Void development on carbonate coasts: creation of anchialine habitats

Anchialine caves in coastal locations develop in two ways: by pseudokarst processes that form talus caves, sea caves, tafoni, fissure caves and lava tubes, and by karst dissolutional processes that form stream caves, flank margin caves, and blue holes. Pseudokarst caves are of minor importance in anchialine cave habitat development, with some lava tubes being notable exceptions. Dissolution caves provide the most extensive, variable, and long-term environments for anchialine habitats. The Carbonate Island Karst Model (CIKM) allows dissolutional cave development in carbonate coasts to be understood as the interplay between freshwater and marine water mixing, sea-level change, rock maturity, and interaction with adjacent non-carbonate rocks. Glacioeustatic sea-level changes of the Quaternary have moved all coastal anchialine cave environments repeatedly through a vertical range of over 100 m, and modern anchialine environments could not develop at their current elevations until ~4,000 years ago when sea level reached its present position. Blue holes form by a variety of mechanisms, but the most common is upward stoping and collapse from deep dissolutional voids. As a result, they provide vertical connection between different levels of horizontal cave development produced by a variety of earlier sea-level positions. Blue holes are overprinted by successive sea-level fluctuations; each sea-level event adds complexity to the habitats within blue holes and the cave systems they connect. Blue holes can reach depths below the deepest glacioeustatic sea-level lowstand, and thereby provide a refugia for anchialine species when cave passages above are drained by Quaternary sea-level fall. Blue holes represent the most significant anchialine cave environment in the world, and may provide clues to anchialine cave species colonization and speciation events.     

Nutrient speciation and hydrography in two anchialine caves in Croatia: tools to understand iodine speciation

Despite iodine being one of the most abundant of the minor elements in oxic seawater, the principal processes controlling its interconversion from iodate to iodide and vice versa, are still either elusive or largely unknown. The two major hypotheses for iodate reduction involve either phytoplankton growth in primary production, or bacteria during regeneration. An earlier study intended to exploit the unusual nature of anchialine environments revealed that iodide is oxidised to iodate in the bottom of such caves, whereas reduction of iodate occurs in the shallower parts of the water column. This investigation was made on the hypothesis that study of the nitrogen and phosphorus nutrient systems within the caves might offer a bridge between the iodine chemistry and the marine bacteria which are assumed to be the agent of change of the iodine in the caves. Accordingly, the hydrography, the nutrient chemistry, and some further iodine studies were made of two anchialine caves on the east coast of the Adriatic Sea in Croatia. Iodate and iodide were determined by differential pulse voltammetry and cathodic stripping square-wave voltammetry, respectively. Total iodine was determined indirectly, as iodate, after oxidation of reduced iodine species with UV irradiation and strong chemical oxidants. Nutrient concentrations were measured by spectrophotometry. Nutrient profiles within the well stratified water columns indicate a relatively short-lived surface source of nitrate and phosphate to the caves, with a more conventional, mid-water, nutrient regeneration system. The latter involves nitrite and ammonium at the bottom of the halocline, suggestive of both autotrophic and heterotrophic microbial activity. High iodate/low iodide deep water, and conservative behaviour of total inorganic iodine were confirmed in both systems. Iodate is reduced to iodide in the hypoxic region where nutrient regeneration occurs. The concentrations of organic iodine were surprisingly high in both systems, generally increasing toward the surface, where it comprised almost 80% of total iodine. As with alkalinity and silica, the results suggest that this refractive iodine component is liberated during dissolution of the surrounding karst rock. A major, natural flushing of one of the caves with fresh water was confirmed, showing that the cave systems offer the opportunity to re-start investigations periodically.     

