TERMS OF USE This pdf is provided by Magnolia Press for private/research use. Commercial sale or deposition in a public library or website is prohibited.

Zootaxa 3700 (1): 159–172 ISSN 1175-5326 (print edition) www.mapress.com/zootaxa/ Article ZOOTAXA Copyright © 2013 Magnolia Press ISSN 1175-5334 (online edition) http://dx.doi.org/10.11646/zootaxa.3700.1.6 http://zoobank.org/urn:lsid:zoobank.org:pub:DD15D14D-DAC2-4305-A21C-2C73F298255A Mantacaprella macaronensis, a new and species of (Crustacea: ) from Canary Islands and Cape Verde

MAITE VÁZQUEZ-LUIS1, JOSÉ M. GUERRA-GARCÍA2, SUSANA CARVALHO3 & LYDIA PNG- GONZALEZ4 1Intituto Español de Oceanografía (IEO), Centre Oceanogràfic de les Balears, Moll de Ponent s/n, 07015 Palma de Mallorca, Spain. E-mail: [email protected] 2Laboratorio de Biología Marina, Dpto. Zoología, Facultad de Biología, Universidad de Sevilla, Avda Reina Mercedes 6, 41012 Sevilla, Spain 3IPMA—Portuguese Institute for the Sea and Atmosphere, Av. 5 de Outubro s/n, 8700-305 Olhao, Portugal 4Department of Biology, Marine Sciences Faculty, Campus Tafira, Universidad de Las Palmas de Gran Canaria, 35017 Tafira, Las Palmas, Spain

Abstract

Mantacaprella macaronensis new genus, new species, is described based on specimens collected from Canary Islands and Cape Verde. Mantacaprella is close to the genera Pariambus, Pseudolirius, Propodalirius and Paracaprella, but can be distinguished by the combination of the following characteristics: pereopods 3, 4 and 5 two-articulate; pereopods 6 and 7 six-articulate; mandibular molar present and palp absent; male abdomen with a pair of well-developed appendages. The new species has been found living in Cymodocea nodosa meadows and Caulerpa prolifera beds from 8.8 to 14.6 m depth in Gran Canaria (Canary Islands), and in natural rocky and artificial habitats (shipwrecks) at 25 m in Sal Island (Cape Verde). Even though the new species is one of the dominant amphipods inhabiting meadows of Cymodocea nodosa in Gran Canaria and in Cape Verde, it had not been described so far. This reflects the lack of knowledge on Macaronesian invertebrates, such as amphipods, and the need of further taxonomical studies to better characterise the whole biodiversity of this region and to design adequate programmes of management and conservation.

Key words: Caprellid, Macaronesia, Cymodocea nodosa, Caulerpa prolifera, natural hard substrata, artificial hard substrata

Introduction

According to Spalding et al. (2007) Marine Ecoregions are areas of relatively homogeneous species composition, clearly distinct from adjacent systems. The species composition is likely to be determined by the predominance of a small number of habitats and/or a distinct suite of oceanographic or topographic features. The dominant biogeographic forcing agents defining the ecoregions vary from location to location, but may include isolation, upwelling, nutrient inputs, freshwater influx, temperature regimes, ice regimes, exposure, sediments, currents, and bathymetric or coastal complexity (Spalding et al., 2007). The Macaronesian region sensu lato includes the Azores, Madeira, Salvage Islands, Canary Islands and Cape Verde. The islands are separated from the European and African continents by waters of depth exceeding 2000 m (Bamber 2012). During the last decade, new amphipod species have been described from this region (Guerra- García 2004, Krapp Schickel and Takeuchi 2005, Bellan-Santini 2007, Rubal and Larsen 2013), probably due to the scarce knowledge about the region and the paucity of studies. Furthermore, caprellid fauna from sediments and seagrasses is still scarcely explored (González et al., 2008; Guerra-García et al., 2013). In fact, even in regions where amphipod fauna has been widely studied, such as the Mediterranean Sea, new caprellids are still being described. For instance, a new caprellid was recently found within a Posidonia oceanica meadow from Sardinia, Italy (Sturaro and Guerra-García 2012). Cape Verde was recognised as one of the 10 marine biodiversity hot-spots most threatened by species extinction (Roberts et al., 2002), encouraging the development of several studies with conservation purposes. As a

Accepted by G. S. Karaman: 17 Jul. 2013; published: 12 Aug. 2013 159 TERMS OF USE This pdf is provided by Magnolia Press for private/research use. Commercial sale or deposition in a public library or website is prohibited. part of a general project assessing the biodiversity associated with natural and artificial rocky bottoms in Sal Island, Cape Verde, in 2009, specimens of an unusual caprellid were collected. Two years later, during another project focused on epifaunal community of Cymodocea nodosa and Caulerpa prolifera meadows carried out in Canary Islands, more individuals of the same taxa were collected. Detailed examination revealed that these specimens belonged to a new genus, here described as Mantacaprella. Ecological information on the new taxa is also presented.

