Sequence comparison of the mitochondrial genomes of Plesionika species (Caridea: Pandalidae), gene rearrangement and phylogenetic relationships of Caridea.

Background: Despite the Caridean shrimps' vast species richness and ecological diversity, controversies persist in their molecular classification. Within Caridea, the Pandalidae family exemplifies significant taxonomic diversity. As of June 25, 2023, GenBank hosts only nine complete mitochondrial genomes (mitogenomes) for this family. The Plesionika genus within Pandalidae is recognized as polyphyletic. To improve our understanding of the mitogenome evolution and phylogenetic relationships of Caridea, this study introduces three novel mitogenome sequences from the Plesionika genus: P.  ortmanni, P. izumiae and P. lophotes. Methods: The complete mitochondrial genomes of three Plesionika species were sequenced utilizing Illumina's next-generation sequencing (NGS) technology. After assembling and annotating the mitogenomes, we conducted structural analyses to examine circular maps, sequence structure characteristics, base composition, amino acid content, and synonymous codon usage frequency. Additionally, phylogenetic analysis was performed by integrating existing mitogenome sequences of true shrimp available in GenBank. Results: The complete mitogenomes of the three Plesionika species encompass 37 canonical genes, comprising 13 protein-coding genes (PCGs), 22 transfer RNAs (tRNAs), two ribosomal RNAs (rRNAs), and one control region (CR). The lengths of these mitogenomes are as follows: 15,908 bp for P. ortmanni, 16,074 bp for P. izumiae and 15,933 bp for P. lophotes. Our analyses extended to their genomic features and structural functions, detailing base composition, gene arrangement, and codon usage. Additionally, we performed selection pressure analysis on the PCGs of all Pandalidae species available in Genbank, indicating evolutionary purification selection acted on the PCGs across Pandalidae species. Compared with the ancestral Caridea, translocation of two tRNA genes, i.e., trnP or trnT, were found in the two newly sequenced Plesionika species-P. izumiae and P. lophotes. We constructed a phylogenetic tree of Caridea using the sequences of 13 PCGs in mitogenomes. The results revealed that family Pandalidae exhibited robust monophyly, while genus Plesionika appeared to be a polyphyletic group. Conclusions: Gene rearrangements within the Pandalidae family were observed for the first time. Furthermore, a significant correlation was discovered between phylogenetics of the Caridea clade and arrangement of mitochondrial genes. Our findings offer a detailed exploration of Plesionika mitogenomes, laying a crucial groundwork for subsequent investigations into genetic diversity, phylogenetic evolution, and selective breeding within this genus.

Multiple transcriptome mining coupled with tissue specific molecular cloning and mass spectrometry provide insights into agatoxin-like peptide conservation in decapod crustaceans.

Over the past decade, in silico genome and transcriptome mining has led to the identification of many new crustacean peptide families, including the agatoxin-like peptides (ALPs), a group named for their structural similarity to agatoxin, a spider venom component. Here, analysis of publicly accessible transcriptomes was used to expand our understanding of crustacean ALPs. Specifically, transcriptome mining was used to investigate the phylogenetic/structural conservation, tissue localization, and putative functions of ALPs in decapod species. Transcripts encoding putative ALP precursors were identified from one or more members of the Penaeoidea (penaeid shrimp), Sergestoidea (sergestid shrimps), Caridea (caridean shrimp), Astacidea (clawed lobsters and freshwater crayfish), Achelata (spiny/slipper lobsters), and Brachyura (true crabs), suggesting a broad, and perhaps ubiquitous, conservation of ALPs in decapods. Comparison of the predicted mature structures of decapod ALPs revealed high levels of amino acid conservation, including eight identically conserved cysteine residues that presumably allow for the formation of four identically positioned disulfide bridges. All decapod ALPs are predicted to have amidated carboxyl-terminals. Two isoforms of ALP appear to be present in most decapod species, one 44 amino acids long and the other 42 amino acids in length, both likely generated by alternative splicing of a single gene. In carideans, a gene or terminal exon duplication appears to have occurred, with alternative splicing producing four ALPs, two 44 and two 42 amino acid isoforms. The identification of ALP precursor-encoding transcripts in nervous system-specific transcriptomes (e.g., Homarus americanus brain, eyestalk ganglia, and cardiac ganglion assemblies, finding confirmed using RT-PCR) suggests that members of this peptide family may serve as locally-released and/or hormonally-delivered neuromodulators in decapods. Their detection in testis- and hepatopancreas-specific transcriptomes suggests that members of the ALP family may also play roles in male reproduction and innate immunity/detoxification.

Crustacean neuroparsins-a mini-review.

Crustacean neuroparsins are poly-cysteine rich neuropeptides that share some similarities with the ovary ecdysteroidogenesis hormone (OEH) of mosquitoes, the N-terminal end of the growth factor binding protein region of the vertebrate and mollusk insulin-like growth factor binding protein and single insulin binding domain protein. Neuroparsins can promote reproduction and neurite outgrowth in various insects. Though many studies have been made in insects, the amount of work reported in crustaceans is still limited. This review emphasizes the neuroparsins found in decapod crustaceans with references to the neuroparsin first discovered in insects. To be more complete in identifying all the neuroparsin members and to understand the structure/function relationship within a single species, we have collected all neuroparsins from the GenBank and our transcriptome datasets. Then, we employed a comparative approach to study the sequence homology, tissue expression patterns, making predictions of their function and the evolutionary relationship particularly in decapod crustaceans. Results from alignment and phylogenetic studies indicated that crustacean neuroparsins consist of unique feature that can be used as criteria for their classification. These features include the presence of 12 cysteine residues in the mature peptide, the strict spacing between these cysteine residues and the size of the mature peptide. Because of the limited data on the expression information, the functions of most neuroparsin are unknown. The review will focus on the site of synthesis, expression, functions, the sequence homology and the evolutionary relationship of this group of neurohormones.

Cloning and characterisation of Na+/K+-ATPase and carbonic anhydrase from oriental river prawn Macrobrachium nipponense.

