Esta reunión de la Sociedad internacional de Crustáceos se incluyó el 11th Colloquium Crustacea Decapoda Mediterranea y la 5th Crustacean Larval Conference.

Trabajo presentado en las VIII Jornadas de Acuicultura en el litoral Suratlántico, celebradas en Cartaya (Huelva) el 7 y 8 de junio de 2016.

8th International Crustacean Congress (ICC-8), 18-23 August 2014, Frankfurt am Main., The main objective of the MEGALOPADN project was to optimize the application of molecular techniques, specifically the analysis of mitochondrial DNA sequences (16S and Cox1), in conjunction with morphological techniques for a correct identification of brachyuran megalopae collected from the plankton around the Iberian Peninsula. The list of Iberian crabs was updated to 136 valid species (from 113 previously). A database was built including the corresponding 16S and/or Cox1DNA barcode sequences for 123 of those species. Thanks to this database, the 90% of the Iberian crabs (adults or larval stages) can now be identified with confidence to species level. The remaining 10% are mainly rare and infrequent species. Furthermore, the megalopae of 57 different species were identified using this database after analyzing morphological and molecularly more than 2000 megalopae collected in the plankton. The megalopal phase was unknown for 11 of these 57 species, and they are being described in detail. Megalopae of the remaining 46 species, of which previous descriptions were available, have been used to revise characters, setation patterns, missing data, correct minor mistakes and analyze the variability found among megalopae from the plankton and those obtained in the laboratory. Megalopae had to be re-described in 3 species because previous descriptions were incomplete or contained too many errors. These new descriptions and redescriptions, along with the existing ones, have enabled the development of an illustrated key for the identification of the megalopae of 78 Iberian crab species. The availability of this database of DNA barcodes has also allowed us to study the phylogenetic relationships of these species and their taxonomic validity. As a result there have been a number of proposed changes in the systematics of some species, comprising synonymization, establishment of new genera and a new subfamily, and changes in the systematic position of the genus Ergasticus (MARCO-HERRERO et al., 2013), This study was funded by the Spanish “Ministerio de Economía y Competitividad, Plan Nacional I+D” and FEDER through project MEGALOPADN (CGL2009–11225)

Study of hermit crabs from Alboran Sea has allowed recognition of two different morphological forms under what had been understood as Pagurus forbesii. Based on morphological observations with various species of Pagurus, and molecular studies, a new species is described as P. pseudosculptimanus. An overview of species of Pagurus from the eastern Atlantic and Mediterranean Sea is provided. © Copyright 2014 Magnolia Press., This work was supported by the Junta de Andalucía RNM-0141 Research Group.

The freshwater shrimp genus Atyaephyra de Brito Capello, 1867 is a complex of cryptic species with morphologically overlapping features, which has led to invalid identifications until recently. The genus is distributed around the entire Mediterranean Basin and surrounding areas. In 2009 we used a molecular approach with 16S and Cox1 genes to identify the European and African populations and to show the existence of different species and groups of haplotypes (García Muñoz et al. 2009). Christodoulou et al. (2012) recently identified four species in Greece (Atyaephyra stankoi, A. thyamisensis, A. strymonensis and A. acheronensis) based on morphology and some partial sequences of Cox1. In the present study we analyse several Greek populations of Atyaephyra spp. using morphology and two mitochondrial markers (16S and Cox1). Our molecular data confirm the taxonomic validity of A. stankoi, A. thyamisensis and A. strymonensis but show that A. acheronensis should be considered a synonym of A. desmarestii. The limited distribution of A. desmarestii in only a few rivers in the Ionian region is probably the result of a posterior introduction from neighbouring European areas. After the genetic identification and as a consequence of the high variability observed within species and between rivers, mainly in A. thyamisensis, we re-describe this species and provide morphological data for the other species. We also give information on the chromatophore pattern in larvae, which could be very useful in the future for recognizing species and their distribution. The distribution of Greek Atyaephyra species seems to be related to the very complex paleogeographical history of the Paratethys and Mediterranean Sea in the Balkan Peninsula. Three areas can be recognized in relation to the distribution of these species: a global region, a western region separated from the eastern region by the Pindos Mountains, and a Macedonia-Thracian region. Copyright © 2014 Magnolia Press., This study was funded by the Spanish Ministry of Science and Education, National Plant I+D and European Regional Development Fund (ERDF)” through project CGL2004–01083/BOS. Collections in Greece were possible thanks to additional funding provided by “CSIC Proyectos Intramurales Especiales” 2004 3 0E 099 and 2006 3 0I 238 to A. Rodríguez and J.A. Cuesta, respectively.