Material and methods

In the Cape Verde Islands, a study was conducted in March and June 2009 to compare the abundance and composition of epibenthic fauna in the Bay of Santa Maria, Sal Island, from natural and artificial areas at two different depths (10 and 25 m) (Fig. 1). At each depth, two areas were sampled, one natural and one artificial (3 replicates of 0.0625 m2 in two spatially separated sites within each area). Consequently, four areas were sampled, Tchukulassa (25 m) and Farol (10 m) in natural areas, and Kwarcit (25 m) and Santo Antão (10 m) in artificial habitats. The sampled area was scraped and the material was collected with the help of a custom made small airlift with bags of 500 µm mesh size.

FIGURE 1. Map of the study area showing the sampling locations in Canary Islands and the Cape Verde. 1: Arinaga (type locality), 2: Risco Verde, 3: Cabrón, 4: Gando 1, 5: Gando 2, 6: Veneguera, 7: Tchukulassa, 8: Farol, 9: Kwarcit and 10: Santo Antão.

160 · Zootaxa 3700 (1) © 2013 Magnolia Press VÁZQUEZ-LUIS ET AL. TERMS OF USE This pdf is provided by Magnolia Press for private/research use. Commercial sale or deposition in a public library or website is prohibited.

In Gran Canaria, Canary Islands, sampling was carried out in November 2011 and October 2012 to characterise the epifaunal community within meadows of the seagrass Cymodocea nodosa and macroalgal beds dominated by the seaweed Caulerpa prolifera (between 8.8 and 14.6 m depth). In both sampling occasions, two areas were sampled in C. nodosa (Arinaga and Veneguera in 2011; and Risco Verde and Cabrón in 2012), and two in C. prolifera-dominated beds (Gando 1 and Gando 2, both in 2011 and 2012). At each habitat and locality 10 replicate samples of 0.04 m2 were taken. Removals were performed by cutting the vegetation immediately above the sediment surface, keeping the associated epifauna in unbleached woven cotton bags (Brearley et al., 2008). Samples were frozen (-20 ºC) until being processed, sieved through a 500 mm mesh sieve and preserved in 70% ethanol. The individuals of the new species were identified and counted, and their abundance was expressed as ind.m-2. The classification of Myers and Lowry (2003) was adopted, considering the family Caprellidae, and the subfamily Caprellinae. The specimens in better conditions were selected and dissected under a microscope Leica DM 1000. All dissected appendages were mounted in polyvinyl-lactophenol. The figures were drawn using a Leica compound microscope equipped with a camera lucida. The symbols used in the present work are: A1, 2 = Antenna 1, 2; UL = Upper lip; LL = Lower lip; LMd = Left mandible; RMd = Right mandible; Mx 1, 2 = Maxilla 1, 2; Mxp = Maxilliped; Gn 1, 2 = Gnathopod 1, 2; P5–7 = Pereopod 5–7; Ab = Abdomen. All the type material is deposited in the Museo Nacional de Ciencias Naturales de Madrid, Spain (MNCN).

Results

Systematics

Family Caprellidae Leach, 1814

Subfamily Caprellinae Leach, 1814

Mantacaprella n. gen.

Diagnosis. Flagellum of antenna 2 two-articulate. Gills present on pereonites 3 and 4. Pereopods 3, 4 and 5 two- articulate. Pereopods 6 and 7 six-articulate. Mandible molar present, palp absent. Male abdomen with a pair of well-developed appendages and a pair of lobes; female abdomen without appendages. Type species: Mantacaprella macaronensis n. sp. Gender: Feminine. Etymology. The term Mantacaprella is dedicated to the Manta Diving Center (www.mantadivingcenter.cv) in Cape Verde. The Manta Diving Center was responsible for the deployment of the first artificial reefs in Cape Verde waters within the framework of the Rebuilding Nature project. It was also by Mantas's initiative that a Portuguese research team started studying the assemblages of the natural and artificial reef assemblages of the Bay of Santa Maria (Sal Island, Cape Verde), where this species was originally identified. Therefore, we would like to highlight the role of all team members (skippers, divers, technical stuff and researchers) involved on this biodiversity project, by associating the Manta Diving Center with the name of the new genus.

Mantacaprella macaronensis n. sp (Figs. 2–7)

Type material: Holotype, mature male (used for drawings of lateral view, antennae, gnathopods, pereopods and abdomen) (vial and two slides, dissected) (vial: MNCN 20.04/9150, slide 1: MNCN 20.04/9151, and slide 2: MNCN 20.04/9152), Arinaga (N 27º51'52.9'', W 015º23'01.0''), Gran Canaria, Canary Islands, 10 meters deep, seagrass Cymodocea nodosa, November 2011. Paratype, mature female, collected together with the holotype (used for drawings of lateral view, gnathopod 2 and abdomen) (vial and one slide, not dissected) (vial: MNCN 20.04/9153, and slide: MNCN 20.04/9154).

A NEW CAPRELLID FROM MACARONESIAN REGION Zootaxa 3700 (1) © 2013 Magnolia Press · 161 TERMS OF USE This pdf is provided by Magnolia Press for private/research use. Commercial sale or deposition in a public library or website is prohibited.