Na+/K+-ATPase (NKA) and carbonic anhydrase (CA) are ion transporters that play important roles in osmotic pressure balance in crustaceans. Oriental river prawn (Macrobrachium nipponense) is one of the most economically important shrimps in China. This species can live in freshwater and low-salt water, but the molecular mechanism of salinity regulation is unclear. In this study, full-length cDNAs of M. nipponense α-NKA and CA were obtained by rapid amplification of cDNA ends (RACE)-PCR (GenBank accession numbers MH378774 and MH827971, respectively), and characterised. Full-length cDNAs of Mnα-NKA (3778bp) and MnCA (1407bp) contain open reading frames of 3030bp and 930bp, respectively, encoding polypeptides of 1009 and 930 amino acids. Bioinformatic analysis showed that Mnα-NKA and MnCA harbour the representative features of NKAs and CAs, and share high homology with orthologs in other crustaceans. Quantitative real-time PCR revealed that Mnα-NKA and MnCA were expressed in various tissues, with highest expression in posterior gills and hepatopancreas. Both Mnα-NKA and MnCA were up-regulated in response to salinity acclimation, but expression patterns differed. Our results provide a theoretical basis for the mechanism regulating salinity acclimation in M. nipponense.

Decapod crustaceans from the state of Ceará, northeastern Brazil: an updated checklist of marine and estuarine species, with 23 new records.

The present study is the first major assessment of the marine decapod fauna of Ceará, northeastern Brazil, since contributions of J. Fausto-Filho in the 1960s-1970s. A fully updated checklist of all decapod crustaceans occurring in marine and estuarine habitats of Ceará is provided, based on literature records, specimens held in two carcinological collections of the Universidade Federal do Ceará (UFC), and material collected mainly by the authors between 2011 and 2014. A total of 337 decapod species are listed, distributed among the following taxa: Achelata (8 species), Anomura (42 species), Astacidea (1 species), Axiidea (11 species), Brachyura (162 species), Caridea (83 species), Dendrobranchiata (20 species), Gebiidea (9 species), and Stenopodidea (1 species). Among them, 23 species represent new records for Ceará, with 14 species, viz. Alpheus peasei (Armstrong, 1940), A. thomasi Hendrix & Gore, 1973, Ambidexter symmetricus Manning & Chace, 1971, Axianassa australis Rodrigues & Shimizu, 1992, Biffarius biformis (Biffar, 1971), B. fragilis (Biffar, 1970), Leptalpheus axianassae Dworschak & Coelho, 1999, L. forceps Williams, 1965, Lysmata bahia Rhyne & Lin, 2006, L. intermedia (Kingsley, 1878), Merhippolyte americana Holthuis, 1961, Neocallichirus maryae Karasawa, 2004, Ogyrides hayi Williams, 1981, and Typton carneus Holthuis, 1951, now having Ceará as the northern-most limit in their distribution range along the Brazilian coastline. One shrimp species, Lysmata lipkei Okuno & Fiedler, 2010, which was also found in the state of Rio Grande do Norte, possibly represents an invasive taxon in Brazil and the western Atlantic, originating from the Indo-West Pacific. Alpheus buckupi Almeida, Terossi, Araújo-Silva & Mantelatto, 2013, previously recorded from Ceará based on a colour photograph, is confirmed from this state, with specimens from several new localities. A few doubtful records from Ceará are briefly discussed. Colour photographs are provided for most of the taxa newly recorded from Ceará; some species are illustrated in colour for the first time.

Morfogeometría del caparazón del cangrejo Damithrax spinosissimus (Decapoda: Mithracidae) en tres islas del Caribe suroccidental / Morfogeometry shell crab Damithrax spinosissimus (Decapoda: Mithracidae) on three islands in the southwestern Caribbean

ResumenEl cangrejo rey del Caribe Damithrax spinosissimus es un recurso pesquero catalogado en estado de vulnerabilidad y de cuya biología se conoce muy poco. Su distribución agregada en mosaico y la gran heterogeneidad ambiental dada por las características oceanográficas, paisajísticas y ecológicas del Caribe, pueden propiciar a que exista variabilidad fenotípica en función de la procedencia geográfica de los individuos. Por esta razón, el objetivo de esta investigación fue determinar si existe variabilidad morfogeométrica en los caparazones de ejemplares procedentes de tres islas del Caribe suroccidental: Providencia, Rosario y San Bernardo. La primera de ellas con una influencia netamente oceánica y las dos siguientes, cercanas al continente, con mayor influencia de las dinámicas costeras. Para ello se capturaron y fotografiaron 276 individuos: 103 hembras ovadas y 173 machos, sobre los cuales se realizaron análisis morfogeométricos, tomando 12 puntos anatómicos de referencia (Landmarks) en una de las mitades del caparazón. Las diferencias de tamaño entre sexos y entre orígenes geográficos se contrastaron mediante comparaciones pareadas de Fisher, el efecto alométrico se estimó mediante Análisis de Regresión Multivariada y el modelo de pendientes alométricas mediante Análisis Multivariado de Covarianza. La conformación promedio entre sexos y entre orígenes geográficos se computó de las deformaciones relativas, obtenidas mediante análisis de componentes principales; asimismo, las distancias Euclidianas entre las conformaciones promedio se utilizaron para construir un árbol con base en el algoritmo de Neighbour- Joining con una significancia calculada sobre 10 000 permutaciones. Los resultados mostraron diferencias en el tamaño y conformación del caparazón entre sexos y entre las tres islas, siendo más evidentes las diferencias en Providencia. Estos resultados pueden ser explicados de manera no excluyente por diferencias genéticas y plasticidad fenotípica debida a la heterogeneidad ambiental del sector. Este estudio, primero en su clase, es un aporte al conocimiento de la especie y de él se concluye que deberían existir estrategias de manejo diferentes para las tres islas.

AbstractThe Caribbean King Crab, Damithrax spinosissimus is a fishery resource, but few biological studies are available. Its patchy distribution, and the high environmental heterogeneity due to the oceanographic, landscape, and ecological characteristics of the Caribbean Sea, can favor the phenotypic variability according to the geographic origin. For this reason, the objective of our study was to determine morphometic variability in the carapace of the crabs from three Southwestern Caribbean islands: Providence, Rosario and San Bernardo. The former has an oceanic influence, whereas the two latter islands, which are closer to the mainland, have more influence of the coastal dynamic. A total of 276 individuals from the three islands were captured and photographed: 103 females and 173 males; their variation was analyzed from 12 anatomical landmarks marked on one half of the carapace. The differences in the carapace size were calculated using a Fisher’s pairwise comparison; the allometric effect was calculated by Multivariate Regression Analysis; and the Allometric model via Multivariate Analysis of Covariance. The average shape was calculated from the relative wraps RW obtained through PCA analysis; and the Euclidian distances between the shape averages, were used to construct a tree using the Neighbour-Joining algorithm over 10 000 permutations. The results showed significant differences in the size and shape of the carapace between sexes and among the three islands. The differences in the shape of the crabs from Providencia were significantly greater than those found between the crabs of Rosario and San Bernardo. These results can be explained inclusively due to the genetic differences and phenotypic plasticity, due to environmental heterogeneity of the sector. This study, the first of its kind, is a contribution to the knowledge of the species. We concluded that different management strategies should be differently adopted in each of the three studied islands. Rev. Biol. Trop. 64 (2): 667-681. Epub 2016 June 01.

