We investigated the taxonomic status of two sympatric morphospecies of squat lobsters from southern South America (Beagle Channel, Strait of Magellan, and Burdwood Bank), Munida gregaria and Munida subrugosa, by DNA sequence analysis of three mitochondrial (mt)DNA gene fragments [416 bp of 16S rDNA(165), 566 bp of cytochrome c oxidase subunit I(COI) and 418 bp of NADH dehydrogenase subunit 1 (ND1)]; and the nuclear rDNA internal transcribed spacer (ITS) 1 (883-952 bp). We obtained a total of 79 sequences from 32 individuals. The 16S sequences of all M. gregaria and M. subrugosa were invariant and identical, whereas COI and ND1 showed 12 and 15 variable sites, respectively. These polymorphisms were shared between morphospecies. Interspecific TamuraNei distances for COI and ND1 sequences were 0.0024 and 0.0032, respectively, and were not significantly different from intraspecific distances (Kruskal-Wallis tests: P = 0.58 and P = 0.69, for COI and ND1, respectively). Similar to the results obtained from the mtDNA sequences, no relationship was found between the ITS1 maximum parsimony tree topology and the morphologic classification of specimens in M. gregaria and M. subrugosa. We conclude that M. gregaria and M. subrugosa from southern South America may either represent a case of a dimorphic species, or a case of incomplete lineage sorting. The fact that these two morphospecies did not show fixed differences over a total of 1947 bp analysed reinforces the hypothesis of a single dimorphic species.
The ''East Pacific Barrier'' has been recognized as the World's largest marine biogeographic barrier. Munida gregaria is the only species of its family with transpacific populations; however, it still remains to be elucidated whether these two distantly located populations belong to the same species. In this study, we investigated the genetic cohesion of M. gregaria across the East Pacific Barrier by analyzing mitochondrial markers. Cytochrome oxidase subunit I and NADH dehydrogenase subunit 1 genes were sequenced for individuals from different areas, i.e., the southeast Pacific, the southern tip of South America, the southwest Atlantic, and the southwest Pacific. A medianjoining network, pairwise F ST s, genetic diversity statistics, and neutrality tests were computed. Our results, i.e., the absence of different haplogroups on both sides of the East Pacific Barrier and existence of shared haplotypes, showed that populations on both sides of this barrier belong to the same species. At a population genetic level, our results suggest that individuals from both regions have been connected, since although some differentiation was found between the southern tip of South America and the southwest Pacific, the southeast Pacific and southwest Atlantic showed no signs of differentiation from the southwest Pacific. In addition, our results provided evidence of a population expansion in southern South America during the Pleistocene. The role of Pleistocene glaciations and the Antarctic Circumpolar Current in shaping the distribution of sub-Antarctic marine invertebrates is discussed.
Resumen.-Se analizó la gametogénesis y la variación temporal en la asignación de energía a diferentes órganos de Aulacomya atra del Canal Beagle (Tierra del Fuego). El periodo de desove de A. atra se extendió desde el final del invierno hasta la primavera, cuando el MGI (Índice gónada-manto) resultó mínimo y el número de individuos desovados, máximo. En los machos maduros el incremento del MGI coincidió con un descenso en la densidad energética del manto-gónada (ED MG), lo que está relacionado con un aumento de masa con menor energía. En hembras maduras el incremento de MGI se relaciona con un aumento de masa con valores más altos de energía, que determina el aumento de la ED MG. Durante la maduración gonadal existen diferentes estrategias de asignación de energía en ambos sexos, aunque se alcanza similar contenido de energía en el manto-gónada; los machos mantienen gónadas más grandes, pero con menos energía por unidad de masa que las hembras. Además, en los individuos maduros desciende el valor del índice de la glándula digestiva (DGI), incrementa el MGI, y tiende a la depleción de la ED y del contenido energético, sugiriendo que este órgano actuaría como reservorio de energía.
Resumen.-Munida gregaria y M. subrugosa fueron originalmente descritas como especies diferentes, pero estudios recientes evidenciaron que ambas especies constituyen una misma especie biológica, y por tanto son referidas como dos morfos de M. gregaria: 'gregaria' y 'subrugosa'. A pesar de su reciente sinonimia taxonómica, la ocurrencia en simpatría de dos morfos representa un caso interesante para estudiar los procesos involucrados en el mantenimiento de los mismos en la naturaleza. Nuestra hipótesis es que las diferentes morfologías de los apéndices bucales de ambos morfos pueden estar relacionadas con hábitos alimentarios diferentes. Se capturaron adultos de M. gregaria, morfos 'gregaria' y 'subrugosa', en el Canal Beagle por medio de lances epibentónicos de arrastre en noviembre de 2004. La superposición dietaria entre los morfos resultó elevada. Ambos ocuparon la misma posición en la trama trófica del Canal Beagle y mostraron los mismos hábitos alimentarios. Como depredadores, se alimentaron principalmente de crustáceos y algas, y como depositívoros ingirieron materia orgánica particulada y organismos unicelulares bentónicos. Sin embargo, la abundancia relativa de crustáceos fue más alta en los estómagos de 'subrugosa' que en los de 'gregaria', una proporción más alta de estómagos de 'gregaria' contenían más algas que los de 'subrugosa', y se encontró una mayor frecuencia de ocurrencia de presas unicelulares en la dieta de 'gregaria'. No obstante estas diferencias, este estudio no provee evidencias que respalden la existencia de un polimorfismo trófico entre los morfos de M. gregaria, al menos sobre la base del hábito alimentario de los adultos. Palabras clave: cangrejos, langostillas, especiación, Atlántico sudoccidentalAbstract.-Munida gregaria and M. subrugosa were originally described as two different species on the basis of several morphological characters, but recent studies evidenced that both former species constitute the same biological species, so hereafter they are referred to as two morphs of M. gregaria: 'gregaria' and 'subrugosa'. Notwithstanding their synonimization, the occurrence of two sympatric morphs represents an interesting case for the study of the processes involved in the maintenance of these two morphotypes in nature. We hypothesized that the distinct shapes of mouthparts of both morphs may be related to different feeding habits. Adults of M. gregaria, morphs 'gregaria' and 'subrugosa', were collected in the Beagle Channel by epibenthic trawling during November 2004. The trophic niche of both morphs highly overlapped. Both occupied the same trophic position in the food web of the Beagle Channel, showing the same feeding habits. As predators they fed mainly on crustaceans and algae, and as deposit feeders they ingested particulate organic matter and organisms associated with the superficial layer of the sediment. However, the relative abundance of crustaceans was higher in stomachs of 'subrugosa' than in those of 'gregaria', a higher proportion of stomachs of 'gregaria' contained more...
