Do Mediterranean fish assemblages associated with marine caves and rocky cliffs differ?

Fish assemblages associated with marine caves and rocky cliffs were investigated in the Salento Peninsula (SE Italy, Mediterranean Sea) by using visual census methods. Sampling was done at three sites, each of which included 4 habitat types: the external and the internal portions of caves, and shallow and deep rocky cliffs. 10 and 13 species were found exclusively inside cave habitats (e.g. Corcyrogobius lichtensteini, Thorogobius ephippiatus and Grammonus ater) or in rocky cliffs (e.g. Diplodus annularis, Sarpa salpa, Sparisoma cretense, Spondyliosoma cantharus), respectively. The four habitat types shared 10 species, and the external portions of the caves shared the most species (both with the internal cave portions and the external rocky cliffs). As a general rule, dissimilarity in the fish assemblage structure between habitats was far greater than dissimilarity between sites. Apogon imberbis (mostly associated with caves) and Chromis chromis (typifying rocky cliffs, mainly the deep ones) mostly contributed to dissimilarities between caves and rocky cliffs. Apogon imberbis (mostly associated with internal caves) and Coris julis (mainly associated to external cave portions) contributed strongly to dissimilarities between internal and external cave portions, while C. chromis, Symphodus mediterraneus and C. julis (associated with the deeper cliffs) and Thalassoma pavo (mostly present in shallow cliffs) differentiated deep and shallow cliffs. Diplodus vulgaris, Oblada melanura and Mullus surmuletus showed a marked increase in density during the cold season in the caves. These results show that fish assemblages associated with rocky reefs rich in marine caves (in terms of relative densities, species composition, species richness, exclusive species and presence of juveniles of some valuable species) may be affected by the peculiar ecological conditions within caves, which could provide additional resources for fishes (e.g. food availability, refuge against predators, sand patches within a rocky matrix) compared to rocky reefs without caves. These results suggest that stretches of rocky coasts rich in marine caves should be considered within management/conservation programs (e.g. when establishing Marine Protected Areas).     

Unexpected records of ‘deep‐sea’ carnivorous sponges Asbestopluma hypogea in the shallow NE Atlantic shed light on new conservation issues

Marine cave communities have been a continued source of ecological surprises, among other things because of their close ecological and evolutionary ties with the deep sea. The discovery of cladorhizid sponges, the deepest occurring poriferan family, in shallow Mediterranean caves in the 1990s was one such surprise, leading to the generally accepted hypothesis that the whole family was carnivorous, an unprecedented feeding mode for sponges. The recent observation of the cave species Asbestopluma hypogea in the Mediterranean bathyal, confirmed the view that some shallow caves can occasionally shelter otherwise deep‐dwelling species. Here we present new distribution data of A. hypogea, from deep Mediterranean locations, and for the first time from Atlantic locations. Among the new Atlantic records, the most surprising ones are located in three different geographic areas (Ria de Arousa, Groix Island and Cherbourg) of the NW European coasts, from the Iberian Peninsula to the English Channel, where A. hypogea reaches SCUBA depths (5–50 m), while not sheltered in marine caves. The carnivorous sponge however reaches its shallowest occurrence (5 m), in a small cave at Groix Island. The ecological significance of these discoveries, particularly the very patchy distribution and peculiar dynamics, are noteworthy, and the shallow occurrence of A. hypogea, together with other deep‐water or uncommon species, constitute unique assemblages that must be considered in conservation plans.     

Occurrence and distribution pattern of Palaemon spp. shrimps in a shallow submarine cave environment: a study case in South‐eastern Italy

The common shrimps of the genus Palaemon are often reported as living in Mediterranean submarine caves. The present study quantifies the distribution pattern of a Palaemon spp. populations throughout 1 year of observations in a shallow submarine cave in South‐eastern Italy, Ionian Sea. The cave was subdivided into six sectors from the entrance towards the inner portion of the ‘blind cave’ and 16 dives (four per season, during both day and night) were performed with SCUBA equipment. The population was estimated in situ using visual census. Four different species of Palaemon were identified. This study has shown that Palaemon spp. assemblages live throughout the year in submarine caves. Moreover, the results suggest submarine caves play an important role in the biology of Palaemon spp. populations, by providing refuge, juvenile recruitment and food resources.     

