Dietary analysis of the marine Amphipoda (Crustacea: Peracarida) from the Iberian Peninsula

Depósito de investigación de la Universidad de Sevilla https://idus.us.es/ “This is an Accepted Manuscript of an article published by Elsevier in Journal of Sea Research on January 2014, available at: https://doi.org/10.1016/j.seares.2013.08.006 .” *Manuscript 61 62 63 64 65 8 1 Dietary analysis of the marine Amphipoda (Crustacea: Peracarida) 1 2 2 from the Iberian Peninsula 3 4 3 5 6 4 J.M. Guerra-García1,2 , J. M. Tierno de Figueroa 2,3 , C. Navarro-Barranco 1,2 , M. Ros 1,2, 7 5 J.E. Sánchez-Moyano1 9 10 and J. Moreira4 11 7 12 13 8 14 15 9 16 17 10 18 19 11 20 21 22 13 23 24 14 25 26 15 27 28 16 29 30 17 31 32 33 19 34 35 20 36 37 21 38 39 22 40 41 23 42 43 24 44 45 25 46 47 26 48 49 27 50 51 28 52 53 54 30 55 56 31 57 58 32 59 1 Departamento de Zoología, Facultad de Biología, Universidad de Sevilla, Avda Reina Mercedes 6, 41012 Sevilla, Spain 2 Jun Zoological Research Center, C/ Los Jazmines nº 15, 18213, Jun, Granada, Spain 3 Departamento de Zoología, Facultad de Ciencias, Universidad de Granada, Campus Fuentenueva, 18071, Granada, Spain 4 Departamento de Zoología, Facultad de Ciencias, Universidad Autónoma de Madrid, C/Darwin 2, 28049, Madrid, Spain Correspondence Laboratorio de Biología Marina, Departamento de Zoología, Facultad de Biología, Universidad de Sevilla, Avda. Reina Mercedes 6, 41012, Sevilla, Spain e-mail: [email protected] Phone: (+34)954556229 Fax: (+34) 954233480 ABSTRACT The gut contents of 2982 specimens of 33 amphipod families, 71 genera and 149 species were examined, representing a high percentage of amphipod diversity in the Iberian Peninsula. Material was collected from sediments, algae, seagrasses, hydroids and bryozoans along the whole coast of the Iberian Peninsula from 1989 to 2011. Although detritus was the dominant food item in the majority of amphipods, gammarideans also included carnivorous (mainly feeding on crustaceans) and herbivorous species (feeding on macroalgal tissues). Our study revealed that general assignment of a type of diet for a whole family is not always adequate. Some families showed a consistent pattern in most of the studied species (Corophiidae, Pontoporeiidae = detritivorous; Oedicerotidae, Phoxocephalidae, Stenothoidae = carnivorous; 6 12 18 29 34 Ampithoidae = primarily herbivorous on macroalgae), but others included species with 1 61 62 63 64 65 4 2 35 3 36 5 37 6 7 38 8 9 39 10 11 40 12 13 41 14 15 16 43 17 18 44 19 20 45 21 22 46 23 24 25 48 26 27 49 28 50 29 30 51 31 32 52 33 34 53 35 36 54 37 38 39 56 40 41 57 42 43 58 44 45 59 46 47 60 48 49 50 62 51 52 63 53 54 64 55 56 65 57 58 66 59 60 totally different feeding strategies. In general terms, detritivorous families were characterised by a stronger mandibular molar, while in carnivorous taxa this feature was less developed or reduced. The percentage of macroalgae in the digestive contents was associated in most cases with a reduction or loss of the mandibular palp. It seems that high trophic diversity in amphipods is a generalised trait along different ecosystems in all latitudes, and could be related to the ecological success of this group in marine benthic communities. Key words Feeding habits; Amphipods; Caprellideans; Gammarideans; Iberian Peninsula 1. Introduction Understanding the dietary habits of benthic invertebrates is pivotal to studies of food webs and energy flows in marine ecosystems, but basic information on the feeding ecology of most taxa is still lacking nowadays (Legeżyńska et al., 2012). Amphipods are the most diverse group of crustaceans with respect to life styles, trophic types, habitats and sizes (De Broyer and Jazdzewski, 1996) and constitute one of the dominant groups of marine rocky habitats and soft bottoms (de-la-Ossa- Carretero et al., 2011). Their high abundance, species richness and wide distribution suggest that amphipods indeed play major roles in the ecology of these habitats (Conlan, 1994). They are an important link between primary and secondary production and higher trophic levels such as fishes, birds and mammals (see Legeżyńska et al., 2012). In studies of marine food webs, amphipods are often grouped into a single trophic guild based on the assumption that they are functionally redundant, but this assumption has been challenged due to contrary evidence provided by feeding assays and gut content analysis (Farlin et al., 2010). As consumers, benthic amphipods are known to have versatile feeding strategies (Carrasco and Arcos, 1984; Sarvala and Uitto, 1991). Amphipods inhabit a variety of marine environments and, in consequence, they show a high diversity of feeding habits; they can feed on debris, detritus, bacteria, aquatic fungi, living animals, carrion and dead plant fragments (Conradi and Cervera, 1995). They also constitute an important food source for a large variety of marine predators (see Vázquez-Luis et al., 2013), hence playing a key role in energy flow through food 42 47 55 61 67 68 webs. Furthermore, amphipods have often been included in ecotoxicological tests and 1 61 62 63 64 65 4 2 69 3 70 5 71 6 7 72 8 9 73 10 11 74 12 13 75 14 76 15 16 77 17 18 78 19 20 79 21 22 80 23 24 25 82 26 27 83 28 29 84 proposed as good bioindicators of the quality of marine habitats (Conradi et al., 1997, Guerra-García and García-Gómez, 2001; Bat, 2005, de-la-Ossa-Carretero et al., 2012) and they can also be an interesting potential resource