Other paratypes (collected together with the holotype): Dissected and/or figured: 2 mature males dissected (2 vials and 2 slides) Male 1, vial: MNCN 20.04/9155, slide: MNCN 20.04/9156; Male 2, vial: MNCN 20.04/9157, slide: MNCN 20.04/9158—used for drawings of mouthparts—and 2 mature females dissected (2 vials and 2 slides). Female 1, vial: MNCN 20.04/9159, slide: MNCN 20.04/9160; Female 2, vial: MNCN 20.04/9161, slide: MNCN 20.04/9162; 3 juveniles (used for drawings of lateral view) MNCN 20.04/9163, MNCN 20.04/9164 and MNCN 20.04/9165. Not dissected, not figured: 5 mature males (MNCN 20.04/9166), 7 mature females (MNCN 20.04/9167). Additional material examined: Canary Islands: 10 mature males, 24 juvenile males, 25 mature females, 13 premature females, 20 juveniles, collected from Arinaga (type locality) together with the type material; 1 mature male, 1 mature female, 2 premature females collected from Veneguera, Gran Canaria, 11 m, seagrass Cymodocea nodosa, November 2011; 7 specimens collected from Gando 2, Gran Canaria, 14–15 m, Caulerpa prolifera dominated beds, November 2011 and October 2012; 9 specimens collected from Risco Verde, Gran Canaria, Canary Islands, 11 m, seagrass Cymodocea nodosa, November 2011; 17 specimens collected from Cabrón, Gran Canaria, 9 m, seagrass Cymodocea nodosa, October 2012. Cape Verde: 10 mature males, 7 juvenile males, 11 mature females, 3 premature females, 11 juveniles collected from Kwarcit, Santa Maria, Sal Island, 25 m, artificial substrate, March 2009; 1 mature male and 2 mature females collected from Tchukulassa, Santa Maria, Sal Island, 25 m, natural substrate, June 2009. Type locality. Arinaga (N 27º51'52.9'', W 015º23'01.0''), Gran Canaria, Canary Islands, 10 meters deep, seagrass Cymodocea nodosa, November 2011 (Fig. 1) Etymology. The specific name refers to the biogeographical area Macaronesia, which includes the Canary Islands, Azores, Madeira, Salvagen Islands and Cape Verde. Macaronesia derives from the greek makárôn nêsoi, which means cheerful or lucky islands. Description. Holotype male (vial: MNCN 20.04/9150, slide 1: MNCN 20.04/9151, and slide 2: MNCN 20.04/ 9152) (Figs. 2–4). Length: 4.5 mm Lateral view (Fig. 2): Body smooth dorsally. Pereonite 1 fused with head, suture present. Pereonites 2, 3 and 4 with an anterolateral and flattened projection at each side. Pereonites 3–5 subequal in length. Pereonite 7 the shortest. Gills (Fig. 2): Elongate, length about 3 times width. Antennae (Figs. 2, 3): Antenna 1 about 0.4 body length; peduncle setose, article 3 of the peduncle shorter than article 1 and 2; flagellum 8-articulate, article 1 being the result of two fused articles. Antenna 2 with short setae (no swimming setae); basal article of the peduncle with a distal projection; flagellum 2-articulate. Gnathopods (Fig. 3): Gnathopod 1 basis as long as ischium, merus and carpus combined; propodus length about 2 times width, palm with denticulate margin, with 1 grasping spine; dactylus provided with rows of setulae and distally bifid. Gnathopod 2 inserted on the anterior half of pereonite 2; basis a little longer than pereonite 2, smooth, lacking serrate carina, without distal projection; ischium rectangular; merus rounded; carpus short and triangular; propodus elongate, length about 3 times width; palm with a proximal projection carrying one grasping spine, a medial acute projection, followed by “U” notch and a distal triangular projection, distally denticulate; dactylus elongate, with a round projection medially, minutely denticulate. Pereopods (Figs. 2, 4): Pereopods 3 and 4 subequal, 2-articulate, length about 1/5 of gills, proximal article the longest, distal article small provided with 2–3 setae. Pereopod 5 with 2 articles, slightly larger than pereopods 3 and 4, proximal article the longest provided with 3 setae, distal article small provided with 3 setae. Pereopod 6 and 7 with 6 articles; pereopod 7 larger and more robust than pereopod 6, propodus proximally with knobs decreasing in size from proximal to distal end, with strong setae proximally. Penes (Fig. 4) rounded, situated laterally Abdomen (Fig. 4) with a pair of appendages 1-articulate and distally serrated, a pair of lateral lobes and a single dorsal lobe, with two plumose setae. Mouthparts (Fig. 5) figured from the male paratype; (Male 2, vial: MNCN 20.04/9157, slide: MNCN 20.04/ 9158). Mandibles without palp; mandibular molar robust; left mandible with incisor and lacinia mobilis 5-toothed followed by two plates; incisor of right mandible 5-toothed, lacinia mobilis transformed in a plate followed by two more plates; molar flake apparently absent. Upper lip smooth. Lower lip smooth, with round and well-demarcated inner lobes. Maxilla 1 outer lobe with 6 robust setae; palp with 4 distal setae and 1 lateral seta. Maxilla 2 inner lobe

162 · Zootaxa 3700 (1) © 2013 Magnolia Press VÁZQUEZ-LUIS ET AL. TERMS OF USE This pdf is provided by Magnolia Press for private/research use. Commercial sale or deposition in a public library or website is prohibited. triangular; outer lobe slightly larger than inner lobe, rectangular. Maxilliped inner plate small and rectangular with two setae; outer plate about 2.5 times as large as inner plate, with 5 setae; palp 4-articulate, article 3 of the palp with a distal projection, article 4 provided with two seta and tiny branch of setulae distally.