Colonisers of the dark: biostalactite‐associated metazoans from “lu Lampiùne” submarine cave (Apulia,Mediterranean Sea)

Metazoan/microbial bioconstructions, or biostalactites (BSTs), discovered in submarine caves of Apulia c. 20 years ago—and later found in several shallow‐water Mediterranean caves—are receiving increasing attention in the last years. Examination of a single BST from the “lu Lampiùne” cave (Apulia), at the limit between the Adriatic and the Ionian seas, has been addressed for the first time in this paper. The BST started growing at c. 6,000 years ago with a rapid accretion of large‐sized serpulids (Protula), slowing down since about 3,000–4,000 years ago with a shift in main bioconstructors, probably caused by environmental changes. The present‐day community on the outer BST surface is dominated by skeletonised epibionts, mostly small‐sized serpulids, bryozoans and foraminifers, which contribute carbonate to the BST growth, by encrusting sponges, and by a few endobionts, including boring bivalves and insinuating sponges. New data remarkably increase biodiversity known for the “lu Lampiùne” cave and the cave habitat in the region. Thirty‐five taxa (16 bryozoans, 10 serpulids, 3 brachiopods, 2 foraminifers, 2 sponges, 1 bivalve and 1 cirriped) are recorded for the first time from Apulian marine caves, highlighting the need for further research in the area. In addition, the BST‐associated community seems to differ from those of individual BSTs from other Mediterranean caves, revealing the individuality of these communities.     

舟山海域外来物种地中海贻贝的自然分布现状及生态影响

以筛选的2个可鉴别的地中海贻贝（Mytilus galloprovincialis）、厚壳贻贝（M.coruscus）及其杂交后代的PCR标记（扩增核基因Glu-5’特异片段PCR标记Me14/Me17和延伸因子1α第一个内含子区段PCR标记引物EFbisF/EFbisR）,对舟山海域3个贻贝主产区（马鞍列岛海域、浪岗山列岛海域、中街山列岛海域）采集的贻贝样本的种类进行鉴别.研究结果显示：100个马鞍列岛海域贻贝个体和30个中街山列岛海域贻贝个体均为厚壳贻贝;而54个浪岗山列岛海域贻贝样本中,16个个体（29.6%）为厚壳贻贝,38个个体（70.4%）为地中海贻贝;3个群体中均未检测到2种贻贝的杂交个体.因此,推测舟山海域地中海贻贝自然分布仅局限于局部海区,但在适宜的海区地中海贻贝可取代厚壳贻贝成为贻贝床的优势物种,舟山海域贻贝床的动态变化值得进一步关注;贻贝样本中没有检测到地中海贻贝与厚壳贻贝的杂交个体,表明调查海域可能没有杂交贻贝分布或数量极少,地中海贻贝通过杂交、基因渐渗污染厚壳贻贝基因组的风险较小.     

Adriatic ‘opisthobranchs’ (Gastropoda,Heterobranchia): shedding light on biodiversity issues

We herein review the Adriatic opisthobranch fauna, provide an updated checklist of 223 species and assess their distribution at regional and country levels. New Adriatic records are provided for 67 opisthobranch taxa, adding three new records for the Italian coastline, five new records for Albania, eight for Croatia and 15 for Montenegro. The presence of Hermaea bifida (Montagu, 1815), Hermaea variopicta (Costa A., 1869) and Facelina annulicornis (Chamisso & Eysenhardt, 1821) is reported for the first time from anywhere in the Adriatic Sea. Including the new findings, our inventory includes 28 species from Albania, 163 from Croatia, 178 from the Italian coastline of the Adriatic, 41 from Montenegro and 74 from Slovenia. No records were available from Bosnia and Hercegovina. Ninety species (40.4%) are widespread, and were recorded from all three main divisions (Northern, Western and Eastern Adriatic), whilst 79 species (35.4%) were reported from only one of them. At sub‐division levels, the Albanian inventory is the most dissimilar to the other country/regional lists, presumably because of the lack of targeted field surveys. The highest similarity is observed amongst the Western Adriatic, Croatia and the Italian Ionian coastline. Seven alien, one possible alien and three cryptogenic opisthobranchs were reliably recorded from the Adriatic Sea. The first Mediterranean records of five species were in the Adriatic Sea, of which four species have not been found elsewhere in the Mediterranean. Shipping and aquaculture are the probable pathways for most of the alien species. A steady rate of one to two alien introductions per decade since the 1970s was evident. The Northern Adriatic coasts are the most affected by alien invasions, probably due to repeated local introductions along with shellfish transfers (oysters, mussels, Manila clams), and heavy maritime transport.     