in aquaculture (Baeza-Rojano et al., 2010, 2012). In spite of the importance of amphipods in marine ecosystems, little is actually known about the feeding habits of these crustaceans, and the diversity of amphipod feeding behaviours in the wild remains an important gap in our knowledge of benthic ecosystems (Farlin et al., 2010). The ecofunctional and trophodynamic roles of epibenthic amphipods are still poorly known (Mancinelli and Rossi, 2002); moreover, trophic roles and functional types have been studied in fewer than about 10% of amphipod species, with very few quantitative approaches so far (Dauby et al., 2001). Traditionally, amphipod feeding preferences have been assessed using in situ and laboratory observations, feeding experiments, gut-content analysis and studies of the functional morphology of feeding appendages (Legeżyńska et al., 2012). Knowledge of amphipod feeding ecology has recently expanded thanks to the use of biomarkers such as lipids and fatty acids (FAs) and stable isotopes (Mancinelli, 2012). In fact, in marine 30 15 13 31 85 food web studies, stable isotopes of nitrogen (δ N) and carbon (δ 32 N) are widely used 33 86 34 35 36 88 37 38 89 39 40 90 41 42 91 43 44 92 45 46 47 94 48 49 95 50 51 96 52 53 97 54 55 98 56 57 58 100 59 60 101 81 87 93 99 68 webs. Furthermore, amphipods have often been included in ecotoxicological tests and 1 61 62 63 64 65 to esti mat e org anis ms’ trop hic leve ls and car bon sou rces , resp ecti vel y (Sø reid e and Ny går d, 201 2). The utili ty of these methods lies in the fact that, in contrast to gut content examination, which provides insight into short-term preferences, they provide dietary information integrated over periods of weeks to months (Legeżyńska et al., 2012). However, the isotope studies require fresh material preventing the use of specimens already fixed in ethanol or formalin, and also depend on the abundance of available material (especially if the species are small) to have enough amounts for chemical analysis. On the other hand, traditional studies dealing with the direct observation of digestive contents are very scarce, probably due to the difficult task of removing the digestive tract by dissection. Recently, a method used by entomologists for studying the digestive contents of insects (e.g. Tierno de Figueroa et al., 2006) was successfully essayed for caprellids (Guerra-García and Tierno de Figueroa, 2009). This study concluded that caprellids feed mainly on detritus based on the examination of 742 specimens of 31 genera and 62 species from all around the world. This pattern has recently been supported by C and N isotope analysis (Jeong et al., 2012). After this, several additional studies using this 102 methodology have been published to explore variations in the diet of tropical caprellids 1 61 62 63 64 65 (Alarcón-Ortega et al., 2012), to describe the diet of amphipods (caprellideans and gammarideans) from soft bottoms submarine caves (Navarro-Barranco et al., 2013) and to assess if the feeding habits of amphipods associated with coastal seaweeds could be affected by the spread of invasive algae (Vázquez-Luis et al., 2013). Although the knowledge of amphipod feeding habits has increased during the last few years, a comprehensive study of gammaridean trophic diversity is still lacking. There is information available for some species (Torrecilla-Roca et al., 2012; Vázquez-Luis et al., 2013), including a recent review on the studies based on stable isotope data (Mancinelli, 2012), but a comprehensive dietary analysis is necessary. In fact, several authors have pointed out that information on the species-specific trophic ecology of amphipods is needed to better understand their potential role in the trophic dynamics and carbon flow of marine ecosystems (Jeong et al., 2012). Consequently, the main objective of the present study is to characterise the diet of amphipods from the Iberian Peninsula, including both Mediterranean and Atlantic species. 2. Material and methods We examined 2982 specimens from 33 families, distributed in 71 genera and 149 species (Table 1), representing a high percentage of amphipod diversity in the Iberian Peninsula. Material was collected mainly from sediments, algae and hydroids along the whole coast of the Iberian Peninsula from 1989 to 2011. Although most sampling efforts were conducted in the Strait of Gibraltar area and the Galician coasts, material collected along the whole coasts of Spain and Portugal was included. The Iberian Peninsula is a very interesting geographical zone since it is the confluence of several biogeographic areas and is located between two different water masses: the Atlantic Ocean and the Mediterranean Sea. We have also included, especially for caprellids, samples collected from Ceuta, located in North Africa at the Strait of Gibraltar, and Alborán Island located in South-eastern Spain. All the material was collected between 0 and 40 meters deep by snorkelling, SCUBA diving and van Veen grabs or corers. The whole list of analyzed material is included in Appendix I. The number of specimens observed for each species depended on the availability of material, but we tried, whenever possible, to study at least 10-20 specimens of each species. Data of those species with a very low number of specimens examined should be 2 103 3 4 104 5 105 6 7 106 8 9 107 10 11 108 12 13 109 14 15 110 16 111 17 18 112 19 20 113 21 22 114 23 24 115 25 26 116 27 28 117 29 30 118 31 32 119 33 34 120 35 36 121 37 122 38 39 123 40 41 124 42 43 125 44 45 126 46 47 127 48 49 128 50 129 51 52 130 53 54 131 55 56 132 57 58 133 59 60 134 135 taken with caution. Families, genera and species’ names are in accordance with the 1 61 62 63 64 65 2 136 World Register of Marine Species – WoRMS 3 4 137 (http://www.marinespecies.org/aphia.php?p=search). 