FIGURE 2. Mantacaprella macaronensis n. sp. Lateral view of holotype male (4.5 mm) and paratype female (2.7 mm). Scale bar: 1 mm.

A NEW CAPRELLID FROM MACARONESIAN REGION Zootaxa 3700 (1) © 2013 Magnolia Press · 163 TERMS OF USE This pdf is provided by Magnolia Press for private/research use. Commercial sale or deposition in a public library or website is prohibited.

FIGURE 3. Mantacaprella macaronensis n. sp. Holotype male antennae, gnathopod 1 and 2. Paratype female gnathopod 2. Scale bars: 0.2 mm. A1=Antenna 1; A2=Antenna 2; Gn1=Gnathopod 1; Gn2= Gnathopod 2.

164 · Zootaxa 3700 (1) © 2013 Magnolia Press VÁZQUEZ-LUIS ET AL. TERMS OF USE This pdf is provided by Magnolia Press for private/research use. Commercial sale or deposition in a public library or website is prohibited.

FIGURE 4. Mantacaprella macaronensis n. sp. Holotype male pereopods 3-7 and abdomen. Paratype female abdomen. Scale bars: 0.1 mm. P3-7=Pereopods 3-7; Ab=Abdomen.

Paratype female (vial: MNCN 20.04/9153, and slide: MNCN 20.04/9154) (Figs. 2, 3, 4) Length: 2.7 mm Similar to the male holotype apart from the following characters: pereonites 3–5 lacking the developed anterolateral projections; flagellum of antenna 1 with 6 articles; pereonite 1 after the suture less elongate than in male; gnathopod 2 propodus more rounded than in male, with a projection proximally with one grasping spines, two smaller projection in the palm and denticulate distally; oostegites on pereonite 3 and 4 with setae; genital openings in the pereonite 5 well-developed; abdomen without appendages, with two lateral lobes and a dorsal lobe.

A NEW CAPRELLID FROM MACARONESIAN REGION Zootaxa 3700 (1) © 2013 Magnolia Press · 165 TERMS OF USE This pdf is provided by Magnolia Press for private/research use. Commercial sale or deposition in a public library or website is prohibited.

FIGURE 5. Mantacaprella macaronensis n. sp. Mouthparts of paratype male. Scale bars: 0.05 mm. LL=Lower lip; UL=Upper lip; Mx1,2=Maxilla 1,2; LMd=Left mandible; RMd=Right mandible; Mxp=Maxilliped.

Intraspecific variation. The morphological characteristics of the species are rather constant in the specimens examined. Mouthparts were similar in the four specimens dissected. The flagellum of antenna 2 was always provided with two articles; the number of articles of antenna 1 flagellum was increasing from 4 (in the smaller juveniles examined) to 8 in the holotype mature male. Insertion of gnathopod 2 was always near proximal end of pereonite 2 regardless of the maturation stage of the specimen (Fig. 6). Length of pereonites 1 and 2 varied

166 · Zootaxa 3700 (1) © 2013 Magnolia Press VÁZQUEZ-LUIS ET AL. TERMS OF USE This pdf is provided by Magnolia Press for private/research use. Commercial sale or deposition in a public library or website is prohibited. according to the stage of development; the distance from the suture to pereonite 2 (pereonite 1 of the cephalon) was increasing from juvenile stages to adult. The smaller juveniles were lacking the anterolateral projections, which develop initially in pereonites 2 and 3 (Fig. 6), and lately in pereonite 4 (Fig. 2). Some juveniles have the pereonite 3 and 4 ventrally expanded (Fig. 6), but in mature males this widening of the body disappear and the pereonites become slender and provided by robust anterolateral and flattened projections (Fig. 2). The morphological variation of male gnathopod 2 is also represented in Fig. 6. In juveniles the propodus palm lacks projections. In juvenile males, a proximal projection with the grasping spine starts to develop together with a medial projection and “U” notch. The notch is more developed in adult males and the distal triangular projection becomes more evident and denticulate. The medial rounded projection in the dactylus is absent in juveniles and juvenile males. The material collected from Canary Islands is very similar morphologically to the material from Cape Verde and no morphological constant differences could be detected among populations of the two archipelagos. Ecological remarks. The specimens of Mantacaprella macaronensis were found in Gran Canaria in both types of habitats sampled. It was especially abundant on C. nodosa meadows in Arinaga (10 m depth), showing a mean abundance of 452.5±50.2 ind.m-2, being absent in the locality of Gando 1 (Fig. 7). In Cape Verde, the species was present both in natural rocky and artificial habitats around 25 meters depth, being absent in the areas sampled at 10 m depth (Fig. 7). The maximum mean abundance in Sal Island was found in Kwarcit with an average of 80.0±56.2 ind.m-2. In Gran Canaria, the amphipod fauna associated with C. nodosa meadows was dominated by Mantacaprella macaronensis, Pseudoprotella phasma and Ampithoe ramondi; while Microdeutopus stationis, Dexamine spinosa, Aora spinicornis, Ischyrocerus inexpectatus and Apherusa bispinosa were more abundant in C. prolifera- dominated beds. In Sal Island, amphipods were among the most abundant faunal groups, representing, together with polychaetes and sipunculids 80.8% of total abundance. The most abundant amphipod species was Stenothoe sp., however, these organisms were only observed in the artificial reef (Kwarcit). Phtisica marina, Ericthonius sp. and Jassa sp. were also highly abundant and present in both natural and artificial areas.