Inversión reproductiva de Aratus pisonii (Decapoda: Sesarmidae): diferencias entre hábitats y análisis de rutas / Reproductive investment of Aratus pisonii (Decapoda: Sesarmidae): differences among habitats and path analysis

Aratus pisonii is an abundant Neotropical mangrove crab that forages on fresh leaf tissue in the canopy and lives in a wide spectrum of habitats. It presents great plasticity in some aspects of its life history becoming an excellent subject for comparative studies. The purpose of this study was to investigate possible reproductive variations among populations of this crab in mangroves with different levels of structural development, and explore the direct and indirect effects of environmental variables on reproductive investment. The study was conducted during the rainy season of 2003 and 2004, in North Western Venezuela. Fecundity (number of eggs/female), dry weight of egg mass (PSH) and egg quality (C/Neggs) were evaluated in five populations of this species, in mangroves under different environmental conditions (estuarine, marine, hypersaline) and diverse degrees of structural development (arboreal, arbustive, scrub). Reproductive output (Ro) was obtained by the ratio Ro=PSH/dry body weight (PSC); other estimations of Ro were tested: Ro1=PSH/carapace length (CL), Ro2=fecundity/CL. A path analysis was used to explore causal relationships between environmental and morphological variables and reproductive investment (PSH, fecundity). Fecundity ranged from 6 463±831 eggs (Mean±SE) in the hypersaline arbustive mangrove to a maximum of 24 584±2 393 eggs in estuarine arboreal mangrove, which presents the greatest structural development. The reproductive output of females showed significant differences among mangroves, with the highest values of Ro1, Ro2 in the estuarine mangrove females. The results of the path analysis indicated a multicausal effect of morphological and environmental variables; in particular, the salinity showed the greatest influence on fecundity and PSH. The condition of the estuarine mangrove forests with lower salinity and higher structural development involving higher quality of mangrove leaves and a wider range of potential resources, may explain the higher reproductive output of females observed in this habitat. Rev. Biol. Trop. 63 (2): 385-399. Epub 2015 June 01.

Aratus pisonii es un cangrejo abundante de los manglares del Neotrópico, que forrajea sobre el tejido foliar fresco del dosel y vive en una amplia variedad de hábitats. Presenta una gran plasticidad en algunos aspectos de su historia de vida, convirtiéndose en un excelente sujeto para estudios comparativos. El propósito de este estudio fue investigar las posibles variaciones reproductivas entre las poblaciones de este cangrejo en manglares con diferentes niveles de desarrollo estructural, y explorar los efectos directos e indirectos de las variables ambientales sobre la inversión reproductiva. El estudio se realizó durante la temporada de lluvias 2003 y 2004, en la región Noroccidental de Venezuela. Se evaluaron la fecundidad (número de huevos/hembra), el peso seco de la masa de huevos (PSH) y la calidad del huevo (C/Nhuevos) en cinco poblaciones de esta especie, que viven en manglares bajo diferentes condiciones ambientales (estuarinos, marinos, hipersalinos) y diferente grados de desarrollo estructural (arbóreo, arbustivo, achaparrado). El rendimiento reproductivo (Ro) se obtuvo por la relación Ro=PSH/peso seco del cuerpo (PSC); se probaron otras estimaciones de Ro: Ro1=PSH/longitud del caparazón (LC), Ro2=fecundidad/LC. Se utilizó un análisis de rutas para explorar las relaciones causales entre las variables ambientales y morfológicas con la inversión reproductiva (PSH, fecundidad). La fecundidad varió de 6 463±831 huevos (media±ES) en el manglar arbustivo hipersalino, hasta un máximo de 24 584±2 393 huevos en el manglar arbóreo estuarino, que presentó el mayor desarrollo estructural. El rendimiento reproductivo de las hembras mostró diferencias significativas entre manglares, encontrándose los más altos valores de Ro1, Ro2 en las hembras de manglar estuarino. Los resultados del análisis de rutas indicaron un efecto multicausal de variables morfológicas y ambientales; en particular, la salinidad mostró la mayor influencia sobre la fecundidad y el PSH. La condición del manglar estuarino con menor salinidad y mayor desarrollo estructural, que de acuerdo con estudios anteriores supone una mayor calidad de las hojas de mangle y una mayor variedad de recursos potenciales, puede explicar la mayor inversión reproductiva observada en las hembras de este hábitat.

Distribution of the Planktonic Shrimp Lucifer (Thompson, 1829) (Decapoda, Sergestoidea) off the Amazon / Distribuição do camarão planctônico Lucifer (Thompson, 1829) (Decapoda, Sergestoidea) na plataforma continental amazônica

Lucifer faxoni (BORRADAILE, 1915) and L. typus (EDWARDS, 1837) are species first identified in the neritic and oceanic waters off the Amazon. Samplings were made aboard the vessel “Antares” at 22 stations in July and August, 2001 with a bongo net (500-µm mesh size). Hydrological data were taken simultaneously for comparative purposes. L. faxoni was present at thirteen of the fourteen neritic stations analysed, as well as at five of the eight oceanic stations. L. typus was present at three of the fourteen neritic stations and in one of the eight oceanic stations. The highest density of L. faxoni in the neritic province was 7,000 ind.m–3 (St. 82) and 159 ind.m–3 (St. 75) in the oceanic area. For L. typus, the highest density observed was 41 ind.m–3 (St. 64) in the neritic province. In the oceanic province, the highest single density value was recorded at station 75 (5 ind.m–3). Overall, L. faxoni typically presented the highest densities.