5 6 138 Animals were fixed in 4% formalin or 70% ethanol depending on the samples 7 8 139 and all material was preserved in 70% ethanol. For the diet study, individuals were 9 140 analyzed following the methodology proposed by Bello and Cabrera (1999) with slight 10 11 141 variations. This method has been used to study the gut contents of different arthropod 12 13 142 groups and other animals, both aquatic and terrestrial forms and both ethanol and 14 15 143 formalin preserved samples, revealing that it as a very appropriate method for gut 16 17 144 content analysis (e.g. Tierno de Figueroa et al., 2006; López-Rodríguez et al., 2009). 18 145 Recently, this method has been successfully used in amphipods (Guerra-García and 19 20 146 Tierno de Figueroa, 2009; Alarcón-Ortega et al., 2012, Torrecilla-Roca and Guerra- 21 22 147 García, 2012; Vázquez-Luis et al., 2013, Navarro-Barranco et al., 2013). Specimens of 23 24 148 each species were introduced in vials with Hertwig’s liquid (consisting of 270 g of 25 26 149 chloral hydrate, 19 ml of chloridric acid 1N, 150 ml of distilled water and 60 ml of 27 28 150 glycerin) and heated in an oven at 65 ºC for 2 to 24 hours depending on the cuticle 29 151 thickness of the specimens. After this, they were mounted on slides for study under the 30 31 152 microscope. The percentage of absolute gut content (at 40x or 100x), as the total area 32 33 153 occupied by the content in the whole digestive tract, and relative gut content (at 100x or 34 35 154 400x), as the area occupied for each component within the total gut content, were 36 37 155 estimated using a microscope equipped with an ocular micrometer. Mean and standard 38 39 156 error of the mean were calculated. 40 157 41 42 158 3. Results 43 44 159 Gut contents of the studied amphipod species included detritus, metazoan preys 45 46 160 (crustaceans, polychaetes, oligochaetes, kinorhynchs and hydroids), macroalgae, 47 48 161 microalgae (e.g. diatoms), dinoflagellates and foraminifers (Table 1). The dominant 49 50 162 component was detritus, followed by crustaceans (mainly copepods), which were the 51 163 dominant prey of carnivorous species, and macroalgal tissues, which were characteristic 52 53 164 of herbivorous amphipods. Dinoflagellates were found in caprellideans but not in 54 55 165 gammarideans, foraminifers were rarely represented and were found mainly in the 56 57 166 family Maeridae. There were oligochaetes and kinorhynchs occasionally. The average 58 59 167 area occupied by content in the whole digestive tract ranged from 6.0 % in Caprella 60 168 andreae to 93.6 % in Ampelisca spinipes. In general terms, detritivorous species had a 169 higher area occupied by content in the digestive tract than carnivorous species. 1 57 58 59 60 61 62 63 64 65 Although amphipods form a trophic continuum from primary herbivores to carnivores (Legeżyńska et al., 2012), this comprehensive work reveals that detritus is 201 202 4 24 35 46 2 170 3 171 5 172 6 7 173 8 9 174 10 11 175 12 13 176 14 177 15 16 178 17 18 179 19 20 180 21 22 181 23 182 25 183 26 27 184 28 29 185 30 31 186 32 33 187 34 188 36 189 37 38 190 39 40 191 41 42 192 43 44 193 45 194 47 195 48 49 196 50 51 197 52 53 198 54 55 199 56 200 Furthermore, empty guts predominated in carnivorous species. For example, we examined 49 individuals of Metaphoxus fultoni and 42 of Stenothoe tergestina, and only 5 and 15 specimens respectively had digestive contents. For some species, such as Amphilochus brunneus, Colomastix pusilla, Cressa cristata, C. mediterranea, Peltocoxa gibbosa, Jassa occia and Pereionotus testudo, all specimens examined had empty guts so diet for these amphipods could not be characterised. When diet was characterised at family level, detritus was also the main component of most of them, apart from Leucothoidae, Stenothoidae, Phoxocephalidae, Oedicerotidae, Liljeborgidae and Pontogeneiidae, in which prey was the dominant item (Fig. 1). Some important families showed a very consistent pattern for all species included (see Table 1), such as Corophiidae or Pontoporeiidae, which can be considered detritivorous families. The 7 species that were studied of the Oedicerotidae family, the 8 species of Phoxocephalidae and the 3 species of Stenothoidae were clearly carnivorous. Most members of the family Ampithoidae were herbivorous, so we could assign herbivory to this family. However, many families included species with different feeding habits, such as Amphilochidae, with carnivorous species (Amphilochus neapolitanus and A. spencebatei), detritivorous (Gitana sarsi) or omnivorous (A. picadurus). Although half of the studied species of the family Hyalidae were clearly herbivorous (Hyale perieri, H. pontica and H. stebbingi), H. schmidti was undoubtedly detritivorous, H. spinidactyla carnivorous and Apohyale prevostii omnivorous (Table 1). In general terms, the families feeding on detritus were characterised by a stronger mandibular molar, while in carnivorous species there seems to be a trend towards a reduction of this structure. The percentage of macroalgae in the digestive contents was associated in most cases to a reduction or loss of mandibular palp (Fig. 1). 