FIGURE 6. Mantacaprella macaronensis n. sp. Juvenile stages showing the development of gnathopod 2 and anterolateral projections. Scale bar: 1 mm.

A NEW CAPRELLID FROM MACARONESIAN REGION Zootaxa 3700 (1) © 2013 Magnolia Press · 167 TERMS OF USE This pdf is provided by Magnolia Press for private/research use. Commercial sale or deposition in a public library or website is prohibited.

FIGURE 7. Mean abundance (number of individuals.m-2 ± SE) of the Mantacaprella macaronensis n. sp. on the different studied localities in Gran Canaria and Sal Island. Ven= Veneguera, Ar= Arinaga, RV= Risco Verde, Cab= Cabrón, G1= Gando 1, G2= Gando 2, Tch= Tchukulassa, Far= Farol, Kw= Kwarcit and AS= Santo Antão.

Discussion

The genus Mantacaprella is close to the genera Paracaprella, Propodalirius, Pariambus and Pseudolirius, being distinguished by the combination of the following characteristics: pereopods 3, 4 and 5 two-articulate; pereopods 6 and 7 six-articulate; mandibular molar present and palp absent; male abdomen with a pair of well-developed appendages. After the examination of the specimens, the first idea was to include the material within the genus Paracaprella based on the similarities in antenna 1 (with a short article 3 of the peduncle), propodus of pereopods 6 and 7 with proximal knobs, mandibles lacking palp (as in some species of Paracaprella) and male abdomen with a pair of well-developed projections. However, the specimens clearly differ from this genus in the morphology of pereopods 5, which is 6-articulate in Paracaprella (McCain, 1968; Ros et al., 2013), but very reduced (only 2- articulate) in our material. The genus Propodalirius was also discarded since its pereopod 5 is 4-articulate (with the suture between the 2 proximal articles indistinct), and the morphology of gnathopod 1, maxilliped and antennae are clearly different (Mayer, 1903; Laubitz, 1991). The two remaining genera, Pariambus and Pseudolirius are very close, both having 2-articulate pereopod 5, as our specimens. The only significant morphological difference between these two genera is the structure of the abdomen (Mayer, 1890, 1903; Krapp-Schickel, 1993). Pseudolirius have 2 pairs of appendages: 1 pair with two articles (one developed basal article and other reduced to a strong setae), and the other pair rudimentary, with only one article bearing short setae. On the other hand, Pariambus is characterised by only one pair of rudimentary appendages (one-articulate), and by the presence of circular median plaque with some setae (Krapp-Schickel, 1993). The abdomen of our specimens has only one pair of appendages,

168 · Zootaxa 3700 (1) © 2013 Magnolia Press VÁZQUEZ-LUIS ET AL. TERMS OF USE This pdf is provided by Magnolia Press for private/research use. Commercial sale or deposition in a public library or website is prohibited. as Pariambus, but these appendages are well-developed, and the median plaque is lacking. Thus, our specimens would be closer to Pariambus than to other genera. The main morphological difference between Pariambus and our material, besides the abdomen structure, is the number of articles of pereopods 3 and 4, being tiny and one- articulate in Pariambus, and larger and two-articulate in our specimens. Moreover, the morphology of antenna 1 (with a very short article 3 of peduncle in our individuals) and pereopods 6 and 7 (with propodus provided with proximal knobs) is clearly different from those of Pariambus. For all these reasons, the erection of the new genus Mantacaprella is justified.

FIGURE 8. Lateral view of male and female of var. armata from Ceuta, North Africa. Scale bar: 1 mm.

During the revision of Mayer’s literature, we found figures of some forms of P. typicus gnathopods 2 (in mature males), which morphologically resemble gnathopods of Mantacaprella macaronensis (Mayer, 1882: page 75, figure 30; Mayer 1903: plate X, figure 7). Therefore, in order to clarify distinct morphological differences preventing further misidentifications, the taxonomical situation of the genus Pariambus is herein discussed. The genus Pariambus was firstly described as Podalirius by Kroyer (1844) with the type species Podalirius typicus.