Lucifer faxoni (BORRADAILE, 1915) e L. typus (EDWARDS, 1837) são espécies identificadas pela primeira vez nas águas neríticas e oceânicas da Amazônia. As coletas foram realizadas a bordo do navio “Antares” em 22 estações em julho e agosto de 2001, com uma rede bongô (500 µm). Dados hidrológicos foram tomados simultaneamente para fins comparativos. L. faxoni esteve presente em 13 das 14 estações neríticas analisadas, bem como em 5 das 8 estações oceânicas. L. typus esteve presente em 3 das 14 estações neríticas e em 1 das 8 estações oceânicas. A maior densidade de L. faxoni na província nerítica foi de 7.000 ind.m–3 (estação 82) e 159 ind.m–3 (estação 75) na área oceânica. Para L. typus, a maior densidade observada foi de 41 ind.m–3 (estação 64), na província nerítica. Na província oceânica, o maior valor de densidade foi registrado na estação 75 (5 ind.m–3). No geral, L. faxoni tipicamente apresentou as maiores densidades.

Do omnivorous shrimp influence mayfly nymph life history traits in a tropical island stream?

Interspecific interactions can play an important role in determining habitat selection and resource use between competing species. We examined interactions between an omnivorous shrimp and a grazing mayfly, two co-dominant taxa found in Puerto Rican headwater streams, to assess how predator presence may influence mayfly resource use and instantaneous growth in a tropical rainforest ecosystem. We conducted a series of behavioral and growth experiments to determine the effects of the freshwater shrimp, Xiphocaris elongata, on the growth rate and resource selection of mayfly nymphs in the family Leptophlebiidae. For resource choice assessments, we conducted a series of five day laboratory experiments where mayflies were given access to two resource substrate choices (cobble vs. leaves) in the presence or absence of shrimp. To assess for the effects of shrimp on mayfly fitness, we measured mayfly growth in laboratory aquaria after five days using four treatments (cobble, leaves, cobble + leaves, no resource) in the presence or absence of shrimp. In resource choice experiments, mayflies showed preference for cobble over leaf substrata (p<0.05) regardless of the presence of shrimps, however, the preference for cobble was significantly greater when shrimp were present in the leaf habitat. In growth experiments, there were no statistical differences in mayfly growth in the presence or absence of shrimp (p=0.07). However, we measured increased mayfly nymph growth in the absence of predators and when both cobble and leaves were available. Our results suggest that interspecific interactions between these taxa could potentially influence organic matter resource dynamics (e.g., leaf litter processing and export) in Puerto Rican streams. Rev. Biol. Trop. 62 (Suppl. 2): 41-51. Epub 2014 April 01.

Las relaciones interespecíficas pueden jugar un papel importante en la determinación del uso de recursos y selección de hábitat entre especies que compiten. Hemos examinado las interacciones entre un camarón omnívoro y un efemeróptero que se alimenta de vegetación , dos taxones codominantes en las partes altas de arroyos de Puerto Rico, para evaluar el efecto de los depredadores en el uso de recursos del efemeróptero y el crecimiento instantáneo en un ecosistema de selva tropical. Examinamos experimentalmente el efecto del camarón Xiphocaris elongata sobre la tasa de crecimiento y selección de recursos en ninfas efemerópteras (Leptophlebiidae). Hicimos experimentos de laboratorio de 5 días, dando a los efemerópteros acceso a dos opciones de sustrato (piedrecillas u hojas) en presencia o ausencia del camarón. Para evaluar los efectos sobre el crecimiento probamos cuatro tratamientos (piedrecillas, hojas , piedrecillas + hojas, sin ambas), en presencia o ausencia del camarón. Los efemerópteros prefirieron las piedrecillas a las hojas ( p < 0,05 ) independientemente de la presencia de los camarones. Esta preferencia fue mayor cuando junto a las hojas había camarones. Los camarones no afectaron el crecimiento de los efemerópteros (p = 0,07). Las ninfas efemerópteras crecieron mejor sin camarones y en acuarios con piedrecillas y hojas. Nuestros resultados sugieren que las interacciones interespecíficas entre estos animales podrían influir en la dinámica de la materia orgánica (por ejemplo, procesamiento de la hojarasca y su exportación) en los arroyos puertoriqueños.

Effects of insect and decapod exclusion and leaf litter species identity on breakdown rates in a tropical headwater stream

High species richness of tropical riparian trees influences the diversity of organic detritus entering streams, creating temporal variability in litter quantity and quality. We examined the influence of species of riparian plants and macroinvertebrate exclusion on leaf-litter breakdown in a headwater stream in the Luquillo Mountains of Puerto Rico. Leaf litter of Dacryodes excelsa (Burseraceae), Guarea guidonia (Meliaceae), Cecropia scheberiana (Moraceae), Manilkara bidentata (Sapotaceae), and Prestoea acuminata (Palmae) were incubated in litter bags in a pool of Quebrada Prieta. Fine mesh bags were used to exclude macroinvertebrates during leaf breakdown, and coarse mesh bags allowed access to decapod crustaceans (juvenile shrimps and crabs) and aquatic insects (mainly mayflies, chironomids, and caddisflies). D. excelsa and G. guidonia (in coarse- and fine-mesh bags) had significantly higher breakdown rates than C. scheberiana, M. bidentata, and P. acuminata. Breakdown rates were significantly faster in coarse-mesh bag treatments for all leaf types, thus indicating a positive contribution of macroinvertebrates in leaf litter breakdown in this headwater stream. After 42 days of incubation, densities of total invertebrates, mayflies and caddisflies, were higher in bags with D. excelsa and G. guidonia, and lower in P. acuminata, C. scheberiana y M. bidentata. Decay rates were positively correlated to insect densities. Our study highlights the importance of leaf identity and macroinvertebrate exclusion on the process of leaf litter breakdown in tropical headwater streams. Rev. Biol. Trop. 62 (Suppl. 2): 143-154. Epub 2014 April 01.

