A review of the distribution of hydrothermal vent communities along the northern Mid-Atlantic Ridge: dispersal vs. environmental controls

Hydrobiologia - Until 1985, seven vent fields were described from the Mid-Atlantic Ridge (MAR). An eighth field, Mount Saldanha (36° N), discovered in 1998, showed unusual geological and...     

The enigmatic,deep‐sea,organic‐walled genera Chitinosiphon,Nodellum and Resigella (Protista,Foraminifera): A taxonomic re‐evaluation

Abstract

Using a morphology‐based approach, we explore the relationships between three poorly understood species of organic‐walled Foraminifera. Thalmann and Bermudez (1954) described Chitinosiphon rufescens as the type species of a new monotypic genus which they compared to the tubular agglutinated foraminiferan Bathysiphon. Loeblich and Tappan (1964), however, considered C. rufescens to be identical to Reophax membranacea Brady 1879, type species of another organic‐walled genus, Nodellum. Based on a re‐examination of the type specimens of both species, new material of C. rufescens from the Lost City hydrothermal field, and new material of N. membranacea from the NE Atlantic and Pacific Oceans, we show that these two deep‐sea species are distinguished by the following features. (1) Chitinosiphon rufescens lacks the distinct, regular constrictions that divide the tubular test of N. membranacea into a series of segments. (2) The proloculus is spindle‐shaped in C. rufescens but sub‐cylindrical in N. membranacea. (3) A distinctive, pocket‐like invagination is developed at the base (proximal end) of the proloculus in N. membranacea but not in C. rufescens. However, a series of undescribed species which occur in deep‐sea sediments blur the distinction between the two genera. We therefore adopt a conservative position and regard Chitinosiphon as a junior synonym of Nodellum. We also examined the holotype and new material of Nodellum moniliforme Resig, 1982, the type species of Resigella Loeblich and Tappan, 1984, in which the organic‐walled test comprises a series of bulbous chambers. This species exhibits a basal invagination, identical to the feature present in Nodellum membranacea. These three remarkable species are united by the basically tubular test and the nature of the test wall which is largely organic, brownish in colour, and exhibits no internal structure when broken sections are examined by SEM. The surface of the organic test of Nodellum rufescens from Lost City is strewn with tiny (≤1μm), needle‐shaped mineral particles, visible only by SEM. More equidimensional, micron‐sized particles are present in the other two species. We agree with Thalmann and Bermudez (1954) that N. rufescens is related to tubular agglutinated taxa such as Bathysiphon. Resigella may have similar affinities, although this needs to be tested using molecular approaches.     

Copepoda from deep-sea hydrothermal vents and cold seeps

Recent explorations of hydrothermal vents in the eastern Pacific (Juan de Fuca spreading zone, Guaymas Basin in the Gulf of California, East Pacific Rise at 21° N and 13° N, and Galapagos Rift) and on the Mid-Atlantic Ridge have revealed many copepods, mostly siphonostomatoids with few poecilostomatoids. In these habitats in depths from 1 808 to 3 650 m water temperatures may reach nearly 15 ° C. Among more than 22 000 copepods from vents examined two new families, 11 new genera, and 32 new species were represented.In addition, two new copepods were found in 3 260 m at cold seeps at the base of the West Florida Escarpment in the Gulf of Mexico, an environment not thermally active, with water temperatures about 4.39 °C.Some of these copepods were associated with host invertebrates such as a Nuculana-like protobranch bivalve, a polychaete, and two species of shrimps. Others were obtained from washings of bivalves or vestimentiferans or by means of corers or slurp guns.     