The present article is the first formal record of pelagic swarms and beach strandings of the squat lobster Munida gregaria in the Beagle Channel, southern South America. To describe size composition and natural diet of beach strandings and swarms, samples from strandings were taken surveys were carried out to describe both, horizontal distribution and school metrics of swarms. Pelagic swarms and beach strandings were composed of 100% individuals of the morph gregaria of M. gregaria. Mean size of males and females in both, beach strandings and swarms, were similar. The diet of pelagic M. gregaria was composed mainly of crustaceans, unicellular algae and small macroalgae, sediment and particulate organic matter. Swarms of pelagic Munida took the shape of extended layers of varying density and height. Their positions in the water column were also variable: swarms were found at different depths, from the subsurface layer to near the sea bottom. Occasionally, some large swarms occupied most of the water column. In the Beagle Channel, SW winds during spring tides could be a forcing factor for M. gregaria strandings. The absence of shoaling animals in the Beagle Channel between 1997 and 2002 and the recent occurrence of swarms are coincident with their appearance in other locations in Patagonia. In consequence, we hypothesize that the factor influencing the formation of swarms must be acting at a regional scale.
Munida gregaria and M. subrugosa have been considered two different species for more than a century; however, after a recent molecular phylogenetic study, they are considered a single polymorphic species. Yet, the use of markers to diagnose species may be misleading when divergence between species is recent, since a speciation event may be obscured by the retention and stochastic sorting of ancestral polymorphisms. The morphs gregaria and subrugosa of Munida gregaria constitute an interesting case for the study of behavioural isolation since they are sympatric, breed at the same time of the year, and might have experienced a recent speciation. Mating behaviour observations and mate choice mating trials were conducted in order to investigate the potential existence of a behavioural prezygotic barrier to gene flow between these two morphs. Since factors involved in mate choice in galatheids are unknown, the four possible combinations of the two different morphs in trios were used to test for the existence of mate choice. Video recordings of all the possible trio combinations revealed that there was cross-attraction between males and females of different morphs. Females bearing partial broods participated in encounters as well as non-ovigerous females. The frequency and duration of homo-and heterotypic encounters were registered, and a reproductive isolation index was calculated for each variable for each trio. The isolation indexes calculated were not different from zero indicating random mating, and were not affected by the composition of the trio or the partial ovigerous condition of females. These results provided evidence of the absence of behavioural prezygotic barriers to gene flow between the morphs gregaria and subrugosa of M. gregaria.
The larval development of the squat lobster Munida subrugosa from subantarctic waters of the Beagle Channel (Tierra del Fuego, Argentina) was studied under controlled laboratory conditions of temperature, salinity, and food supply. Developmental times, survival, and growth of larvae and early juveniles were investigated. Hatching of the entire brood always occurred during one night. Larvae were kept in 100ml individual bowls with filtered seawater at 8 ± 0.5°C and fed with Artemia spp. nauplii three times a week. Larvae passed through 6 zoeal instars and one megalopa. Previously, only five zoeal instars were known from this species. Mean cumulative durations of the zoeal stages I to VI were: 20.5 ± 2.5, 33.9 ± 4.1, 43.3 ± 5.4, 52.6 ± 5, 61.2 ± 3.9, and 83days, respectively. By adding the 28days that a single megalopa took to metamorphose to crab I stage, the complete larval development lasted 111days. Highest mortality occurred prior to the moult from the zoea I to zoea II stage (79.21% ± 18.65%) and during the moult from zoea VI to megalopa (92.86%). Carapace length was 1.64 ± 0.06, 1.52 ± 0.16, 1.57 ± 0.26, 1.64 ± 0.21, 2.11 ± 0.35, and 2.58 ± 0.19mm, for zoeal stages I to VI, respectively. Carapace length of megalopae and crab I instars was similar (2.85 ± 0.28 and the 2.84 ± 0.05, respectively). Unlike other subantarctic decapods, which show a tendency towards abbreviated larval development and/or some degree of endotrophy, M. subrugosa shows an extended planktotrophic larval development synchronized with short seasonal plankton production in austral summers.
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