Mitochondrial OXPHOS genes provides insights into genetics basis of hypoxia adaptation in anchialine cave shrimps

Cave shrimps from the genera Typhlatya, Stygiocaris and Typhlopatsa (TST complex) comprises twenty cave-adapted taxa, which mainly occur in the anchialine environment. Anchialine habitats may undergo drastic environmental fluctuations, including spatial and temporal changes in salinity, temperature, and dissolved oxygen content. Previous studies of crustaceans from anchialine caves suggest that they have possessed morphological, behavioral, and physiological adaptations to cope with the extreme conditions, similar to other cave-dwelling crustaceans. However, the genetic basis has not been thoroughly explored in crustaceans from anchialine habitats, which can experience hypoxic regimes. To test whether the TST shrimp-complex hypoxia adaptations matched adaptive evolution of mitochondrial OXPHOS genes. The 13 OXPHOS genes from mitochondrial genomes of 98 shrimps and 1 outgroup were examined. For each of these genes was investigated and compared to orthologous sequences using both gene (i.e. branch-site and Datamonkey) and protein (i.e. TreeSAAP) level approaches. Positive selection was detected in 11 of the 13 candidate genes, and the radical amino acid changes sites scattered throughout the entire TST complex phylogeny. Additionally, a series of parallel/convergent amino acid substitutions were identified in mitochondrial OXPHOS genes of TST complex shrimps, which reflect functional convergence or similar genetic mechanisms of cave adaptation. The extensive occurrence of positive selection is suggestive of their essential role in adaptation to hypoxic anchialine environment, and further implying that TST complex shrimps might have acquired a finely capacity for energy metabolism. These results provided some new insights into the genetic basis of anchialine hypoxia adaptation.     

Genetic exchange between anchialine cave populations by means of larval dispersal: the case of a new gastropod species Neritilia cavernicola

Despite being limited to caves, many anchialine taxa have disjunct insular distributions, which raises questions about their origins and colonization history. This study deals with the new gastropod Neritilia cavernicola sp. n. (Neritopsina: Neritiliidae) from anchialine caves on two islands in the Philippines that are separated by the deep Bohol Strait and situated 200 km apart along the coastline of Cebu Island. Neritilia cavernicola is an obligate stygobiont and most closely resembles Neritilia littoralis , which lives in interstitial waters of the Nansei-shoto Islands, Japan. Its eggs and larval shells are identical to those of other Neritilia species, despite their different adult habitats. This suggests a marine planktotrophic phase (as occurs in amphidromous riverine species of Neritilia ), and consequent migration between islands via ocean currents. Here we present the first genetic structure for anchialine cave organisms; comparisons of 1276 bp sequences from mitochondrial cytochrome oxidase I show no evidence of genetic isolation between the islands. All individuals evidently are part of a panmictic population and the low vagility of adults and their seemingly isolated cave habitats do not limit gene flow in N. cavernicola . This migration model, based on marine larval dispersal, may be widely applicable to anchialine stygobites with insular distributions, as many such organisms (including shrimps, crabs and fishes) are phylogenetically allied to amphidromous species.     

Phytoplankton of cenotes and anchialine caves along a distance gradient from the northeastern coast of Quintana Roo,Yucatan Peninsula

This work details the taxonomic composition of suspended algae (phytoplankton and tycoplankton) communities in five cenotes (sinkholes) and two anchialine caves in northeastern Quintana Roo, Mexico. The sample set of cenotes are Casa, Nohoch Nah Chich, Maya Blue, Cristal, and Carwash, as well as the two associated caves leading from the cenotes of Maya Blue and Cristal. The site distribution represents a distance gradient with respect to the coastline with which we observe the effects of tidal movement and the mixing of waters (e.g. saline water and freshwater) on the composition of the suspended algae communities. Two sample sets were taken, one at the end of the dry season (March–April 1995) and the second at the end of the rainy season (September–October 1995) with the goal of comparing the contrasting climatic conditions of the region. A total of 79 species were identified, of which, diatoms were the most important with respect to species richness with a total of 75% of species. The floristic composition is very similar between the freshwater cenotes. The distance of a cenote site with respect to the coastline was a determining factor in the species composition. Casa Cenote is the most distinct of the sample set for the presence of marine species due to its proximity to the coastline. The tides are a large determining factor of the floristic composition of Casa Cenote with 24% all species identified in this study found exclusively in this system. The anchialine system species are transported from the cenotes and the adjacent cave systems. The largest percentage or species (95%) are freshwater, and only 5% of the total number of identified species are of marine origin. It is recognized that the most distant cenotes from the coast, Carwash and Cristal, as well as Maya Blue and Nohoch Nah Chich, are the most similar, despite being part of different cave systems. In these inland systems the marine species decreased drastically (2.4% in Nohoch Nah Chich and no marine species in the remaining cenotes). Marine species are found at the halocline of the caves.     