La gran variedad de especies vegetales junto a los ríos tropicales afecta la diversidad de detritos orgánicos que llegan a los arroyos, creando variaciones temporales en cantidad y calidad de hojarasca. Examinamos la influencia de las especies vegetales y de la exclusión de macroinvertebrados sobre la descomposición de hojarasca en la parte alta de un arroyo en las montañas Luquillo de Puerto Rico. Incubamos hojarasca de Dacryodes excelsa (Burseraceae), Guarea guidonia (Meliaceae), Cecropia scheberiana (Moraceae), Manilkara bidentata (Sapotaceae) y Prestoea acuminata (Palmae) en bolsas descomposición que colocamos en una poza de Quebrada Prieta. Usamos bolsas de malla fina para excluir los macroinvertebrados y de malla gruesa para permitir la entrada a crustáceos decápodos (camarones y cangrejos jóvenes) e insectos acuáticos (principalmente efemerópteros, quironómidos y tricópteros). D. excelsa y G. guidonia (en ambos tipos de malla) tuvieron tasas de descomposición más altas que C. scheberiana, M. bidentata y P. acuminata. La descomposición fue más rápida para todas las especies de hoja con la malla gruesa, resaltando el papel de los descomponedores en este hábitat. Tras 42 días de esta incubación, las densidades totales de invertebrados, efemerópteros y tricópteros fueron mayores en las bolsas con hojas de D. excelsa y G. guidonia, e inferiores en P. acuminata, C. scheberiana y M. bidentata. A mayor densidad de insectos hubo mayor tasa de descomposición. Nuestros resultados indican la importancia de la especie de las hojas y de la presencia de macroinvertebrados en el proceso de descomposición de hojarasca en partes altas de arroyos tropicales.

Population biology of the mangrove crab Ucides cordatus (Decapoda: Ucididae) in an estuary from semiarid Northeastern Brazil / Biología poblacional del cangrejo de manglar Ucides cordatus (Decapoda: Ucididae) en un estuario tropical semiárido

The mangrove crab Ucides cordatus is an important resource of estuarine regions along the Brazilian coast. U. cordatus is distributed from Florida, USA, to the coast of Santa Catarina, Brazil. The species plays an important role in processing leaf litter in the mangroves, which optimizes the processes of energy transfer and nutrient cycling, and is considered a keystone species in the ecosystem. Population declines have been reported in different parts of the Brazilian coast. In the present study we evaluated aspects of the population structure, sex ratio and size at morphological sexual maturity. We analyzed 977 specimens collected monthly over 24 months (2010-2012), in a mangrove of the Jaguaribe River, in the municipality of Aracati on the East coast of Ceará state, Northeastern Brazil. The study area has a mild semiarid tropical climate, with mean temperatures between 26 and 28oC. The area is located within the eco-region of the semiarid Northeast coast, where mangroves occur in small areas and estuaries are affected by mesomareal regimes. The population structure was evaluated by the frequency distribution of size classes in each month, and the overall sex ratio was analyzed using the chi-square test. Size at morphological sexual maturity was estimated based on the allometry of the cheliped of the males and the abdomen width of the females, using the program REGRANS. The size-frequency distribution was unimodal in both sexes. The overall sex ratio (M: F) (1:0.6) was significantly different from 1:1. Analysis of the sex ratio by size class showed that the proportion of males increased significantly from size class 55-60mm upward, and this pattern persisted in the larger size classes. In the smaller size classes the sex ratio did not differ from 1:1. The size at morphological sexual maturity was estimated at a carapace width (CW) of 52mm and 45mm for males and females, respectively. Analysis of the population parameters indicated that ...

El cangrejo Ucides cordatus es un importante recurso pesquero de las regiones de estuarios a lo largo de la costa brasileña. Disminuciones de las poblaciones han sido reportadas en diferentes regiones del país. El presente estudio evaluó aspectos de la estructura poblacional, proporción sexual y talla en la madurez sexual. Se analizó un total de 977 ejemplares capturados mensualmente, por 24 meses, en lo manglar del río Jaguaribe, en el municipio de Aracati, en la costa este de Ceará, nordeste del Brasil. El área de estudio tiene un clima semiárido tropical, con temperaturas medias entre los 26 y 28°C. La estructura poblacional fue evaluada para determinar la distribución de frecuencia de clases de talla. Para analizar diferencias significativas respecto a la proporción sexual se aplicó la prueba Chi-quadrado. La talla de la madurez sexual morfológica fue estimada con base en la alometría del propodito del quelípedo de los machos y de la anchura del abdomen de las hembras, utilizando el programa REGRANS. La distribución de frecuencia de talla fue unimodal para los dos sexos. La proporción sexual total (M:H)(1:0.6), se desvió significativamente de la igualdad. El análisis de la proporción entre sexos, relativo a las clases de tallas mostró que la proporción de machos se tornó significativamente más alta desde la clase de 55-60mm de talla, y este se mantuvo en las clases de talla más grande. No obstante, en las clases más pequeñas la proporción entre los sexos no fue distinta de 1:1. La talla de la madurez sexual morfológica fue de 52mm de ancho del caparazón y 45mm para los machos y hembras, respectivamente. Los resultados de los atributos analizados permitieran deducir que la población de U. cordatus en el manglar del río Jaguaribe se encuentran en una situación de estabilidad. Sin embargo, el monitoreo constante de la población es necesario para detectar alteraciones en los atributos de la población que puedan afectar esta situación de estabilidad.

Composición y estructura de la macrofauna asociada con agregaciones de dos especies de bivalvos en Isla de Cubagua, Venezuela / Composition and structure of the macrofauna associated with beds of two bivalve species in Cubagua Island, Venezuela

Bivalve aggregations constitute a microhabitat for a wide variety of organisms in intertidal, subtidal and deep-water marine benthic habitats. Increase in density of bivalve beds could offer more crevices and substratum for the associated fauna, affecting community composition. Beds of the Atlantic Pearl Oyster (Pinctada imbricata) and the Turkey Wing (Arca zebra) of contrasting population densities were evaluated to determine the composition and structure of the associated macrofauna of three taxa (Crustacea Decapoda, Mollusca and Echinodermata). We evaluated plots of three levels of bed density in both species, the associated fauna were identified and counted. Other species were collected by qualitative samples. Univariate and multivariate descriptors were tested comparing the associated fauna between the beds of two species at three levels of density. In these beds a total of 104 species belonging to 58 families were recorded. Mithraculus forceps (Majidae), Crucibulum auricula (Calyptraeidae) and Ophiotrix angulata (Ophiothrichidae) were the most common species found in these assemblages. The medium and high-density bivalve beds exhibited greater species number, abundance, Shannon diversity, evenness, taxonomic diversity, and taxonomic distinctness of associated fauna, than low-density bivalve beds. Moreover, multivariate analysis detected different assemblages of associated fauna between beds with different densities. Additionally, similarities were found in the communities of macrofauna in both beds of P. imbricata and Arca zebra. Our results suggest that bivalve aggregations at Cubagua Island provide additional habitat for macrofauna living in other shallow habitats such as Thalassia beds, corals and rocky environments. Bed density, associated with topographic complexity, represents an important factor for the composition and complexity of the associated fauna.