4. Discussion 4.1. The importance of detritus in amphipod diet The trophic diversity of amphipod species measured in the present study indicates that they use different food resources within their microhabitats. This suggests that they play species-specific functional roles as mediators in trophic pathways from producers to higher-level consumers of the marine ecosystems they inhabit. 203 the main food item in the majority of species. The importance of detritus in benthic 1 57 58 59 60 61 62 63 64 65 235 trophic diversity, with a major contribution of algal material in Gammarella fucicola communities has often been reported in the literature. More energy and materials flow through detritus food webs than through grazer food webs in most freshwater, estuarine and marine ecosystems, and many animals can use detritus directly because it is highly nutritious after a short period of microbial colonization (Mann, 1998). The data obtained herein for amphipods of the Iberian Peninsula (gammarideans and caprellideans) support previous observations obtained for worldwide caprellids (Guerra- García and Tierno de Figueroa, 2009). The percentage of species feeding on metazoan prey was higher than that of species feeding on macroalgae. Animal matter is more easily assimilated than vegetal matter. In fact, marine plants are little used by animals that graze directly on them because they can have a relatively high content of indigestible fibers and low nitrogen content (Mann, 1998). Thus, our study shows that even in species with high contents of macroalgae in their digestive tracts, detritus was the main gut component (Fig. 1). Howard (1982) reported that the inclusion of epibenthic gammaridean amphipods in a laboratory microcosm containing eelgrass, substantially reduced the load of attached epiphytes and settled detritus on eelgrass leaves, when compared to a microcosm without amphipods. Duffy and Hay (2000), based on mesocosm experiments, reported that grazing amphipods, which are ubiquitous in marine vegetation but poorly understood ecologically, may have strong impacts and play important roles in the organization of benthic communities, particularly where predation pressure is low. In addition to their abundance and rates of consumption, the way in which mesograzers consume plants is of great importance for predicting their effects on plant fitness (Poore, 1994). Several experimental studies of mainly herbivorous amphipods have compared feeding impacts of amphipods on the plant (and invertebrate) communities of which they are a part and often found quite different impacts (Duffy, 1990; Duffy and Harvilicz, 2001; Best and Stachowicz, 2012). The method used in the present study let us clearly differentiate the presence of detritus from other types of items such as crustaceans, microalgae, macroalgae, etc. However, detritus feeders can display relatively different diets depending on the vegetal or animal origin of the organic matter. In this sense, isotopic values for amphipods associated with leaf litter of the seagrass Posidonia oceanica showed a certain degree of 2 204 3 4 205 5 206 6 7 207 8 9 208 10 11 209 12 13 210 14 15 211 16 212 17 18 213 19 20 214 21 22 215 23 24 216 25 26 217 27 218 28 29 219 30 31 220 32 33 221 34 35 222 36 37 223 38 39 224 40 225 41 42 226 43 44 227 45 46 228 47 48 229 49 50 230 51 231 52 53 232 54 55 233 56 234 438 Caine, E.A., 1974. Comparative functional morphology of feeding in three species of 1 46 47 48 49 50 51 52 53 54 55 56 57 58 59 60 61 62 63 64 65 rade de Villefranche-sur-Mer. Trav. Sta. Zool. Villefranche 19, 103-105. 464 465 466 Dauby, P., Scailteur, Y., De Broyer, C., 2001. Trophic diversity within the eastern Weddell Sea amphipod community. Hydrobiologia 443, 69-86. 467 468 469 Dauvin, J.C., 1984. Dynamique d’écosystèmes macrobenthiques des fonds sédimentaires de la baie de Morlaix et leur perturbation par les hydrocarbures de l’Amoco-Cádiz. PhD Thesis, Univ. P et M Curie, Paris, pp. 446. 470 471 caprellids (Crustacea: Amphipoda) from the North western Florida Gulf Coast. J. Exp. Mar. Biol. Ecol. 15, 81-96. Caine, E.A., 1977. Feeding mechanisms and possible resource partitioning of the Caprellidae (Crustacea: Amphipoda) from Puget Sound, USA. Mar. Biol. 42, 331-336. Carrasco, F.D., Arcos, D.F., 1984. Life history of a cold temperate population of the sublittoral amphipod Ampelisca araucana. Mar. Ecol. Prog. Ser. 14,: 245-252. Coleman, C.O., 1989. On the nutrition of two Antarctic Acanthonotozomatidae (Crustacea: Amphipoda). Gut contents and functional morphology of mouthparts. Polar Biol. 9, 287-294. Conlan, K.E., 1994. Amphipod crustaceans and environmental disturbance: a review. J. Nat. Hist. 28, 519-554. Conradi, M., Cervera, L., 1995. Variability in trophic dominance of amphipods associated with the bryozoan Bugula neritina (L., 1758) in Algeciras Bay (Southern Iberian Peninsula). Pol. Arch. Hydrobiol. 42, 483-494. 43 44 462 45 463 Conradi, M., López-González, P.J., García-Gómez, J.C., 1997. The Amphipod community as bioindicator in Algeciras Bay (Southern Iberian Peninsula) based on a spatio-temporal distribution. PSZN Mar. Ecol. 18, 97-111. Costa, S., 1960. Note préliminaire sur l’éthologie alimentaire de deux Caprellidés de la 2 439 3 4 440 5 6 441 7 8 442 9 443 10 11 444 12 13 445 14 15 446 16 17 447 18 448 19 20 449 21 22 450 23 24 451 25 26 452 27 28 453 29 454 30 31 455 32 33 456 34 35 457 36 37 458 38 459 39 40 460 41 42 461 472 1 46 47 48 49 50 51 52 53 54 55 56 57 58 59 60 61 62 63 64 65 498 Duffy, J.E., Harvilickz, A., 2001. Species-specific impacts of grazing amphipods in an eelgrass-bed community. Mar. Ecol. Prog. Ser. 223, 201-211. 