Breeding sites of Phlebotomus sergenti, the sand fly vector of cutaneous leishmaniasis in the Judean Desert

Phlebotomine sand flies transmit Leishmania, phlebo-viruses and Bartonella to humans. A prominent gap in our knowledge of sand fly biology remains the ecology of their immature stages. Sand flies, unlike mosquitoes do not breed in water and only small numbers of larvae have been recovered from diverse habitats that provide stable temperatures, high humidity and decaying organic matter. We describe studies designed to identify and characterize sand fly breeding habitats in a Judean Desert focus of cutaneous leishmaniasis. To detect breeding habitats we constructed emergence traps comprising sand fly-proof netting covering defined areas or cave openings. Large size horizontal sticky traps within the confined spaces were used to trap the sand flies. Newly eclosed male sand flies were identified based on their un-rotated genitalia. Cumulative results show that Phlebotomus sergenti the vector of Leishmania tropica rests and breeds inside caves that are also home to rock hyraxes (the reservoir hosts of L. tropica) and several rodent species. Emerging sand flies were also trapped outside covered caves, probably arriving from other caves or from smaller, concealed cracks in the rocky ledges close by. Man-made support walls constructed with large boulders were also identified as breeding habitats for Ph. sergenti albeit less important than caves. Soil samples obtained from caves and burrows were rich in organic matter and salt content. In this study we developed and put into practice a generalized experimental scheme for identifying sand fly breeding habitats and for assessing the quantities of flies that emerge from them. An improved understanding of sand fly larval ecology should facilitate the implementation of effective control strategies of sand fly vectors of Leishmania.     

Global biodiversity and phylogenetic evaluation of remipedia (crustacea)

Remipedia is one of the most recently discovered classes of crustaceans, first described in 1981 from anchialine caves in the Bahamas Archipelago. The class is divided into the order Enantiopoda, represented by two fossil species, and Nectiopoda, which contains all known extant remipedes. Since their discovery, the number of nectiopodan species has increased to 24, half of which were described during the last decade. Nectiopoda exhibit a disjunct global distribution pattern, with the highest abundance and diversity in the Caribbean region, and isolated species in the Canary Islands and in Western Australia. Our review of Remipedia provides an overview of their ecological characteristics, including a detailed list of all anchialine marine caves, from which species have been recorded. We discuss alternative hypotheses of the phylogenetic position of Remipedia within Arthropoda, and present first results of an ongoing molecular-phylogenetic analysis that do not support the monophyly of several nectiopodan taxa. We believe that a taxonomic revision of Remipedia is absolutely essential, and that a comprehensive revision should include a reappraisal of the fossil record.     

Recently discovered landlocked basins in Indonesia reveal high habitat diversity in anchialine systems

In this article, the variability of physical settings of anchialine systems in Indonesia is discussed together with the consequences these settings have for the environment and biota within the systems. Exploration in two karstic areas (Berau, East Kalimantan and Raja Ampat, West Papua) has resulted in the discovery of 20 previously unknown anchialine systems in Indonesia. Based on parameters such as bathymetry, size, coastline, salinity, water temperature, pH, degree of connection to the sea, and the presence-absence of selected key taxa we distinguish three types of (non-cave) anchialine systems in the Indo-Pacific: (1) Marine lakes with large and deep basins containing brackish to almost fully marine waters. Marine lakes show a range in the degree of connection to the sea with the result that the higher the connection the more the lake resembles a lagoon in both water chemistry and biota, while the more isolated lakes have brackish water and contain unique species that are rarely found in the adjacent sea. (2) Anchialine pools with small and shallow basins containing brackish water and low diversity of macrofauna. (3) Blue pools in chasms that contain water with a clear halocline and are possibly connected to anchialine caves. Study of the many unique features of anchialine systems will enhance our understanding of the physical and ecological processes responsible for diversification in tropical shallow marine environments.     