The Crustacean Society & Latin American Association of Carcinology Summer Meeting, Costa Rica, San José, 07-11 July 2013, Many marine invertebrates present complex life cycles comprising several developmental stages which clearly differ morphologically from those finally reached by adults. This is the case in the brachyuran crustacean decapods. Most brachyuran crabs pass through a planktonic larval period with two phases, zoea and megalopa, which are very different from each other and from the adult form. This represents an important drawback in the identification of the brachyuran larvae collected from plankton. The main objective of the present study is the use of molecular techniques, DNA barcodes (mitochondrial DNA sequences 16S and Cox1), and morphological analysis, in the accurate identification of Iberian brachyuran megalopae obtained directly from planktonic samples. We have presently identified 36 different planktonic megalopae after examination of more than 2100 specimens. Twenty-six of them are megalopae of species already described from laboratory cultures that have been revised in order to look for differences with respect to wild specimens. Moreover, some of them also need to be re-described, considering that previous published descriptions were inaccurate or incomplete (i.e. Rhithropanopeus harrisii, Dyspanopeus sayi, Percnon gibbesi). Ten megalopae unknown until now (Sirpus zariquieyi, Derilambrus angulifrons, Polybius zariquieyi, Ergasticus clouei, Afropinnotheres monodi, between others) will be described in detail. These new descriptions, together with the formerly existing ones, will allow the creation of an illustrated identification key, as a final objective, addressed to researchers and technicians, which is intended to assist in the correct identification of this important group of planktonic organisms

The Crustacean Society Summer Meeting (TCSSM) - 10th Colloquium Crustacea Decapoda Mediterranea (CCDM), 3-7 June 2012, Athens, Greece, Many marine invertebrates present complex life cycles comprising several developmental stages which clearly differ morphologically from those finally reached by adults. This is the case of the brachyuran crustacean decapods. Most brachyuran crabs, with the exception of a few species that show direct development, pass through a planktonic larval period with two phases, zoea and megalopa, which are very different from each other and from the adult form. This represents an important drawback in the identification of the brachyuran larvae collected from plankton, which is further complicated by the fact that reliable larval descriptions are only available for a small amount of known brachyuran species, most of them based in laboratory-reared specimens. This gap in the knowledge of brachyuran larval morphology is in turn responsible of an important amount of problems encountered by researchers when studying population dynamics, recruitment events, larval dispersal and colonization, functioning of planktonic trophic webs (inter-specific interactions) and, overall, any kind of biodiversity research. The main objective of the present study is the optimization of the joint applicability of molecular techniques, DNA barcode, particularly the analysis of mitochondrial DNA sequences (16S and Cox1), and morphological analysis, in the accurate identification of Iberian brachyuran megalopae obtained directly from planktonic samples. The first step needed to achieve this objective, is the setting-up of a complete database containing the sequences of the above-mentioned markers, for the 140 bachyuran species recorded in the Iberian Peninsula. Remarks We have presently obtained the DNA barcode for more than 80% of the Iberian crabs, and identified 26 different planktonic megalopae after examination of more than 800 specimens. Sixteen of them are megalopae of species already described from laboratory cultures that have been revised in order to look for differences with respect to wild specimens. Moreover, some of them also need to be re-described, considering that previous published descriptions were inaccurate or incomplete (i.e. Rhithropanopeus harrisii, Dyspanopeus sayi, Percnon gibbesi). Nine megalopae unknown until now (Sirpus zariquieyi, Parthenope angulifrons, Polybius zariquieyi, Ergasticus clouei, Afropinnotheres monodi, between others) will be described in detail. These new descriptions, together with the formerly existing ones, will allow the creation of an illustrated identification key, as a final objective, addressed to researchers and technicians, which is intended to assist in the correct identification of this important group of planktonic organisms, This study was funded by the Spanish “Ministerio de Ciencia e Innovación, Plan Nacional I+D” and FEDER through project MEGALOPADN (CGL2009–11225)