A NEW CAPRELLID FROM MACARONESIAN REGION Zootaxa 3700 (1) © 2013 Magnolia Press · 169 TERMS OF USE This pdf is provided by Magnolia Press for private/research use. Commercial sale or deposition in a public library or website is prohibited.

Haller (1879) described a second species in the genus, Podalirius kroyeri, and Mayer (1882) completed the descriptions of these two species and described a third one, Podalirius minutus. Mayer (1890) transferred the species Podalirius kroyeri to the genus Pseudolirius and considered Podalirius minutus as a juvenile form of Podalirius typicus (Krapp-Schickel, 1993). Sars (1895) reported that Stebbing (1888) had changed the genus Podalirius to Pariambus, since the name Podalirius was already appropriated in Zoology (see also Lowry et al., 2013). However, Mayer (1903) still used the term Podalirius for this genus, and taking into account that the males of P. typicus were polymorphic, he described three forms: (1) α or armata, (2) β or inermis and (3) γ or cumana. The form armata is the most common variety, widely distributed in the Atlantic Ocean from Scotland to Cape Verde, and Algeria (Mediterranean Sea). The form inermis is distributed along British Isles, English Channel and French Atlantic coasts. The form cumana has been only recorded from Cuma (Italy). Further details on localities distribution are included in Mayer (1903) and McCain & Steinberg (1970). The form armata is characterised by the presence of grasping spines on the propodus of pereopods 6 and 7, together with the presence of a proximal tubercle on the third article of antenna 1 peduncle; the form inermis lack grasping spines and tubercle in antenna 1; and the form cumana has grasping spines but the antenna 1 tubercle is lacking. The main morphological differences among forms are summarized in Mayer (1903), Chevreux (1925) and Krapp-Shickel (1993). Geographically, P. typicus f. armata overlaps in its distribution with the new species. In fact, the presence of P. typicus has been confirmed from Canary Islands (Riera et al. 2012) and Madeira (Ramos, pers. com.). To provide morphological evidence for a proper identification, we have included a lateral view of Pariambus typicus f. armata collected from the north of Africa (Fig. 8). A detailed comparison between males of Fig. 2 (Mantacaprella macaronensis) and Fig. 8 (Pariambus typicus f. armata) reveal clear differences in the lateral view, which prevent misidentifications: (1) the body is robust in Pariambus typicus f. armata and slender in M. macaronensis (2) the head is rounded and smooth in M. macaronensis, while straight and provided with two small anterodorsal tubercles in P. typicus f. armata, (3) the article 3 of the peduncle is clearly shorter in M. macaronensis, and it is provided by a row of setae and a tubercle in P. typicus f. armata, (4) pereonites 2–4 have anterolateral projections in M. macaronensis, which are absent in P. typicus f. armata, (5) pereopods 3 and 4 are two-articulate in M. macaronensis, and tiny (one-articulate) in P. typicus f. armata, (6) pereopod 5 is inserted in the half of the pereonite 5 in P. typicus f. armata, and in the distal end of the pereonite in M. macaronensis, (7) in M. macaronensis the proximal article of pereonite 5 is larger and more robust than the distal one, while in P. typicus f. armata the proximal article is small and thin and the distal is larger, (8) the pereopods are robust and the propodus is provided with proximal knobs in M. macaronensis, while they are slender and without knobs in P. typicus f. armata. Most of these differences are also valid to distinguish between females of both species. It is important to point out that Pariambus is still waiting for a full morphological and molecular revision to clarify the status of the three forms (armata, inermis and cumana) of this species reported in the literature. For instance, recent molecular studies have demonstrated that forms or varieties of the complex of C. penantis belong to a different species (Cabezas et al., 2013) and detailed morphological studies have also revealed a new appraisal of species ranking for Caprella scauroides inside the complex of Caprella californica sensu lato (Takeuchi & Oyamada, 2013). The new species Mantacaprella macaronensis has been found in shallow waters with relatively high abundances in Canary Islands and Cape Verde, reflecting the lack of detailed studies on benthic fauna in the region, namely on amphipods. Therefore, this new species is expected to be present also in other islands of the Macaronesian region. On the other hand, during the identification of Cape Verde organisms, several specimens of the groups Polychaeta and Amphipoda could not been accurately assigned to an already described species, suggesting that other species new to science might had been collected. Further taxonomical studies are necessary to characterise the biodiversity and distribution of species in this region. This is necessary to design adequate programmes of management and conservation.

Acknowledgements

Samples of Canary Islands were collected within the framework of the European project “Changes in submerged vegetation: assessing how ecosystems services shift from frondose to depauperate systems dominated by opportunistic seaweeds (ECOSERVEG)” conducted by F. Tuya. Additional financial support was provided by the

170 · Zootaxa 3700 (1) © 2013 Magnolia Press VÁZQUEZ-LUIS ET AL. TERMS OF USE This pdf is provided by Magnolia Press for private/research use. Commercial sale or deposition in a public library or website is prohibited.