Evolution of cave suspension feeding in Protodrilidae (Annelida)

Protodrilidae belongs in a lineage that until now entirely consisted of deposit‐feeding, highly adapted interstitial annelids. Except for a pair of anterior palps, all protodrilids lack appendages, parapodia and chaetae; and have slender bodies adapted to glide between the sand grains by ciliary motion. The first exception to these characteristics is Megadrilus pelagicus n. sp. inhabiting the water column of the anchialine La Corona cave system in Lanzarote. Its morphology and evolutionary history are here investigated by combining observations from in vivo video recordings and advanced microscopy with phylogenetic analyses. Our studies revealed a unique pelagic, suspension feeding behaviour attained by its long ciliated palps in combination with an autopomorphic dorsal ciliated keel and several longitudinal and transverse ciliary bands. Phylogenetic analyses recovered Megadrilus pelagicus n. sp. nested within Protodrilidae indicating that its unique traits are derived within the family. These traits are traced in the tree topologies in correlation to cave colonization. The evolution of these traits can be functionally explained by the different demands of a pelagic suspension feeding strategy compared to the ancestral deposit‐feeding guild of the family. The origin of this suspension feeding strategy was presumably favoured by the partial isolation of the anchialine ecosystem, connected to the sea only through the highly porous volcanic subterranean bedrock. This crevicular connection limits the amount of predators and turbulence in the cave, but allows continuous water flow into the system carrying organic particles, which is the main source of food when photosynthetic primary production does not occur and sedimentation is limited. These conditions may select for pelagic suspension feeding as the most feasible life‐strategy in anchialine caves, which the dominance of pelagic, suspension feeding crustaceans and annelids in anchialine cave assemblages may also reflect. For species of ancestrally deposit‐feeding lineages entering the cave system, such as the annelid families Protodrilidae and Nerillidae, an adaptive‐shift from interstitial to crevicular habitats seemingly correlates with dramatic morphological changes and speciation. The dramatic changes observed in these primarily interstitial lineages compared to their relatives, point to alternative adaptive evolutionary pathways related to ecological fitness contrary to the previously proposed theories focusing on geological or stochastic processes.     

Combining otolith chemistry and telemetry to assess diadromous migration in pinkeye mullet, <Emphasis Type="Italic">Trachystoma petardi</Emphasis> (Actinopterygii,Mugiliformes)

Little is currently known regarding microbial community structure, and the environmental factors influencing it, within the anchialine ecosystem, defined as near-shore, land-locked water bodies with subsurface connections to the ocean and groundwater aquifer. The Hawaiian Archipelago is home to numerous anchialine habitats, with some on the islands of Maui and Hawaii harboring unique, laminated orange cyanobacterial–bacterial crusts that independently assembled in relatively young basalt fields. Here, benthic and water column bacterial and micro-eukaryotic communities from nine anchialine habitats on Oahu, Maui, and Hawaii were surveyed using high-throughput amplicon sequencing of the V6 (Bacteria-specific) and V9 (Eukarya-biased) hypervariable regions of the 16S- and 18S-rDNA genes, respectively. While benthic communities from habitats with cyanobacterial–bacterial crusts were more similar to each other than to ones lacking it on the same island, each habitat had distinct benthic and water column microbial communities. Analyses of the survey data in the context of environmental factors identified salinity, site, aquifer, and watershed as having the highest explanatory power for the observed variation in microbial diversity and community structure, with lesser drivers being annual rainfall, longitude, ammonium, and dissolved organic carbon. Our results epitomize the abiotic and biotic uniqueness characteristic of individual habitats comprising the Hawaiian anchialine ecosystem.     