499 500 501 Duffy, J.E., Hay, M.E., 2000. Strong impacts of grazing amphipods on the organization of a benthic community. Ecol. Mon. 2000, 237-263. 502 503 504 4 24 2 473 3 474 5 475 6 7 476 8 9 477 10 11 478 12 13 479 14 480 15 16 481 17 18 482 19 20 483 21 22 484 23 485 25 486 26 27 487 28 29 488 30 31 489 De Broyer, C. and Jazdzewski, K., 1996. Biodiversity of the Southern Ocean: towards a new synthesis for the Amphipoda (Crustacea). Boll. Mus. Civ. Stor. Nat. Verona 20, 547- 568. De Broyer, C., Costello, M., Bellan-Santini, D., 2012. Amphilochidae. In: Lowry J. World Amphipoda database. Accessed through: Wold Register of Marine Species at http://www.marinespecies.org/aphia.php?p=taxdetailsandid=101365 on 2012-11-16. de-la-Ossa-Carretero, J.A., Dauvin, J.C., Del-Pilar-Ruso, Y., Giménez-Casalduero, F. and Sánchez-Lizaso, J.L., 2011. Inventory of benthic amphipods from fine sand community of the Iberian Peninsula east coast (Spain), western Mediterranean, with new records. Mar. Biod. Rec. 3 e119, 1-10. de-la-Ossa-Carretero, J.A., Del-Pilar-Ruso, Y., Giménez-Casaldurero, F., Sánchez- Lizaso, J.L., Dauvin, J.C., 2012. Sensitivity of amphipods to seawage pollution. Est. Coast. Shelf Sci. 73, 630-638. Dekker, R., 1989. The macrozoobenthos of the subtidal western dutch wadden sea I. Biomass and species richness. Netherlands J. Sea Res. 23, 57-68. 43 44 496 45 497 Dixon, I.M.T., Moore, P.G., 1997. A comparative study on the tubes and feeding behaviour of eight species of corophioid Amphipoda and their bearing on phylogenetic relationships within the Corophioidea. Phil. Trans. R. Soc. Lond. 352, 93-112. Duffy, J.E., 1990. Amphipods on seaweeds: partners or pests ? Oecologia 83, 267-276. 32 33 490 34 35 491 36 37 492 38 493 39 40 494 41 42 495 505 Eleftheriou, A, Basford D.J., 1989. The macrobenthic infauna of the offshore northern 1 46 47 48 49 50 51 52 53 54 55 56 57 58 59 60 61 62 63 64 65 359. 531 532 533 Howard, R.K., 1982. Impact of feeding activities of epibenthic amphipods on surfacefouling of eelgrass leaves. Aquatic Bot. 14, 91-97. 534 535 536 Jeong, S.J., Suh, H-L., Kang, C-K., 2012. Trophic diversity in amphipods within a temperate eelgrass ecosystem as determined by gut contents and C and N isotope analysis. Mar. Biol. 159, 1943-1954. 537 538 4 24 35 2 506 3 507 5 508 6 7 509 8 9 510 10 11 511 12 13 512 14 513 15 16 514 17 18 515 19 20 516 21 22 517 23 518 25 519 26 27 520 28 29 521 30 31 522 32 33 523 34 524 36 525 37 38 526 39 40 527 41 42 528 43 44 529 45 530 North Sea. J. Mar. Biol. Ass. UK 69, 123-143. Enequist, P., 1949. Studies on the soft-bottom amphipods of the Skagerak. Zool. Bidrag. Fran. Uppsala 28, 297-492. Fanelli, E., Cartes, J.E., Badalamenti, F., Rumolo, P., Sprovieri, M., 2009. Trophodynamics of suprabenthic fauna on coastal muddy bottoms of the southern Tyrrhenian Sea (western Mediterranean). J. Sea Res. 61, 174-187. Farlin, J.P., Lewis, L.S., Anderson, T.W., Lai, C.T., 2010. Functional diversity in amphipods revealed by stable isotopes in an eelgrass ecosystem. Mar. Ecol. Prog. Ser. 420, 277-281. Greze, I.I., 1968. Feeding habits and food requirements of some amphipods in the Black Sea. Mar. Biol. 1, 316-321. Guerra-García, J.M., García-Gómez, J.C., 2001. The spatial distribution of Caprellidae (Crustacea: Amphipoda): a stress bioindicator in Ceuta (North Africa, Gibraltar Area). PSZN Mar. Ecol. 22, 357-367. Guerra-García, J.M., Tierno de Figueroa, J.M., 2009. What do caprellids feed on? Mar. Biol. 156, 1881-1890. Hily, C., 1984. Variabilité de la macrofaune benthique dans les milieux hypertrophiques de la rade de Brest. PhD Thesis, Université de Bretagne Occidentale, pp. 539 1 58 59 60 61 62 63 64 65 571 Sci. 114, 130-139. 572 4 24 35 46 57 2 540 3 541 5 542 6 7 543 8 9 544 10 11 545 12 13 546 14 547 15 16 548 17 18 549 19 20 550 21 22 551 23 552 25 553 26 27 554 28 29 555 30 31 556 32 33 557 34 558 36 559 37 38 560 39 40 561 41 42 562 43 44 563 45 564 47 565 48 49 566 50 51 567 52 53 568 54 55 569 56 570 Lastra, M., Palacio, J., Sánchez-Mata, A., Mora, J., 1991. Estructura trófica infralitoral de la Bahía de Santander. Cah. Biol. Mar. 32, 333-351. Legeżyńska, J., Kędra, M., Walkusz, W., 2012. When season does not matter: summer and winter trophic ecology of Arctic amphipods. Hydrobiologia 684, 189-214. Lepoint, G., Cox, A.S., Dauby, P., Poulicek, M., Gobert, S., 2006. Food sources of two detritivore amphipods associated with the seagrass Posidonia oceanica leaf litter. Mar. Biol. Res. 2, 355-365. Lewis, J.B., 1992. Abundance, distribution and behaviour of a commensal amphipod Stenothoe valida Dana on the hydrocoral Millepora complanata Lamarck. Bull. Mar. Sci. 51, 245-249. Lincoln, R.J., 1979. British Marine Amphipoda: Gammaridea. British Museum (Natural History): London, UK. 658 pp. López-Rodríguez, M.J., Tierno de Figueroa, J.M., Fenoglio, S., Bo, T., Alba-Tercedor, J., 2009. Life strategies of three Perlodidae species (Plecoptera, Insecta) in a Mediterranean seasonal stream of Southern Europe. J. North Am. Benth. Soc. 28, 611- 625. Lowry, J.K., Myers, A.A., 2013. A phylogeny and classification of the Senticaudata subord. nov. (Crustacea: Amphipoda). Zootaxa, 