The dynamics of biogeographic ranges in the deep sea

Anthropogenic disturbances such as fishing, mining, oil drilling, bioprospecting, warming, and acidification in the deep sea are increasing, yet generalities about deep-sea biogeography remain elusive. Owing to the lack of perceived environmental variability and geographical barriers, ranges of deep-sea species were traditionally assumed to be exceedingly large. In contrast, seamount and chemosynthetic habitats with reported high endemicity challenge the broad applicability of a single biogeographic paradigm for the deep sea. New research benefiting from higher resolution sampling, molecular methods and public databases can now more rigorously examine dispersal distances and species ranges on the vast ocean floor. Here, we explore the major outstanding questions in deep-sea biogeography. Based on current evidence, many taxa appear broadly distributed across the deep sea, a pattern replicated in both the abyssal plains and specialized environments such as hydrothermal vents. Cold waters may slow larval metabolism and development augmenting the great intrinsic ability for dispersal among many deep-sea species. Currents, environmental shifts, and topography can prove to be dispersal barriers but are often semipermeable. Evidence of historical events such as points of faunal origin and climatic fluctuations are also evident in contemporary biogeographic ranges. Continued synthetic analysis, database construction, theoretical advancement and field sampling will be required to further refine hypotheses regarding deep-sea biogeography.     

Bathymetric and geographic population structure in the pan-Atlantic deep-sea bivalve Deminucula atacellana (Schenck, 1939)

The deep-sea soft-sediment environment hosts a diverse and highly endemic fauna of uncertain origin. We know little about how this fauna evolved because geographic patterns of genetic variation, the essential information for inferring patterns of population differentiation and speciation are poorly understood. Using formalin-fixed specimens from archival collections, we quantify patterns of genetic variation in the protobranch bivalve Deminucula atacellana, a species widespread throughout the Atlantic Ocean at bathyal and abyssal depths. Samples were taken from 18 localities in the North American, West European and Argentine basins. A hypervariable region of mitochondrial 16S rDNA was amplified by polymerase chain reaction (PCR) and sequenced from 130 individuals revealing 21 haplotypes. Except for several important exceptions, haplotypes are unique to each basin. Overall gene diversity is high (h = 0.73) with pronounced population structure (Phi(ST) = 0.877) and highly significant geographic associations (P < 0.0001). Sequences cluster into four major clades corresponding to differences in geography and depth. Genetic divergence was much greater among populations at different depths within the same basin, than among those at similar depths but separated by thousands of kilometres. Isolation by distance probably explains much of the interbasin variation. Depth-related divergence may reflect historical patterns of colonization or strong environmental selective gradients. Broadly distributed deep-sea organisms can possess highly genetically divergent populations, despite the lack of any morphological divergence.     

High connectivity across the fragmented chemosynthetic ecosystems of the deep Atlantic Equatorial Belt: efficient dispersal mechanisms or questionable endemism?

Chemosynthetic ecosystems are distributed worldwide in fragmented habitats harbouring seemingly highly specialized communities. Yet, shared taxa have been reported from highly distant chemosynthetic communities. These habitats are distributed in distinct biogeographical regions, one of these being the so‐called Atlantic Equatorial Belt (AEB). Here, we combined genetic data (COI) from several taxa to assess the possible existence of cryptic or synonymous species and to detect the possible occurrence of contemporary gene flow among populations of chemosynthetic species located on both sides of the Atlantic. Several Evolutionary Significant Units (ESUs) of Alvinocarididae shrimp and Vesicomyidae bivalves were found to be shared across seeps of the AEB. Some were also common to hydrothermal vent communities of the Mid‐Atlantic Ridge (MAR), encompassing taxa morphologically described as distinct species or even genera. The hypothesis of current or very recent large‐scale gene flow among seeps and vents was supported by microsatellite analysis of the shrimp species Alvinocaris muricola/Alvinocaris markensis across the AEB and MAR. Two nonmutually exclusive hypotheses may explain these findings. The dispersion of larvae or adults following strong deep‐sea currents, possibly combined with biochemical cues influencing the duration of larval development and timing of metamorphosis, may result in large‐scale effective migration among distant spots scattered on the oceanic seafloor. Alternatively, these results may arise from the prevailing lack of knowledge on the ocean seabed, apart from emblematic ecosystems (chemosynthetic ecosystems, coral reefs or seamounts), where the widespread classification of endemism associated with many chemosynthetic taxa might hide wider distributions in overlooked parts of the deep sea.