Spanish Ministerio de Economía y Competitividad (Project CGL2011-22474) cofinanced by FEDER funds. Samples of Cape Verde were collected within the scope of the project Rebuilding Nature - “Criac?ão de recifes artificiais em Cabo Verde” (“Establishment of artificial reefs in Cape Verde”), promoted by Manta Diving Center and sponsored by Soltrópico, Banco Comercial Atla?ntico, Grupo Oásis Atla?ntico, CV Telecom and Supersub- Beuchat. We would like to acknowledge the Manta Diving Center scientific diving team (Nuno Marques da Slva, Miguel Neves dos Santos, Miguel Tiago Oliveira, João Cúrdia, Pedro Lino, João Góis and Rute Portugal) for collecting the samples.

References

Bamber, R.N. (2012) Littoral Tanaidacea (Crustacea: Peracarida) from Macaronesia: allopatry and provenance in recent habitats. Journal of the Marine Biological Association of the United Kingdom, 92 (5), 1095–1116. http://dx.doi.org/10.1017/s0025315412000252 Bellan-Santini, D. (2007) New amphipods of hydrothermal vent environments on the Mid-Atlantic Ridge, Azores Triple junction zone. Journal of Natural History, 41 (9–12), 567–596 pp. Brearley, A., Kendrick, A.J. & Walker, D. (2008) How does burrowing by the isopod Limnoria agrostisa (Crustacea: Limnoriidae) affect the leaf canopy of the southern Australian seagrass Amphibolis griffithii? Marine Biology, 156, 65–77. http://dx.doi.org/10.1007/s00227-008-1065-1 Cabezas, M.P., Cabezas, P., Machordom, A. & Guerra-García, J.M. (2013) Hidden diversity and cryptic speciation refute cosmopolitan distribution in Caprella penantis (Crustacea: Amphipoda: Caprellidae). Journal of Zoological Systematics and Evolutionary Research, 51 (2), 85–99. http://dx.doi.org/10.1111/jzs.12010 Chevreaeux, E. & Fage, L. (1925) Amphipodes. Faune de France, 9, 1–488. Emerson, B.C. (2002) Evolution on oceanic islands: molecular phylogenetic approaches to understanding pattern and process. Molecular Ecology, 11, 951–966. http://dx.doi.org/10.1046/j.1365-294x.2002.01507.x González, A.R., Guerra-García, J.M., Maestre, M.J., Ruiz-Tabares, A., Espinosa, F., Gordillo, I., Sánchez-Moyano, J.E. & García-Gómez, J.C. (2008) Community structure of caprellids (Crustacea: Amphipoda: Caprellidae) on seagrasses from southern Spain. Helgoland Marine Research, 62, 189–199. http://dx.doi.org/10.1007/s10152-008-0107-x Guerra-García, J.M. (2004) Deep-sea Caprellidea (Crustacea, Amphipoda) from Azores with the description of three new species. Zoosystema, 26 (2), 235–262. Guerra-García, J.M., Navarro-Barranco, C., Corzo, J., Cobos-Muñoz, V., García-Adiego, E.M., Sempere Giménez, F. & García-Gómez, J.C. (2013) An illustrated key to the soft-bottom caprellids (Crustacea: Amphipoda) of the Iberian Peninsula and remarks to their ecological distribution along the Andalusian coast. Helgoland Marine Research, 67, 321– 336. http://dx.doi.org/10.1007/s10152-012-0324-1 Haller, G., (1879) Vorläufige Notizen über die Systematik der im Mittelmeer vorkommenden Caprelliden. Zoologischer Anzeiger, 2, 230–233. Krapp-Schickel, T. (1993). Suborder Caprellidea. In: Ruffo, S. (Ed.), The Amphipoda of the Mediterranean. Memoires de l'Institut Oceanographique (Monaco), vol. 13, pp. 773–809. Krapp-Schickel, T. & Takeuchi, I. (2005) A new species of Caprella (Amphipoda: Caprellidae) from the Cape Verde Islands (Atlantic), with note on Mediterranean Caprella liparotensis. Journal of Marine Biological Association of the UK, 85, 93– 100. http://dx.doi.org/10.1017/s002531540501088xh Kroyer, H. (1844–1845) Karcinologiske bidrag. Naturhistorisk tidsskrift, (N.R.) l, 283–345, pl. 2–3 ; p. 403; pp. 453–638, p1. 6–7. Laubitz, D.R. (1991) Crustacea Amphipoda Caprellidea: Caprellids from the western Pacific (New Caledonia, Indonesia and the Philippines). Resultats des campagnes Musorstom, 9, 101–123. Lowry, J., Costello, M. & Bellan-Santini, D. (2013) Pariambus Stebbing, 1888. In: Lowry, J. (2013) World Amphipoda database. Accessed through: World Register of Marine Species at http://www.marinespecies.org/aphia.php?p=taxdetails&id=101433 (Accessed 5 May 2013) Mayer, P. (1882) Monographie: Caprelliden. Fauna und flora des golfes von Neapel und der angrerzenden Meeres-Abschnitte, 6, i–x,1–201, pls I–X. http://dx.doi.org/10.5962/bhl.title.53624 Mayer, P. (1890) Die Caprelliden des Golfes von Neapel und der angrenzenden Meeres-Abschnitte. Nachtrag zur Monographie derselben. Fauna Flora Golf. Neapel, 17, i–viii+1–157. http://dx.doi.org/10.5962/bhl.title.53624

A NEW CAPRELLID FROM MACARONESIAN REGION Zootaxa 3700 (1) © 2013 Magnolia Press · 171 TERMS OF USE This pdf is provided by Magnolia Press for private/research use. Commercial sale or deposition in a public library or website is prohibited.