Metazoan/microbial biostalactites from present‐day submarine caves in the Mediterranean Sea

Biostalactites formed by metazoan–microbialite associations from three submerged marine caves in the Plemmirio Peninsula (south of Syracuse, Ionian Sea) are randomly distributed and show different sizes and morphologies, as well as variations in surface roughness/smoothness. The biostalactites consist of crusts a few centimeters thick of small serpulids and other metazoans, associated with fine‐grained carbonate; the larger ones often include a nucleus of serpulid tubes (Protula). The metazoans include mainly serpuloideans, sponges, bryozoans and foraminifers but microbial carbonates are also significant components. The composition of both the living communities and thanatocoenoses on the outer surfaces, as well as the composition and fabric of the internal framework, were analysed and used to reconstruct the history of the caves. All of the identified sessile faunas mainly consist of cryptic and sciaphilic dwellers that reflect cave conditions and their variations through time. The distribution pattern, composition and abundance of the present‐day dwellers largely depend on the degree of roughness of the biostalactite surfaces and their positions within the caves. It has been suggested that the Protula specimens in the nuclei represent pioneer populations that formed aggregates during the early cave colonization phase, in response to relatively high food supply from seawater inflow and intruding continental waters. By contrast, the outer metazoan–microbialite carbonates reflect more confined conditions in the caves caused by Holocene sea‐level rises. Hypotheses are proposed for biostalactite growth, taking into account information about the growth rates of some constituents, and evidence of dissolution effects. Similarities and differences between these biostalactites and other Holocene deposits previously described from submarine caves in the Mediterranean Sea and in tropical reefs are discussed.     

Back to the future: reflections and directions of South African marine bioinvasion research

Biological invasions continue to increase around the world, with impacts on many coastal marine systems. Here we review the South African marine invasion literature which, despite the field being relatively new, has grown to have significant presence in both the local and international arenas. Of the 79 papers reviewed, 70% focused on the establishment and spread of alien species, with modes of transport and introduction largely overlooked. An emphasis was also apparent towards field studies, in particular survey work, with few experimental studies. The overwhelming majority of papers focused on a single species, the Mediterranean mussel Mytilus galloprovincialis, reflecting the scale of this invasion and the tractable nature of rocky shores as study systems. With the exception of this one species, the impacts of marine alien species have rarely been quantified. We suggest that future research extends the taxonomic coverage of present work and develops a better understanding of the mechanisms of introduction, establishment and spread of marine alien species. Through an experimental approach, the drivers of altered ecological patterns and processes resulting from invasions should be addressed, providing insight into associated impacts. This approach will maintain the local applicability and international relevance of South African marine invasion research.     

Molecular and distribution data on the poorly known,elusive, cave mysid Harmelinella mariannae (Crustacea: Mysida)

Mediterranean underwater marine caves harbour abundant populations of several species of mysids that are increasingly used as biological models in ecological and evolutionary studies. One exception is the species Harmelinella mariannae, described in 1989 and then hardly ever again reported in the literature. We here provide the first data on the distribution of this poorly known taxon that, contrary to expectations for a rare brooding cave dweller, we now report from Madeira Island in the nearby Atlantic, to the easternmost parts of the Mediterranean. Brief behavioural observations are added, particularly its atypical solitary habits and its feeding behaviour as a high trophic level carnivore. Molecular characterization of the different specimens captured provided three sorts of information. Mitochondrial COI and 16S haplotypes suggest different colonization waves in the Mediterranean, with one group in the Eastern Basin, two in the Marseille region in the NW part of this sea, and another group with a very wide extension from Madeira to Liguria and Malta. Mitochondrial data also support that one of the groups in Marseille might have diverged as a cryptic species of Harmelinella. 18S rRNA gene displays a single common sequence to all specimens from the four groups, and seems to confirm the original proposed placement of this taxon within the subfamily Heteromysinae, not Leptomysinae.     

Feeding habits of amphipods (Crustacea: Malacostraca) from shallow soft bottom communities: Comparison between marine caves and open habitats