Anchialine Gyprididae (Ostracoda) from the Galapagos Islands, with a review of the subfamily Paracypridinae

The five species of Cyprididae collected from anchialine habitats of the Galapagos Islands include Paracypris crispa, previously described from marine caves and reefs on Bermuda, and the freshwater species Strandesia stocki, which is widely distributed in wells of the West Indies. Three new species, Dolerocypria ensigera, Mungava recta and Hansacypris galapagosensis, contribute to more precise understanding of these circumtropical genera. A review of known soft-anatomical characters for 68 species of Paracypridinae, five of which are transferred to different genera, reveals too many inconsistencies for a simple hypothesis. The current classification of Paracypridinae into three tribes and 18 genera may be overly subdivided, and elevation of Paracypridinae to family rank is not warranted.     

贵州董背洞和水江洞内动物群落结构与部分环境因子的相关性

在2002—2005年的2月和7月共5次赴荔波董背洞和水江洞对肉眼能见到的软体动物、节肢动物和脊索动物进行了观察和采集,在董背洞共获标本440号,隶属3门5纲10目20科39种或类群;在水江洞共获标本498号,隶属3门6纲11目20科25种或类群。根据上述两洞内各光带中动物种类和数量组成不同,将其划分为6个动物群落,经群落多样性分析,物种丰富度、群落多样性、最大多样性、均匀度、优势度和相似性指数最高的分别是群落B(4·1059)、H(2·4716)、B(3·3322)、E(0·9042)、C(0·3442)和A—C(0·5251)。此外还研究了群落多样性与部分环境因子的相关性,如温度、湿度、空气中CO2和N2含量、土壤有机质和部分无机盐含量等。用Pearson相关系数进行分析,结果显示土壤中有机质的含量与物种数、物种丰富度和群落最大多样性指数都呈极显著正相关,相关系数分别为0·885、0·909和0·868(双尾显著性检验均P≤0·05),与群落多样性指数呈显著正相关,相关系数为0·611(双尾显著性检验,P≥0·1),由此证明土壤有机质的含量是影响洞穴动物群落变化的重要因子之一;在地表,温、湿度是影响动物群落多样性变化的重要因子,但在特殊的溶洞内由于洞尾段的温度四季较稳定和整个洞穴内的湿度均较高,故与群落多样性的相关性不显著。     

Search for Bermuda’s deep water caves

The mid-Atlantic islands of Bermuda harbor one of the richest and most diverse anchialine communities known from anywhere on Earth. However, all known anchialine caves in Bermuda (maximum depth—26 m) were dry during the last glacial period extending from approximately 9,000 to 115,000 years ago when glacial sea levels were as much as 127 m lower. Since it is highly unlikely that Bermuda’s endemic cave species evolved since the caves were flooded by sea level rise, alternate deeper habitats must have existed to shelter anchiane fauna for prolonged periods of lower sea level during the Pleistocene. In order to systematically search for such now deep water cave habitats, high-resolution multibeam sonar and remotely operated vehicles were used to map and explore the seafloor off Bermuda in 60–200 m depths along the outer shelf break edge of the submarine escarpment surrounding the Bermuda Platform and an adjacent seamount. Specific goals were to discover deep water cave and/or crevicular habitats and to characterize the nature, geological stratification and composition, and sea level history of the platform margin, in particular focusing on features directly relating to Pleistocene low sea stand events. During this sea floor survey, clearly defined paleo-shoreline features generated by wave and current erosion were found to encircle the Bermuda seamount and Challenger Bank at 60 and 120 m depths.     