Las agregaciones de bivalvos constituyen un microhábitat para una gran variedad de organismos en el ambiente intermareal, submareal y en aguas profundas. Agregaciones de la ostra perla (Pinctada imbricata) y pepitona (Arca zebra) a diferentes densidades poblacionales se evaluaron para determinar la composición y estructura comunitaria de la macrofauna asociada en tres taxa (Crustacea Decapoda, Mollusca y Echinodermata). La hipótesis nula de no diferencias en descriptores univariados y multivariados fue probada comparando la fauna asociada entre las agregaciones de las dos especies a tres niveles de densidad. En estas agregaciones se identificaron 102 especies de 55 familias. Mithraculus forceps (Majidae), Crucibulum auricula (Calyptraeidae) y Ophiotrix angulata (Ophiothrichidae) fueron las especies más comunes encontradas en estas asociaciones. Las densidades medias y altas de las agregaciones de bivalvos presentaron mayor número de especies, abundancia, diversidad de Shannon, equidad, diversidad taxonómica y distinción taxonómica de la fauna asociada que las agregaciones de baja densidad poblacional. Análisis multivariados detectaron diferentes estructuras de los ensambles de la fauna asociada en agregaciones de bivalvos con densidad baja en comparación con los de densidad media y alta. Adicionalmente no se detectaron diferencias en la fauna asociada entre las especies. La densidad de las agregaciones de bivalvos, asociada a la complejidad topográfica, es un factor importante para la composición de la fauna asociada.

Phylogenetic relationships among genera of the Periclimenes complex (Crustacea: Decapoda: Pontoniinae) based on mitochondrial and nuclear DNA.

The genus Periclimenes Costa, 1844 is the most species-rich genus in the subfamily Pontoniinae. Recent studies have suggested that it might be a polyphyletic taxon and could be further subdivided. In this study, three protein-coding nuclear genes and one mitochondrial ribosomal gene were used to analyze the phylogenetic relationships among the genera of the Periclimenes complex, includes the genus Periclimenes and 15 related genera, viz. Ancylomenes, Brucecaris, Crinotonia, Cuapetes, Harpiliopsis, Harpilius, Laomenes, Leptomenaeus, Manipontonia, Palaemonella, Periclimenella, Philarius, Phycomenes, Unguicaris and Vir. Based on both independent and combined data analyses, the results support that the genus Periclimenes is a polyphyletic group. Furthermore, the studied Periclimenes species could be divided into several independent groups, and the taxonomic status of P. commensalis, P. brevicarpalis and P. digitalis may need to be reconsidered. Besides, the majority of the related genera of Periclimenes are suggested to be monophyletic. Our analyses also reveal that these genera approximately form two main clades, despite some deep relationships are still obscure.

Freshwater decapods of Puerto Rico: a checklist and reports of new localities.

An updated checklist of the freshwater decapod species of Puerto Rico is presented based on records of shrimp and crab species whose presence has been confirmed in Puerto Rico as a result of extensive field collections, examination of carcinological collections, literature review, and personal communications from researchers. The freshwater decapods fauna of Puerto Rico consists of 18 species of shrimps belonging to eight genera and three families, and one species of crab belonging to the family Pseudothelphusidae.

Evolution and phylogeny of the mud shrimps (Crustacea: Decapoda) revealed from complete mitochondrial genomes.

BACKGROUND: The evolutionary history and relationships of the mud shrimps (Crustacea: Decapoda: Gebiidea and Axiidea) are contentious, with previous attempts revealing mixed results. The mud shrimps were once classified in the infraorder Thalassinidea. Recent molecular phylogenetic analyses, however, suggest separation of the group into two individual infraorders, Gebiidea and Axiidea. Mitochondrial (mt) genome sequence and structure can be especially powerful in resolving higher systematic relationships that may offer new insights into the phylogeny of the mud shrimps and the other decapod infraorders, and test the hypothesis of dividing the mud shrimps into two infraorders. RESULTS: We present the complete mitochondrial genome sequences of five mud shrimps, Austinogebia edulis, Upogebia major, Thalassina kelanang (Gebiidea), Nihonotrypaea thermophilus and Neaxius glyptocercus (Axiidea). All five genomes encode a standard set of 13 protein-coding genes, two ribosomal RNA genes, 22 transfer RNA genes and a putative control region. Except for T. kelanang, mud shrimp mitochondrial genomes exhibited rearrangements and novel patterns compared to the pancrustacean ground pattern. Each of the two Gebiidea species (A. edulis and U. major) and two Axiidea species (N. glyptocercus and N. thermophiles) share unique gene order specific to their infraorders and analyses further suggest these two derived gene orders have evolved independently. Phylogenetic analyses based on the concatenated nucleotide and amino acid sequences of 13 protein-coding genes indicate the possible polyphyly of mud shrimps, supporting the division of the group into two infraorders. However, the infraordinal relationships among the Gebiidea and Axiidea, and other reptants are poorly resolved. The inclusion of mt genome from more taxa, in particular the reptant infraorders Polychelida and Glypheidea is required in further analysis. CONCLUSIONS: Phylogenetic analyses on the mt genome sequences and the distinct gene orders provide further evidences for the divergence between the two mud shrimp infraorders, Gebiidea and Axiidea, corroborating previous molecular phylogeny and justifying their infraordinal status. Mitochondrial genome sequences appear to be promising markers for resolving phylogenetic issues concerning decapod crustaceans that warrant further investigations and our present study has also provided further information concerning the mt genome evolution of the Decapoda.

A new Lepeophtheirus (Copepoda: Siphonostomatoida: Caligidae) from Isla del Coco National Park, Costa Rica, Eastern Tropical Pacific / Un nuevo Lepeophtheirus (Copepoda: Siphonostomatoida: Caligidae) del Parque Nacional Isla del Coco, Costa Rica, Pacífico Tropical Oriental

Among the several groups of copepods that are teleost parasites, the siphonostomatoid family Caligidae is by far the most widespread and diverse. With more than 108 nominal species, the caligid genus Lepeophtheirus von Nordmann is one of the most speciose. There are no reports of this genus in Costa Rican waters. A new species of Lepeophtheirus is herein described based on female specimens collected from plankton samples in waters off Bahía Wafer, isla del Coco, an oceanic island in the Eastern Tropical Pacific. The new species, L. alvaroi sp. nov., has some affinities with other congeners bearing a relatively short abdomen, a wider than long genital complex and a 3-segmented exopod of leg 4. it differs from most of these species by the presence of an unbranched maxillular process and by the relative lengths of the terminal claws of leg 4, with two equally long elements. it is most closely related to two other Eastern Pacific species: L. dissimulatus Wilson, 1905 and L. clarionensis Shiino, 1959. it differs from these species by the proportions and shape of the genital complex, the shape of the sternal furca, the relative length of the maxillar segments, the absence of a pectiniform process on the distal maxillar segment, the length of leg 4 and the armature of leg 5. The new species represents the first Lepeophtheirus described from Costa Rican waters of the Pacific. The low diversity of this genus in this tropi- cal region is explained by its tendency to prefer hosts from temperate latitudes. Until further evidence is found, the host of this Lepeophtheirus species remains unknown.