Marine caves are environments of great interest since the organisms that inhabit them are forced to develop specific adaptations to high constraint conditions. Because of some of these particular conditions, such as light absence or oligotrophy, it can be expected that feeding strategies into caves differ from that present outside them. Nevertheless, no studies have been done to compare the trophic structure of marine caves and open habitats, at least for amphipod communities, considering their importance both inside and outside of the caves. In this study, the diet of the dominant amphipod species living on shallow sediments, both inside and outside of six marine caves in western Mediterranean, was characterized. Thereby, the gut content of 17 amphipod species was studied, being this study the first attempt to establish the feeding habit of most of these species. Analysis of digestive contents of the species showed that amphipod diet is less diverse in sediments than in other environments, such as algae and seagrasses. No herbivorous species were found in the sediment and carnivorous amphipods showed a little variety of prey, feeding mainly on crustaceans. Differences in the trophic structure were also found between marine caves and open habitats sediments: while outside the caves detritivorous was the dominant group (both in number of species and number of individuals), amphipods mainly play the role of carnivorous inside the caves. No detritivorous species were found into the caves, where carnivorous represents almost 60% of amphipods species and more than 80% of amphipod individuals. This pattern obtained in amphipods differ from the general trend observed in marine cave organisms, for which a generalist diet, such as omnivory, usually is an advantage in these oligotrophic conditions. The possible causes of this pattern are discussed.     

Organic pollution of an inland marine cave from Bermuda

Inland marine caves have served as biological refugia and, as such, frequently contain unique assemblages of taxonomically significant relict populations known only from a single cave or cave system. However, the absence of photosynthesis in this totally lightless environment and the presence of a highly stratified water column and long residence times for cave waters, characteristically result in depleted dissolved oxygen levels. Under such circumstances, organic pollution, even at low levels, can produce anoxic conditions and may consequently result in the extinction of entire species of cavernicolous fauna. We here present data on the normal environment of marine caves in Bermuda and cite as an example a cave which has become grossly polluted as a result of indiscriminate dumping.     

Movement patterns of the sea urchin Paracentrotus lividus in a marine reserve and an unprotected area in the NW Mediterranean

Sea urchins can alter the composition and dynamics of algal communities by grazing. Changes in their displacement capability can influence their grazing and thus their effect on algal communities. The daily and monthly movement of Paracentrotus lividus inside and outside a marine reserve in the NW Mediterranean Sea was studied in order to determine the role of predation in sea urchin movements and its potential grazing impacts. During the 3‐month study, the total distance travelled by sea urchins ranged from 71.8 to 673.6 cm (mean = 379.2 cm, SD = 221.3). Home ranges (distances from the initial position) were between 50 and 302 cm (mean = 150.7 cm, SD = 99.4). Movements were not directional but random, with a mean of 150 cm around the initial position. The activity of Paracentrotus lividus was significantly higher at night than during daylight. Total displacement at the end of the experiment was significantly higher in the unprotected area than in the reserve, as was the home range. The linear distance travelled in a 24‐h period ranged between 6 and 220 cm with a mean home range of 51 cm. These results agree with those of other authors, and allow us to rule out major migratory processes of sea urchin populations in this area. Nevertheless, these results underline the role of predatory fishes in reducing herbivory pressure by trophic cascades, reinforcing the evidence that sea urchin escape behaviour may be an important factor in structuring algal communities.     

Mycobiota associated with the rhodophyte alien species Asparagopsis taxiformis (Delile) Trevisan de Saint‐Léon in the Mediterranean Sea

This study is the first to investigate and characterize the mycobiota associated with the alien species Asparagopsis taxiformis, a rhodophyte classified as one of the ‘100 worst invasive species’ in the Mediterranean Sea by the Hellenic Center for Marine Research, as it threatens biodiversity. Fungal endophyte and epiphyte communities were investigated on algal specimens from two sampling sites on the island of Linosa (Marine Protected Area Isole Pelagie, Italy). 87% of the 24 specimens that were analysed for epiphytes displayed a microfungal colonization. No endophytes were found. Only a small amount of microfungi was found to be associated with this alga. Only five fungal taxa were isolated; two of which are sporadically associated with the alga, while three, Eurotium rubrum, Cladosporium cladosporioides and Cladosporium pseudocladosporioides, seem to be related to A. taxiformis. This scarcity could be related to algal chemical composition.     

Marine conservation challenges in an era of economic crisis and geopolitical instability: The case of the Mediterranean Sea

In the Mediterranean Sea, socio-economic drivers may accelerate the process of exclusive economic zone (EEZ) declarations. Despite the challenges, the EEZ declarations may provide important opportunities for leveraging change to national policy towards the development of large-scale conservation of marine ecosystems and biodiversity in this zone. Using the Mediterranean Sea as a case study, we aim to highlight a set of best practices that will maximize the potential for the development of large-scale marine conservation initiatives. These include a range of approaches, such as using surrogates to fill the many biodiversity data gaps in the region, further the development of consistent and open access databases, and the utilization of technological developments to improve monitoring, research and surveillance of less accessible and under-explored marine areas. The integration of Mediterranean-wide and local conservation efforts, the facilitation of transboundary collaboration, and the establishment of regional funds for conservation will further enhance opportunities for marine conservation in this region.     