Milyeringa Veritas (Eleotridae), a remarkably versatile Cave Fish from the Arid Tropics of Northwestern Australia

The blind cave gudgeon Milyeringa veritas is restricted to groundwaters of Cape Range and Barrow Island, northwestern Australia. It occurs in freshwater caves and in seawater in anchialine systems. It is associated with the only other stygobitic cave vertebrate in Australia, the blind cave eel, Ophisternon candidum, the world's longest cave fish, and a diverse stygofauna comprising lineages with tethyan tracks and widely disjunct distributions, often from North Atlantic caves. The cave gudgeon inhabits a karst wetland developed in Miocene limestones in an arid area. There is an almost complete lack of information on the basic biology of this cave fish, despite it being listed as threatened under the Western Australian Wildlife Conservation Act. Allozyme frequencies and distributions indicate significant population sub-structuring on the Cape Range peninsula such that the populations are essentially isolated genetically suggesting that more than one biological species is present. Further, they suggest that the vicariant events may have been associated with a series of eustatic low sealevels. Analysis of intestinal contents indicates that they are opportunistic feeders, preying on stygofauna and accidentals trapped in the water, at least at the sites sampled which were open to the surface, a conclusion supported by the results of stable isotope ratio analysis. The gudgeons are found in freshwater caves and throughout deep anchialine systems in which they occur in vertically stratified water columns in which there is a polymodal distribution of water chemistries (temperature, pH, salinity, dissolved oxygen, redox, dissolved inorganic nitrogen series, hydrogen sulphide).     

Rocks and clocks: linking geologic history and rates of genetic differentiation in anchialine organisms

The geologic history of a region can significantly impact the development of its flora and fauna, with past events shaping community patterns and evolutionary trajectories of species. In this context, islands are excellent “natural laboratories” for studying the fundamental processes of evolution due to their discrete geographical nature and dynamic geologic histories. An island system meeting these criteria is the Hawaiian Archipelago, which is ideal for testing how island geologic history influences the processes leading to population genetic variation and differentiation. One Hawaiian endemic whose evolutionary history is closely tied to the geology of the islands is the anchialine atyid shrimp Halocaridina, whose mitochondrial cytochrome oxidase I (COI) gene is hypothesized to be evolving at the rate of 20% per million years. To validate this rapid evolutionary rate, time since divergence estimates between geographically close, yet genetically distinct, populations were calculated for Halocaridina from anchialine habitats on the islands of Hawai’i, Maui, and O’ahu. On the younger (i.e., <1.5 million years) islands of Hawai’i and Maui, where all anchialine habitats occur in basalt, application of the Halocaridina molecular clock identified a strong correlation between levels of genetic divergence and the geologic age of the region inhabited by those populations. In contrast, this relationship weakened when similar analyses were conducted for Halocaridina from limestone anchialine habitats on the older (i.e., >2.75 million years) island of O’ahu. These results suggest geologic age, basin origin and/or composition are important factors that should be taken into consideration when conducting molecular clock analyses on anchialine flora and fauna as well as island populations in general.     

Diurnal dynamics of the microbial loop in peatlands: structure, function and relationship to environmental parameters

Globally, introductions of alien species are increasingly common, with invasive predators potentially having detrimental effects via predation on native species. However, native prey may avoid predation by adopting new behaviors. To determine whether invasive fish populations consume endemic shrimp in invaded Hawaiian anchialine habitats or if adopted patterns of diel migration prevents predation as previously hypothesized, a total of 183 invasive poeciliids (158 Gambusia affinis and 25 Poecilia reticulata) were collected for gut content analyses from four anchialine sites during wet and dry seasons on the islands of Hawai‘i and Maui. Predation on shrimp was not detected in habitats where they retreat exclusively into the underlying aquifer diurnally and only emerge nocturnally. However, low levels of predation were detected (7/65 fishes, only by Gambusia affinis) at Waianapanapa Cave, Maui, where shrimp retreat into both the aquifer and a cave during the day. Thus, adopted behavioral responses to invasive fishes generally, though not universally, prevent predation on endemic Hawaiian anchialine shrimps. However, non-consumptive effects resulting from behavioral modification of shrimps may have appreciable impacts on the Hawaiian anchialine ecosystem and warrant further study.     