3610: 1-80. Macneil, C., Dick, J.T.A., Elwood, R.W., 1997. The trophic ecology of freshwater Gammarus spp. (Crustacea: Amphipoda): problems and perspectives concerning the functional feeding group concept. Biol. Rev. 72, 349-364. Mancinelli, G., 2012. On the trophic ecology of Gammaridea (Crustacea: Amphipoda) in coastal waters: a European-scale analysis of stable isotopes data. Est. Coast. Shelf 573 Mancinelli, G., Rossi, L., 2002. The influence of allochthonous leaf detritus on the 1 58 59 60 61 62 63 64 65 4 24 35 46 57 2 574 3 575 5 576 6 7 577 8 9 578 10 11 579 12 13 580 14 581 15 16 582 17 18 583 19 20 584 21 22 585 23 586 25 587 26 27 588 28 29 589 30 31 590 32 33 591 34 592 36 593 37 38 594 39 40 595 41 42 596 43 44 597 45 598 47 599 48 49 600 50 51 601 52 53 602 54 55 603 56 604 occurrence of crustacean detritivores in the soft-bottom macrobenthos of the Po River Delta Area (northwestern Adriatic Sea). Est. Coast. Shelf Sci. 54, 849-861. Mann, K.H., 1998. Production and use of detritus in various freshwater, estuarine and coastal marine ecosystems. Limnol. Oceanogr. 33, 910-930. Mayer, G., Maier, G., Maas, A., Waloszek, D., 2008. Mouthparts of the ponto-caspian invader Dikerogammarus villosus (Amphipoda: Pontogammaridae). J. Crust. Biol. 28, 1- 15. Minshall, G.W., 1988. Stream ecosystem theory: a global perspective. J. North Amer. Benthol. Soc. 7, 263-288. Myers, A.A., Lowry, J.K., 2003. A phylogeny and a new classification of the Corophiidea Leach, 1814 (Amphipoda). J. Crust. Biol. 23, 443-485. Navarro-Barranco, C., Tierno de Figueroa, J.M., Guerra-García, J.M., Sánchez Tocino, L., García-Gómez, J.C., 2013. Feeding habits of amphipods (Crustacea: Malacostraca) from shallow soft bottom communities; comparison between marine caves and open habitats. J. Sea Res. 78, 1-7. Oakden, J.M., 1984. Feeding and substrate preference in five species of phoxocephalid amphipods from central California. J. Crust. Biol. 4, 233-247. Oliver, J.S., Oakden, J.M., Slattery, P.N., 1982. Phoxocephalid amphipod crustaceans as predators on larvae and juveniles in marine soft-bottom communities. Mar. Ecol. Prog. Ser. 7, 179-184. Poore, G., 1994. Selective herbivory by amphipods inhabiting the brown alga Zonaria angustata. Mar. Ecol. Prog. Ser. 107, 113-123. 605 Procaccini, G.., Scipione, M.B., 1991. Observations on the spatial-temporal distribution 1 57 58 59 60 61 62 63 64 65 637 4 24 35 46 2 606 3 607 5 608 6 7 609 8 9 610 10 11 611 12 13 612 14 613 15 16 614 17 18 615 19 20 616 21 22 617 23 618 25 619 26 27 620 28 29 621 30 31 622 32 33 623 34 624 36 625 37 38 626 39 40 627 41 42 628 43 44 629 45 630 47 631 48 49 632 50 51 633 52 53 634 54 55 635 56 636 of crustacean amphipods in the Fusaro Coastal Lagoon (Central Tyrrhenian Sea, Italy) and some notes on their presence in Mediterranean lagoons. Mar. Ecol. 13, 203-224. Ruffo, S., 1982-1998. The Amphipoda of the Mediterranean: Parts 1-4. Mémoires de l’Institut Océanographique, Monaco. Sainte-Marie, B., 1986. Feeding and swimming of lysianassid amphipods in a shallow cold-water bay. Mar. Biol. 91, 219-229. Sánchez-Mata, A., Lastra, M., Mora, J., 1993. Macrobenthic crustacean characterization of an estuarine area. Crustaceana 64, 337-355. Sarvala, J., Uitto, A., 1991. Production of the benthic amphipod Pontoporeia affinis and P. femorata in a Baltic archipelago. Ophelia 34, 71-90. Saunders, C.G., 1966. Dietary analysis of caprellids (Amphipoda). Crustaceana 10, 314- 316. Scipione, M.B., 1989. Trophic behaviour of Amphipoda in some benthic systems. Oebalia 15, 249-260. Sparla, M.P., Scipione, M.B., Riggio, S., 1993. Peracarid Crustacea inhiting aegagropylae of the red alga Rytiphloea tinctoria (Clemente) C. Ag. in the Stagnone Sound, Western Sicily, Italy. Crustaceana 64, 1-17. Shillaker, R.O., Moore, P.G., 1987. The feeding habits of the amphipods Lembos websteri Bate and Corophium bonnellii Milne Edwards. J. Exp. Mar Biol. Ecol. 110, 93-112. Søreide, J.E., Nygård, H., 2012. Challenges using stable isotopes for estimating trophic levels in marine amphipods. Polar Biol. 35, 447-453. 638 Tierno de Figueroa J.M., Vera, A., López-Rodríguez, M.J., 2006. Adult and nymphal 1 57 58 59 60 61 62 63 64 65 4 24 35 2 639 3 640 5 641 6 7 642 8 9 643 10 11 644 12 13 645 14 646 15 16 647 17 18 648 19 20 649 21 22 650 23 651 25 652 26 27 653 28 29 654 30 31 655 32 33 656 34 657 36 658 37 38 659 39 40 41 42 43 44 45 46 47 48 49 50 51 52 53 54 55 56 feeding in the stonefly species Antarctoperla michaelseni and Limnoperla jaffueli from Central Chile (Plecoptera: Gripopterygidae). Entomol. Gen. 29, 39-45. Torrecilla-Roca, I., Guerra-García, J.M., 2012. Feeding habits of the peracarid crustaceans associated to the alga Fucus spiralis in Tarifa Island, Cádiz (Southern Spain). Zoo. Baet. 23, 39-47. Vázquez-Luis, M., Sánchez-Jerez, P., Bayle-Sempere, J.T., 2013. Does the invasion of Caulerpa racemosa var. cylindracea affect the feeding habits of amphipods (Crustacea: Amphipoda). J. Mar. Biol. Ass. UK 93, 87-94. Watling, L., 1993. Functional morphology of the amphipod mandible. J. Nat. Hist. 27, 837-849. Yu, O.H, Suh, H.L., 2002. Secondary production of Synchelidium lenorostralum (Amphipoda, Oedicerotidae) on a temperate sandy shore, southern Korea. J. Crust. Biol. 22, 467-473. Zimmerman, R., Gibson, R., Harrington, J., 1979. Herbivory and detritivory among gammaridean amphipods from a Florida seagrass community. Mar. Biol. 54, 41-47. 