Entre los varios grupos de copépodos que son parásitos de teleósteos, la familia sifonostomatoide Caligidae incluye los más dispersos y diversos. Con más de 108 especies nominales, el género de calígidos Lepeophtheirus von Nordmann es uno de los más diversos. No existen registros previos de este género en aguas de Costa Rica. Se describe una nueva especie de copépodo calígido del género Lepeophtheirus con base en especímenes femeninos recolectados a partir de muestras de plancton en aguas de Bahía Wafer, isla del Coco, una isla oceánica del Pacífico Tropical Oriental. La nueva especie, L. alvaroi sp. nov., tiene algunas afinidades con congéneres que poseen un abdomen relativamente corto, un complejo genital más ancho que largo y un exópodo de la pata 4 con 3 segmentos. Esta especie difiere de la mayoría de estas especies por la presencia de un proceso maxilular no ramificado y por el tamaño relativo de las garras terminales de la pata 4, con dos elementos igualmente largos. Está más cercanamente relacionada con otras dos especies del Pacífico Oriental: L. dissimulatus Wilson, 1905 y L. clarionensis Shiino, 1959. Difiere de estas especies por las proporciones y forma del complejo genital, la forma de la furca esternal, la longitud relativa de los segmentos maxilares, la ausencia de un proceso pectiniforme en el segmento maxilar distal, la longitud de la pata 4 y la armadura de la pata 5. La nueva especie representa el primer Lepeophtheirus descrito en aguas del Pacífico de Costa Rica. La baja diversidad del género en esta región tropical se explica por su tendencia a preferir hospederos de latitudes templadas. Hasta encontrar más evidencia, el hospedero de esta especie de Lepeophtheirus permanece desconocida.

Faunistic analysis of the caridean shrimps inhabiting seagrasses along the NW coast of the Gulf of Mexico and Caribbean Sea

Seagrass meadows are highly productive and ecologically important habitats in estuaries and coastal lagoons, and contain a variety of faunal communities, from which the caridean shrimps are a dominant component. The purpose of this work was to analyze the environmental parameters of water and sediments, with the biological components in seagrass epifaunal communities, from the Western Gulf of Mexico and the Caribbean Sea. For this, density and diversity of caridean shrimps were analyzed and correlated with environmental parameters and seagrass biomass, and zoogeographic affinities were determined. The spatial distribution of caridean shrimps was recorded for 12 localities with Halodule wrightii and Thalassia testudinum monospecific seagrass meadows. A total of 72 158 individuals of 16 taxa were collected. Among results, the Hippolytidae resulted the most abundant group (92.3%) with eight species, and was followed by Palaemonidae with 7.6% of the abundance and seven species, and the Alpheidae with only one genus. From the total of collected carideans, a 37.3% was found in H. wrightii and 62.7% in T. testudinum. The dominant species were Hippolyte zostericola (12.39ind./m2), Tozeuma carolinense (9.5ind./m2), Thor dobkini (4.84ind./m2) and Palaemonetes vulgaris (4.87ind./m2). The zoogeographic distribution of the carideans presented two groups: species of the Virginian-Carolinean province representing its Southern limit (43.75%) and species of the Brazilian-Caribbean province representing its Northern limit (56.25%). The species H. zostericola, T. carolinense, P. vulgaris, P. pugio and P. intermedius are widely distributed along the Western Atlantic coast. This study has base line information for seagrass habitats, the community of epifaunal carideans and their ecological affinities, previous to the oil spill in the Gulf of Mexico.

Las praderas de pastos marinos son hábitats altamente productivos y ecológicamente importantes a lo largo de las costas y estuarios, en estas se encuentra una gran variedad de comunidades faunísticas, donde los camarones carideos son un componente dominante por su densidad. Los parámetros ambientales del agua y sedimento y los componentes biológicos de biomasa de pastos marinos y comunidad epifaunística fueron recolectados a lo largo del occidente del Golfo de México y Mar Caribe. La densidady diversidad de los camarones carideos fueron analizadas y correlacionadas con los parámetros ambientales y biomasa de pastos, también se determinaron las afinidades zoogeográficas de las especies. La distribución espacial de los camarones carideos fue registrada en 12 localidades con praderas monoespecíficas de los pastos Halodule wrightii y Thalassia testudinum. Un total de 72 158 individuos pertenecientes a 16 taxa fueron recolectados. La familia Hippolytidae incluyó a ocho especies y representó el 92.3% de la abundancia, la familia Palaemonidae comprendió a siete especies y el 7.6%, y la familia Alpheidae estuvo representada por un solo género. Del total de carideos recolectados, el 37.3% se capturó en H. wrightii y el 62.7% en T. testudinum. Las especies dominantes fueron Hippolyte zostericola (12.39ind./m2), Tozeuma carolinense (9.5ind./m2), Thor dobkini (4.84ind./m2) y Palaemonetes vulgaris (4.87ind./m2). La composición zoogeográfica de los carideos estudiados estuvo representada por dos grupos: el primero constituido por especies afín a la provincia Virginiana- Carolineana en su límite más sureño (43.75%), y el segundo por especies de la provincia Brasileña-Caribeña con su límite más norteño (56.25%). Las especies H. zostericola, T. carolinense, P. vulgaris, P. pugio y P. intermedius tienen una amplia distribución a lo largo de la costa noreste Atlántica. Este estudio constituye la línea base de información sobre los hábitats de pastos marinos, la comunidad de camarones carideos epifaunales y sus afinidades ecológicas de recolectas previas al derrame de petróleo en el noreste del Golfo de México.

Adaptive evolution of crustin antimicrobial peptides in decapods.