Distributional dynamics of a specialized subterranean community oppose the classical understanding of the preferred subterranean habitats

Troglobionts are organisms that are specialized for living in a subterranean environment. These organisms reside prevalently in the deepest zones of caves and in shallow subterranean habitats, and complete their entire life cycles therein. Because troglobionts in most caves depend on organic matter resources from the surface, we hypothesized that they would also select the sections of caves nearest the surface, as long as environmental conditions were favorable. Over 1 year, we analyzed, in monthly intervals, the annual distributional dynamics of a subterranean community consisting of 17 troglobiont species, in relation to multiple environmental factors. Cumulative standardized annual species richness and diversity clearly indicated the existence of two ecotones within the cave: between soil and shallow subterranean habitats, inhabited by soil and shallow troglobionts; and between the transition and inner cave zones, where the spatial niches of shallow and deep troglobionts overlap. The mean standardized annual species richness and diversity showed inverse relationships, but both contributed to a better insight into the dynamics of subterranean fauna. Regression analyses revealed that temperatures in the range 7–10°C, high moisture content of substrate, large cross section of the cave, and high pH of substrate were the most important ecological drivers governing the spatiotemporal dynamics of troglobionts. Overall, this study shows general trends in the annual distributional dynamics of troglobionts in shallow caves and reveals that the distribution patterns of troglobionts within subterranean habitats may be more complex than commonly assumed.     

Micropacteridae, a new family of Remipedia (Crustacea) from the Turks and Caicos Islands

Recent diving explorations of anchialine caves on the Turks and Caicos Islands yielded a rather small and slender new species of Remipedia. Micropacter yagerae n. gen., n. sp. is distinguished from all other species of nectiopod remipedes by a number of autapomorphic characters, including an oval body terminus with fused segments, unequal pairs of terminal claws on maxilla and maxilliped, an almost complete reduction of sternal bars and pleurotergites, molar processes with relatively few, but strong spines, and frontal filaments with bifurcate processes. Based on the unique combination of derived and primitive characters, we propose to erect a new family, Micropacteridae, for this new species and genus of Remipedia. Taxonomic diagnoses for the class Remipedia, order Nectiopoda (emended due to discovery that the maxilliped is 9-segmented), and for the families Speleonectidae and Godzilliidae are presented and discussed.     

Evolution of cave Axiokebuita and Speleobregma (Scalibregmatidae,Annelida)

The evolutionary history of Axiokebuita and Speleobregma, two poorly known lineages of annelids exclusive from deep‐sea or marine caves but always from crevicular habitats, is explored here. Speleobregma lanzaroteum Bertelsen, 1986, and Axiokebuita cavernicola sp. n. are described from anchialine and marine caves of the Canary Islands using light and electron microscopy. Speleobregma lanzaroteum is previously known only from a single specimen from the water column of an anchialine cave in Lanzarote. Emended diagnosis, details on the ciliary patterns and behavioural observations are provided based on newly collected material and in situ observations. Axiokebuita cavernicola sp. n. is found in Pleistocene gravel deposits in a shallow water marine cave in Tenerife (Canary Islands). The new species is characterized by the presence of dorsal ciliary bands and short knob‐like neuropodial cirri from segment two. The porosity and permeability of the gravelly environment of A. cavernicola sp. n. are shown to be equivalent to the water column or crevices of Speleobregma and other Axiokebuita spp. Phylogenetic analyses of five gene fragments and 44 terminals using maximum‐likelihood and Bayesian methods support a derived position of A. cavernicola sp. n. within Axiokebuita and confirm a sister‐group relationship of Axiokebuita with Speleobregma with high nodal support. The Axiokebuita–Speleobregma clade is morphologically characterized by a globular pygidium with adhesive glands and ventral ungrooved ciliated palps. Our results support two independent cave colonization events, favoured by the preadaptation of the members of Axiokebuita–Speleobregma lineage to crevicular habitats.