660 TABLE AND FIGURE LEGENDS 1 46 47 48 49 50 51 52 53 54 55 56 57 58 59 60 61 62 63 64 65 686 687 688 689 690 691 Table 1. Gut contents of the studied amphipod species. N: number of specimens of each species examined, n: number of specimens with detected digestive contents. % Abs: total area occupied by the content in the whole digestive tract. Det: detritus, Cru: crustaceans, Pol: polychaetes, Oli: oligochaetes, Kin: kinorhynchs, Hyd: hydroids, MAlg: Macroalgae, µalg: microalgae, Din: dinoflagellates, For: foraminifers. The habitat (Hab) where the species was collected is indicated: Sed (sediments), Alg (algae), Hyd (hydroids). Mean values with standard errors of the mean (in parentheses) are included. Fig. 1. Percentage of the three dominant food items in the diet of the studied amphipod families. The values were obtained as the mean of the different species studied of each family. Representative mandibles of different families are also included. Figures redrawn from Ruffo (1982-1998) 2 661 3 4 662 5 663 6 7 664 8 9 665 10 11 666 12 13 667 14 15 668 16 17 669 18 670 19 20 671 21 22 672 23 24 673 25 26 674 27 675 28 29 676 30 31 677 32 33 678 34 35 679 36 37 680 38 681 39 40 682 41 42 683 43 44 684 45 685 Table 1 Table 1 Components (100%) Hab N/n %Abs %Det %Cru %Pol %Oli %Kin %Hyd %MAlg %µalg %Din %For CAPRELLIDEANS Caprellidae Caprella acanthifera Alg 28/14 18.6(5.1) 93.2(4.1) - 6.6(4.1) - - - - 0.2(0.1) - - Caprella andreae Alg 5/4 6.0(1.7) 100(0) - - - - - - - - - Caprella danilevskii Alg 30/18 9.0(2.3) 90.4(5.4) 8.6(6.3) - - - - - 0.7(0.4) 0.3(0.2) - Caprella dilatata Hyd 41/40 80.0(2.5) 87.8(2.7) 11.0(2.7) - - - 0.4(0.2) - 0.2(0.1) 0.6(0.3) - Caprella equilibra Hyd 64/61 82.8(2.4) 89.7(2.4) 9.6(2.4) - - - 0.4(0.2) - - 0.3(0.1) Caprella erethizon Hyd 6/1 15.0(-) 100(-) - - - - - - - - - Caprella grandimana Alg 13/13 74.1(9.3) 92.8(4.7) 5.3(4.4) - - - - - 1.9(0.5) - - Caprella hirsuta Alg 24/18 50.8(7.8) 97.8(0.8) - - - - - - 1.3(0.5) 0.9(0.4) - Caprella liparotensis Alg 22/9 14.1(5.5) 67.8(13.5 21.1(11.2) - - - 11.1(6.9) - - - - Caprella penantis Alg 82/79 50.7(3.2) 85.4(2.4) 11.0(1.9) - - - - 3.1(1.4) 0.1(0.1) 0.4(0.2) - Caprella sabulensis Sed 7/7 37.2(4.3) 100(0) - - - - - - - - - Caprella santosrosai Hyd 14/12 21.2(4.8) 100(0) - - - - - - - - - Caprella tuberculata Hyd 8/4 9.0(6.6) 100(0) - - - - - - - - - Pariambus typicus Sed 14/11 24.6(7.1) 82.3(8.9) 17.7(8.4) - - - - - - - - Parvipalpus onubensis Sed 2/2 13.5(4.6) 100(0) - - - - - - - - - Pedoculina garciagomezi Alg 9/4 8.2(4.1) 96.2(3.2) - - - - - 3.8(3.2) - - - Phtisica marina Alg, Sed 61/36 17.9(3.4) 93.2(2.6) 5.8(2.4) - - - - - 0.3(0.1) 0.7(0.3) - Pseudolirius kroyeri Sed 5/5 32.0(4.6) 100(0) - - - - - - - - - Pseudoprotella phasma Hyd 32/18 7.8(2.0) 80.4(6.7) 11.5(5.7) 3.9(3.2) - - 3.7(2.3) - 0.5(0.3) - - GAMMARIDEANS Ampeliscidae Ampelisca brevicornis Sed 12/11 60.8(8.5) 92.5(4.5) 4.2(4.2) 3.3(3.3) - - - - - - - Ampelisca diadema Sed 29/29 84.7(3.6) 99.8(0.2) - - - - - - 0.2(0.2) - - Ampelisca sarsi Sed 4/4 65.0(9.6) 87.5(12.5) 12.5(12.5) - - - - - - - - Ampelisca serraticauda Alg 11/11 75.5(4.5) 83.2(7.9) 15.9(8.0) - - - - 0.9(0.9) - - - Ampelisca spinifer Sed 14/14 76.4(6.4) 100(0) - - - - - - - - - Ampelisca spinipes Sed 7/7 93.6(4.0) 100(0) - - - - - - - - - Ampelisca tenuicornis Sed 11/11 77.3(7.0) 100(0) - - - - - - - - - Ampelisca typica Sed 16/16 74.7(5.0) 100(0) - - - - - - - - - Amphilochidae Amphilochus brunneus Sed 9/0 - - - - - - - - - - - Amphilochus neapolitanus Alg 21/14 9.4(1.5) 28.6(12.5) 71.4(12.5) - - - - - - - - Amphilochus picadurus Alg 18/16 12.1(0.9) 92.5(5.2) - - - - 7.5(5.2) - - - - Amphilochus spencebatei Sed 22/18 14.6(4.1) 72.2(10.1) 22.2(9.2) 5.6(5.6) - - - - - - - Gitana sarsi Alg 8/2 10(0) 100(0) - - - - - - - - - Ampithoidae Ampithoe ferox Alg 9/7 42.1(10.0) 60.0(11.9) 25.0(14.9) - - - - 15.0(6.1) - - - Ampithoe gammaroides Alg 2/2 50.0(30) 10.0(0) - 45(45) - - - 35(35) - - - Ampithoe helleri Alg 11/5 25.0(8.9) 96.0(4.0) - 4.0(4.0) - - - - - - - Ampithoe ramondi Alg, Sed 37/34 69.9(3.6) 52.8(5.7) 3.9(2.8) - - - - 40.0(5.9) 3.3(2.5) - - Ampithoe riedli Alg 26/23 52.6(4.5) 43.5(8.3) 4.3(4.3) - - - - 52.2(8.4) - - - Aoridae Aora spinicornis Alg 17/17 39.6(8.1) 100(0) - - - - - - - - - Lembos websteri Alg 23/23 73.0(5.0) 97.4(1.8) 2.6(1.8) - - - - - - - - Microdeutopus anomalus Sed 9/9 81.1(4.9) 100(0) - - - - - - - - - Microdeutopus chelifer Alg 9/9 31.3(7.0) 87.8(11.0) - - - - 11.1(11.1) 1.1(1.1) - - - Microdeutopus versiculatus Sed 17/17 62.6(8.0) 94.7(5.3) 5.3(5.3) - - - - - - - - Atylidae Atylus falcatus Sed 11/8 37.5(8.2) 96.3(2.6) - - - - - 3.7(2.6) - - - Atylus guttatus Sed 6/5 53.0(15.1) 66.0(17.2) - - - - - 34.0(17.2) - - - Atylus vedlomensis Sed 20/15 34.3(7.2) 100(0) - - - - - - - - - Calliopiidae Apherusa alacris Sed 15/15 66.7(6.9) 100(0) - - - - - - - - - Apherusa bispinosa Alg 22/21 44.7(5.7) 96.2(3.8) 3.8(3.8) - - - - - - - - Apherusa jurinei Alg 19/19 54.7(4.5) 88.4(5.8) 2.6(2.6) 1.1(1.1) - - - 5.3(5.3) 2.6(1.7) - - Colomastigidae Colomastix pusilla Alg 1/0 - - - - - - - - - - - Corophiidae Apocorophium acutum Alg 22/16 49.4(5.8) 100(0) - - - - - - - - - Corophium multisetosum Sed 19/17 46.8(6.0) 100(0) - - - - - - - - - Leptocheirus hirsutimanus Sed 25/17 43.5(8.3) 100(0) - - - - - - - - - Leptocheirus pectinatus Sed 23/22 83.9(5.0) 100(0) - - - - - - - - - Medicorophium annulatum Sed 25/23 51.5(5.1) 100(0) - - - - - - - - - Mediocorophium longisetosum Sed 14/12 58.8(6.5) 100(0) - - - - - - - - - Medicorophium