Antimicrobial peptides (AMPs) are present in a wide range of taxonomic groups and played a crucial role in host adaptation to a diverse array of ever-changing pathogens. Crustin, a cysteine-rich cationic AMP, is known to exhibit antimicrobial activity against Gram-positive and Gram-negative bacteria in decapods. Given their important role in host-immune defense, a large proportion of amino acid substitutions in crustin AMPs are expected to be fixed by natural selection. Utilizing the complete coding nucleotide sequence data of crustin, the present study revealed the pervasive role of positive Darwinian selection in the evolution and divergence of crustin AMPs in decapods. Approximately, 20-35 % of codons in two phylogenetically distinct groups of closely related crustins in Penaeid shrimps are shown to have evolved under positive selection. Interestingly, several of these positively selected sites are located at the carboxyl-terminal region, the region that directly interacts with the invading pathogens. Pathogen-mediated selection pressure could be the likely cause for such an accelerated rate of amino acid substitutions and could have contributed to the structural and functional diversification of crustin AMPs in several taxa.

Invertebrados asociados al coral constructor de arrecifes Pocillopora damicornis en Playa Blanca, Bahía Culebra, Costa Rica / Invertebrates associated with the reef-building coral Pocillopora damicornis at Playa Blanca, Bahía Culebra, Costa Rica

Invertebrates associated with the reef-building coral Pocillopora damicornis at Playa Blanca, Bahía Culebra, Costa Rica. The coral reefs are one of the most diverse ecosystems in the planet, not only because their reef-building species but also because of the species that live above, on, inside and below them. Corals of the genus Pocillopora are recognized globally by harboring an important fauna of invertebrates among their branches, many of those considered obligate symbionts. The present investigation describes the fauna of invertebrates associated with the reef-building coral P. damicornis in Bahía Culebra, Costa Rica, their densities, frequencies, richness and diversities from 2003 to 2004. Five coral colonies were collected every 3-4 months at Playa Blanca within Bahía Culebra. A total of 448 individuals in 35 species were found: Harpiliopsis depressa, Trapezia ferruginea, Alpheus lottini, Fennera chacei, and Petrolisthes haigae, and the bivalve Lithophaga aristata, the predominant species. November was the month with the highest richness, diversity and taxonomic distinctiveness, while August had the lowest values. In general terms, the rainy season showed greater richness of species than the dry season. The species collected and their densities and abundances were similar to other sites along the Eastern Tropical Pacific. Bahía Culebra has been losing coral cover, which could have consequences on the diversity and abundances of associated corals organisms. These consequences include loss in the fertility of those organisms, a reduction in its function as cleaners and protection from coral predators, putting in risk their diversity, affecting the stocks of predatory fish that depends on them. Carrying out permanent monitoring of the associated cryptofauna of the coral Pocillopora is crucial to quantify losses or recoveries in the composition of associated invertebrates, and therefore in the functions that they perform in this ecosystem.

Los arrecifes coralinos son uno de los ecosistemas más diversos en el planeta, tanto por los organismos constructores como por aquellos que viven encima, sobre, dentro y debajo de ellos. Los corales del genero Pocillopora son reconocidos mundialmente por albergar una importante fauna de invertebrados entre sus ramas, los cuales son considerados como simbiontes obligatorios en una gran cantidad de casos. La presente investigación describe la fauna de invertebrados asociados al coral Pocillopora damicornis en Bahía Culebra, Costa Rica, describiendo sus densidades, frecuencias, riquezas y diversidades a través del tiempo. Para esto se colectaron 5 colonias cada 3-4 meses en Playa Blanca, Bahía Culebra. En total se encontraron 448 individuos en 35 especies, siendo Harpiliopsis depressa, Lithophaga aristata, Trapezia ferruginea, Alpheus lottini, Fennera chacei, y Petrolisthes haigae las especies predominantes. Noviembre fue el mes en el que se encontraron los mayores valores en los índices de riqueza, diversidad y diferenciación taxonómica, mientras que agosto fue el que presento los valores más bajos de todos. En términos generales, la época lluviosa mostro mayor riqueza de especies que la época seca. Así mismo, las especies colectados y los valores obtenidos son muy similares a otras zonas del Pacifico Oriental Tropical. Culebra ha venido sufriendo una perdida en la cobertura coralina, que podría tener consecuencias en la diversidad y abundancia de organismos asociados a corales. Estas consecuencias incluyen perdida en la fecundidad de estos organismos, una reducción en su función como limpiadores y protectores de depredadores del coral, poniendo en riesgo su diversidad, lo que puede afectar los stocks de peces depredadores que depende de ellos. Realizar monitoreos permanentes de la criptofauna asociada al coral Pocillopora va a ser determinante para cuantificar perdidas o recuperaciones en la composición de invertebrados asociados, y por ende en las funciones que estos desempeñan en este ecosistema.

Nuevas adiciones a la fauna de crustáceos decápodos de Bahía Culebra, Guanacaste, Costa Rica / New additions to the decapod crustacean fauna of Bahía Culebra, Guanacaste, Costa Rica

New additions to the crustacean decapod fauna of Bahía Culebra, Guanacaste, Costa Rica. The crustacean fauna of Bahía Culebra, Guanacaste, north Pacific, Costa Rica, was studied before the 1940’s. Based on the collections, mainly of the last two decades, deposited in the Zoology Museum of the University of Costa Rica, 15 species of decapods crustaceans belonging in 11 families are added to the 94 species previously reported for Bahía Culebra. This results in a total of 109 decapod species, and 130 crustacean species for the bay. Probably more species are present since several environments of the bay have not been studied, as well as the benthic microcrustaceans. This type of baseline data studies are important because of the rapid changes taking place in Bahía Culebra.

La fauna de crustáceos de Bahía Culebra, Guanacaste, Pacífico norte de Costa Rica, fue estudiada antes de la década de 1940. Basado en recolectas, principalmente de las últimas dos décadas y depositadas en el Museo de Zoología, Universidad de Costa Rica, se presentan aquí 15 nuevos registros de crustáceos decápodos pertenecientes a 11 familias. Estos registros se suman a las 94 especies conocidas previamente para la bahía, para un total de 109 especies de crustáceos decápodos y 130 de crustáceos. Probablemente hay más especies ya que algunos ambientes no han sido estudiados, como tampoco los microcrustáceos bénticos. Este tipo de estudio de línea base es importante especialmente en regiones donde se están dando alteraciones ambientales muy rápidas.