Mayer, P. (1890) Monographic: nachtrag zur den caprelliden. Fauna und flora des golfes von neapel und der angrerzenden Meeres-Abschnitte, 17, i–vii, 1–157, pls I–VII. http://dx.doi.org/10.5962/bhl.title.53624 Mayer, P. (1903) Die Caprellidae der Siboga-Expedition. Siboga-Expeditie, 34, 1–160. http://dx.doi.org/10.5962/bhl.title.53742 McCain, J.C. (1968) The Caprellidae (Crustacea: Amphipoda) of the western North Atlantic. United States National Museum Bulletin, 278, i–iv, 1–147. http://dx.doi.org/10.5479/si.03629236.278 McCain, J.C. & Steinberg, J.E. (1970) Amphipoda 1 Caprellidea 1. Fam. Caprellidae. Gruner HE, Holthuis LB (ed.) Catalogue, 2, 1–78. Myers, A.A. & Lowry, J.K. (2003) A phylogeny and a new classification of the Corophiidea Leach, 1814 (Amphipoda). Journal of Crustacean Biology, 23, 443–485. http://dx.doi.org/10.1651/0278-0372(2003)023[0443:apaanc]2.0.co;2 Riera, R., Delgado, J.D., Rodríguez, M., Monterroso, O. & Ramos, E. (2012) Macrofaunal communities of threatened subtidal maërl seabeds on Tenerife (Canary Islands, north-east Atlantic Ocean) in summer. Acta Oceanologica, 31 (1), 98–105. http://dx.doi.org/10.1007/s13131-012-0181-4 Roberts, C.M., McClean, C.J., Veron, J.E.N., Hawkins, J.P., Allen, G.R., McAllister, D.E., Mittermeier, C.G., Schueler, F.W., Spalding, M., Wells, F., Vynne, C. & Werner, T.B. (2002) Marine biodiversity hotspots and conservation priorities for tropical reefs. Science, 295 (5558), 1280. http://dx.doi.org/10.1126/science.1067728 Ros, M., Vázquez-Luis, M. & Guerra-García, J.M. (2013) The tropical caprellid amphipod Paracaprella pusilla: a new alien crustacean in the Mediterranean Sea. Helgoland Marine Research. http://dx.doi.org/10.1007/s10152-013-0353-4 Rubal, M. & Larsen, K. (2013) A new species of Ingolfiellidae (Peracarida, Amphipoda, Crustacea) from the Azores, Portugal. Helgoland Marine Research, 67, 149–154. http://dx.doi.org/10.1007/s10152-012-0311-6 Sars, G.O. (1895) Amphipoda. An account of the Crustacea of Norway. Christiania and Copenhagen. 1, 1–711, pls. 1–248. [Caprellidae : 644–667, 700–701, pls. 233–239] Spalding, M.D., Fox, H.E., Allen, G. R., Davidson, N., Ferdaña, Z.A., Finlayson, M., Halpern, B.S., Jorge, M.A., Lombana, A., Lourie, S.A., Martin, K.D., Mcmanus, E., Molnar, J., Recchia, C.A. & Robertson, J. (2007) Marine Ecoregions of the World: A Bioregionalization of Coastal and Shelf Areas. BioScience, 57 (7), 573–583. http://dx.doi.org/10.1641/b570707 Stebbing, T.R.R. (1888) Report on the Amphipoda collected by the H.M "Challenger" during the years 1873–76. Rep. Scient. Results "Challenger". 29 (1), xxxiv-872, (2), 873–1737. (3), i–xiii, pls. 1–210. [Caprellidae: 1226–1268, pls. 139–145. Sturaro, N. & Guerra-García, J.M. (2012) A new species of Caprella (Crustacea: Amphipoda) from the Mediterranean Sea. Helgoland Marine Research, 66, 33–42. http://dx.doi.org/10.1007/s10152-011-0244-5 Takeuchi, I. & Oyamada, A. (2013) Description of two species of Caprella (Crustacea: Amphipoda: Caprellidae) from the North Pacific; C. californica Stimpson, 1857 and C. scauroides Mayer, 1903, with a new appraisal of species ranking for C. scauroides. Helgoland Marine Research, 67 (2), 371–381. http://dx.doi.org/10.1007/s10152-012-0329-9

172 · Zootaxa 3700 (1) © 2013 Magnolia Press VÁZQUEZ-LUIS ET AL.