XVII Simposio Ibérico de Estudios de Biología Marina (SIEBM), 11-14 September 2012, Donostia-San Sebastián, La correcta identificación de las larvas planctónicas de los malacostráceos decápodos es fundamental para la realización de estudios sobre la biodiversidad de este importante grupo de artrópodos marinos, así como para la comprensión de procesos relacionados con la dinámica de poblaciones, periodos de reclutamiento, mecanismos de dispersión o funcionamiento de las redes tróficas. Sin embargo, existen grandes diferencias entre la morfología, el comportamiento y el hábitat de larvas y adultos, lo que supone un inconveniente a la hora de identificar los estados larvarios colectados en el plancton, ya que a menudo falta información morfológica que relacione los distintos estadios larvarios planctónicos con la especie a la que pertenecen. De las 140 especies de braquiuros ibéricos conocidas sólo se dispone de descripciones fiables de sus estadios larvarios, basadas principalmente en cultivos realizados en laboratorio, para el 41 % de las especies, siendo menos frecuente (65 especies) la descripción del último estadio larvario, denominado megalopa. La megalopa representa la transición entre las fases planctónica y bentónica, es por tanto una fase larvaria muy importante en el ciclo de vida de los braquiuros. La finalidad del presente estudio es la identificación de megalopas obtenidas del plancton mediante su código de barras genético, en concreto secuencias parciales de ADN mitocondrial de dos marcadores, los genes 16S y Cox1. Hasta el momento se han obtenido las secuencias del 65% de las especies de braquiuros de la Peninsula Ibérica lo que ha permitido la identificación de las megalopas de 26 especies, de las que de 7 se desconocía su morfología (Afropinnotheres monodi, Derilambrus angulifrons, Ergasticus clouei, Macropodia czernjwaskii, Polybius vernalis, Polybius zariquieyi y Sirpus zariquieyi) y 19 fueron previamente descritas en base a cultivos en laboratorio. De las primeras se han realizado las descripciones detalladas de su morfología, y del resto se han comparado con las descripciones previas. Las megalopas de Rhithropanopeus harrisii, Dyspanopeus sayi y Percnon gibbesi se han redescrito a la vista de lo incompleto, en base a criterios actuales, de las descripciones existentes. Hasta la fecha sólo existen 2 claves para la identificación de megalopas, una para especies atlánticas (Ingle, 1992) y otra para especies mediterráneas (Pessani et al., 2004). Entre ambas se pueden identificar las megalopas de 57 especies de la Península Ibérica. Las nuevas descripciones y redescripciones obtenidas en el presente proyecto, junto a las ya existentes, nos permiten en el momento actual elaborar una clave ilustrada para la identificación rápida y precisa de 74 megalopas en base a caracteres morfológicos externos, Estudio financiado por el Ministerio de Ciencia e Innovación (MICIIN) y el Fondo Europeo de Desarrollo Regional (FEDER): Proyectos CGL2009-11225, CTM2005-00024, y beca FPI

I Congreso Ibérico de Sistemática Animal (CISA), 17-19 de enero de 2012, Madrid, La correcta identificación de las larvas planctónicas de los crustáceos decápodos es fundamental para la realización de estudios sobre la biodiversidad de este importante grupo de macroinvertebrados marinos, así como para la comprensión de procesos como su dinámica de poblaciones, periodos de reclutamiento, mecanismos de dispersión o funcionamiento de las redes tróficas. Sin embargo, existen grandes diferencias entre la morfología, el comportamiento y el hábitat ocupado por larvas y adultos, lo que supone un problema a la hora de identificar los estados larvarios colectados en el plancton, ya que no hay datos a priori que los relacionen con la especie a la que pertenecen. En el caso de los braquiuros ibéricos, de las 140 especies conocidas, sólo para el 58 % se dispone de descripciones fiables de sus estadios larvarios, basadas principalmente en cultivos realizados en laboratorio, siendo la descripción de estadios más avanzados, como la megalopa, menos frecuente (65 especies). La megalopa representa la transición entre las fases planctónica y bentónica, es por tanto, una fase larvaria muy importante en el ciclo de vida de los braquiuros. La finalidad del presente estudio es la identificación de megalopas obtenidas del plancton mediante su código de barras genético, en concreto secuencias parciales de ADN mitocondrial de dos marcadores, los genes 16S y Cox1. Hasta el momento se han identificado 19 especies, 7 de las que se desconocía su morfología y 12 previamente descritas a partir de cultivos en laboratorio. De las primeras se están realizando las descripciones detalladas de su morfología y del resto se analizan sus diferencias con las descripciones previas. Estas nuevas descripciones, junto a las ya existentes, permitirán la elaboración de una clave ilustrada para la identificación rápida y precisa de las megalopas en base a caracteres morfológicos externos

Viral Encephalopathy and Retinopathy (VER), is caused by a nodavirus included within the Betanodavirus genus of the Nodaviridae family. This disease affects more than 30 marine fish species worldwide and has been a major obstacle in the aquaculture industry; control of the disease is based on virus detection, essentially in carrier specimens. This study describes a real time PCR procedure for viral nervous necrosis virus detection from several organs of sea bass, Senegalese sole, and gilt-head sea bream, from fish displaying either clinical symptoms or asymptomatic cases. The sensitivity of this technique was about 106-fold higher than that of the conventional RT-PCR. The newly designed primers detected nodavirus isolates belonging to the RGNNV and SJNNV genotypes. © 2010 Blackwell Verlag, Berlin., This study has been supported by the Spanish-Tunisian cooperation project (A/8793/07) granted to Dr Borrego and Dr Hammami.