runcicorne Sed 15/12 51.7(4.6) 100(0) - - - - - - - - - Monocorophium acherusicum Sed 24/18 50.6(5.0) 100(0) - - - - - - - - - Monocorophium insidiosum Sed 24/23 64.6(4.5) 100(0) - - - - - - - - - Table 1 (Cont.) Cressidae Cressa cristata Alg 1/0 - - - - - - - - - - - Cressa mediterranea Alg 8/0 - - - - - - - - - - - Cyproideidae Peltocoxa gibbosa Alg 1/0 - - - - - - - - - - - Peltocoxa marioni Alg 2/1 5(-) 80(-) - - - - 20(-) - - - - Cheirocratidae Cheirocratus intermedius Sed 12/12 84.6(3.5) 100(0) - - - - - - - - - Cheirocratus sundevalli Sed 9/9 72.2(8.1) 100(0) - - - - - - - - - Dexaminidae Dexamine spiniventris Alg 12/6 25.8(9.5) 92.5(5.1) - - - - - 7.5(5.1) - - - Dexamine spinosa Alg 23/14 44.6(9.5) 54.3(13.1) - - - - - 45.7(13.1) - - - Guernea coalita Alg 17/6 32.5(8.1) 33.3(21.1) 41.7(20.1) - 25.0(17.1) - - - - - - Tritaeta gibossa Alg 20/14 23.2(5.9) 100(0) - - - - - - - - - Hyalidae Apohyale prevostii Alg 17/6 40.0(8.9) 76.7(16.7) 5.0(5.0) 6.7(6.7) - - - 11.7(11.7) - - - Hyale perieri Alg 286/233 50.9(4.3) 7.6(2.9) 14.7(5.4) - - - - 62.1(5.6) 4.0(0.7) 10.6(1.4) - Hyale pontica Alg 40/3 66.7(8.8) 56.7(29.6) - - - - - 43.3(29.6) - - - Hyale schmidti Alg 33/31 53.8(5.3) 100(0) - - - - - - - - - Hyale spinidactyla Alg 20/12 53.3(5.1) 48.9(13.3) 50.8(13.3) - - - - - 0.3(0.3) - - Hyale stebbingi Alg 26/17 59.1(8.2) 46.1(12.0) 6.3(6.3) - - - - 47.2(12.1) 0.4(0.3) - - Iphimediidae Iphimedia minuta Alg 9/7 33.6(7.8) 97.1(2.9) - - - - - - 2.9(2.9) - - Ischyroceridae Ericthonius punctatus Alg, Sed 35/27 45.0(5.3) 100(0) - - - - - - - - - Ischyrocerus inexpectatus Alg 9/7 64.3(6.5) 100(0) - - - - - - - - - Jassa cadetta Alg 50/35 53.3(4.7) 100(0) - - - - - - - - - Jassa ocia Alg 2/0 - - - - - - - - - - - Jassa pusilla Alg 4/4 60.0(10.8) 100(0) - - - - - - - - - Parajassa pelagica Alg 53/49 63.9(3.0) 88.3(2.8) 11.1(2.7) - - - - 0.6(0.6) - - - Siphonoecetes kroyeranus Sed 55/47 33.5(3.5) 100(0) - - - - - - - - - Siphonoecetes sabatieri Sed 70/60 41.7(4.0) 100(0) - - - - - - - - - Leucothoidae Leucothoe incisa Sed 33/15 52.0(6.9) 86.7(6.4) 9.3(5.6) 4.0(4.0) - - - - - - - Leucothoe lilljeborgi Sed 2/2 60.0(10.0) 40.0(10.0) 60.0(10.0) - - - - - - - - Leucothoe oboa Sed 3/2 40.0(30.0) 55.0(45.0) 45.0(45.0) - - - - - - - - Leucothoe spinicarpa Alg 13/10 46.5(5.8) 15.0(10.7) 85.0(10.7) - - - - - - - - Liljeborgiidae Liljeborgia dellavallei Alg 2/1 30.0(-) 30.0(-) 70.0(-) - - - - - - - - Lysianassidae Hippomedon denticulatus Sed 8/8 76.3(9.9) 100(0) - - - - - - - - - Hippomedon massiliensis Sed 30/17 69.4(6.0) 58.2(9.9) 41.5(9.9) - - - - - 0.3(0.3) - - Hippomedon oculatus Sed 2/2 40.0(30.0) 100(0) - - - - - - - - - Lepidepecreum longicornis Sed 10/3 64.0(28.5) 100(0) - - - - - - - - - Lysianassa costae Alg 15/8 28.1(6.2) 66.6(15.9) 5.0(5.0) - - - - 27.3(15.1) 1.1(0.6) - - Orchomenella nana Sed 9/2 30.0(20.0) 100(0) - - - - - - - - - Socarnes erythrophthalmus Sed 27/23 60.2(4.4) 58.7(7.9) 41.3(7.9) - - - - - - - - Socarnes filicornis Sed 10/8 38.8(5.8) 96.2(3.8) - - 3.8(3.8) - - - - - - Tryphosella minima Sed 2/2 37.5(32.5) 100(0) - - - - - - - - - Maeridae Animoceradocus semiserratus Sed 26/26 62.3(4.2) 64.4(6.4) 19.8(5.4) - - - - 8.8(3.9) 0.8(0.4) - 6.0(2.9) Elasmopus pocillimanus Alg 7/6 43.3(12.6) 90.0(10.0) - - - - - 10.0(10.0) - - - Elasmopus rapax Alg 9/8 41.3(8.3) 93.8(4.2) 2.5(2.5) - - - - 3.1(3.1) - - 0.6(0.6) Elasmopus vachoni Alg 29/26 53.1(5.3) 54.8(6.8) 6.2(2.8) - - - - 36.3(6.7) - - 2.7(1.3) Maera grossimana Sed 7/7 72.9(8.4) 96.4(2.8) - - - - - 3.6(2.8) - - - Orthomaera othonis Sed 16/16 80.6(4.4) 85.6(4.5) - - - - - 14.4(4.5) - - - Megaluropidae Megaluropus agilis Sed 6/6 74.2(4.2) 100(0) - - - - - - - - - Megaluropus massiliensis Sed 3/3 83.3(6.7) 90.0(10.0) 10.0(10.0) - - - - - - - - Megaluropus monasteriensis Sed 6/6 73.3(11.1) 100(0) - - - - - - - - - Melitidae Abludomelita gladiosa Sed 22/22 70.5(2.6) 96.8(2.4) 3.2(2.4) - - - - - - - - Abludomelita obtusata Sed 2/2 80.0(10.0) 100(0) - - - - - - - - - Gammarella fucicola Alg, Sed 8/7 40.0(10.5) 85.7(7.2) 14.3(7.2) - - - - - - - - Maerella tenuimana Sed 6/6 37.5(13.1) 98.3(1.7) - - - - - - - - 1.7(1.7) Melita hergensis Sed 14/12 54.2(5.1) 93.3(4.3) 6.7(4.3) - - - - - - - - Melita palmata Sed 19/19 54.7(4.1) 96.8(2.2) 3.2(2.2) - - - - - - - - Microprotopidae Microprotopus longimanus Alg 18/9 24.4(7.1) 100(0) - - - - - - - - - Oedicerotidae Deflexilodes griseus Sed 7/4 12.5(2.5) 32.5(11.8) 67.5(11.8) - - - - - - - - Monoculodes carinatus Sed 9/9 47.8(10.2) 80.0(10.4) 20.0(10.4) - - - - - - - - Pereioculodes aequimanus Alg 19/7 28.6(7.6) 50.0(18.9) 50.0(18.9) - - - - - - - - Perioculodes longimanus Sed 25/16 16.9(3.1) 29.4(7.3) 70.6(7.3) - - - - - - - - Pontocrates arenarius Sed 24/13 15.4(2.0) 26.5(11.1) 72.7(11.4) - - - - - 0.8(0.8) - - Synchelidium haplocheles Sed 10/6 34.2(9.9) 16.7(16.7) 83.3(16.7) - - - - - - - - Synchelidium longidigitatum Alg 14/4 51.3(12.3) 30.0(23.8) 70.0(23.8) - - - - - - - - Phliantidae Pereionotus testudo Alg, Sed 16/0 - - - - - - - - - - - Photidae Gammaropsis maculata Alg 21/14 45.9(7.0) 95.6(3.1) 4.3(3.1) - - - - - 0.1(0.1) - - Gammaropsis ostroumowi Alg 7/7 85.7(3.2) 99.0(0.7) 1.0(0.7) - - - - - - - - Gammaropsis palmata Alg 28/26 62.5(5.5) 97.7(2.3) 2.3(2.3) - - - - - - - - Megamphopus cornutus Sed 15/15 82.7(5.5) 100(0) - - - - - - - - - Photis longicaudata Sed 37/29 41.6(4.1) 91.4(5.0) 8.6(5.0) - - - - - - - - Photis longipes Sed 41/30 29.2(5.2) 100(0) - - - - - - - - -