气候变化对海洋生物多样性的影响

本文针对国内外有关气候变化对海洋生物多样性影响的研究情况，分别从温度、CO2浓度变化、海平面上升、降雨量、海洋水文结构和海流变化以及紫外线辐射增强等方面探讨气候变化对海洋生物多样性的影响，并从病原生物传播、浮游生物群落结构、海洋鱼类群落结构变化等方面分析气候变化对海洋生态系统的影响．针对全球气候变化对我国海洋生物多样性保护带来的挑战，提出了今后的研究重点．     

中国海洋浮游植物和赤潮物种的生物多样性研究进展(一):渤海

渤海是我国最大的半封闭型内海,也是重要的海洋生物产卵场、育幼场和索饵场。其陆源径流丰富,环境异质性高,具有很高的浮游植物物种多样性,其中包括大量的赤潮物种。该海域赤潮频发对水产养殖、生态环境、旅游业等造成了很大影响。因此,针对该海域浮游植物物种多样性的研究具有极其重要的意义。本文整理了1957—2019年间46个主要航次的调查资料,比较分析了这些航次中鉴定的浮游植物组成以及相对丰度,并跟踪分析了近20年来渤海的主要赤潮事件,重点描述了赤潮物种多样性,并详细阐述了常见赤潮物种。在航次鉴定的140个赤潮物种中,包括硅藻78种,甲藻52种,定鞭藻5种,绿藻4种和蓝藻1种。本文为比较研究包括渤海海域在内的不同海域浮游植物和赤潮物种的多样性,探索海洋生物多样性的起源和演化规律,解读宏基因组学及宏条形码分析结果提供了基础和参考。     

Spatial patterns of biodiversity in the Black Sea: An assessment using benthic polychaetes

The current study broadens the biodiversity information available for the Black Sea and neighbouring regions and improves our knowledge about the polychaete biogeographic patterns to be discerned in them. There appears to be a well-defined zoogeocline from the Marmara Sea and Bosphorus Strait to the inner parts of the region (Azov Sea), depicted both as a multivariate pattern and in terms of species (or taxa) numbers. The emergent multivariate pattern complies, to a certain extent, with Jakubova’s (1935) views: three main sectors can be defined in the basin: (a) Prebosphoric, (b) the Black Sea and, (c) the Azov Sea, whereas the Bosphorus Strait and Marmara Sea show less faunal affinities with the afore-mentioned sectors. Patterns derived both from the cosmopolitan and Atlanto-Mediterranean species closely follow the one coming from the polychaete species and genera inventories. As a general trend, species numbers decrease along with the decrease in salinity towards the inner parts of the region. The trend is homologous to that seen in the benthic invertebrate inventories of all the major European semi-enclosed regional seas. Salinity and food availability appear to be the dominant abiotic factors correlated, though weakly, with the various patterns deriving from the taxonomic/zoogeographic categories. With the exception of the Anatolia, polychaete inventories from all sectors appear to be random samples of the total inventory of the region, in terms of taxonomic distinctness values. Therefore, these sectoral inventories can be used for future biodiversity/environmental impact assessment studies. A massive invasion of Mediterranean species after the opening of the Black Sea, in the lower Quaternary period, appears to be the likely biogeographic mechanism through which the old Sarmatic fauna was almost completely replaced by species of marine origin.     

黄东海间遗传连通性多物种案例分析与启示

黄东海是太平洋重要的边缘海,拥有丰富的海洋生物多样性资源,但在气候变化及人类活动的双重胁迫下,黄东海面临生物多样性衰退等问题,亟需加强保护。保护区网络被认为可有效防止或减缓人类活动和气候变化对生物多样性的负面影响,但连通性作为保护区网络的重要参数却很难直接测量。因此,本研究希望通过梳理已发表的黄东海间遗传连通性文献来揭示黄东海区域内的保护区是否形成网络。通过搜集已发表的137篇文献,共获得175个研究案例。超过半数的案例(65.71%)认为黄东海之间具有显著的遗传分化,这也意味着跨生态区群体间具有较低的遗传连通性。在总结影响因素时发现长江冲淡水、生境异质性、空间距离、进化过程、生活史(较短的浮游幼体期)、洋流等6个因素阻碍了黄东海群体间的遗传连通性,而洋流、生活史(较长的浮游幼体期)等2个因素促进了遗传连通性。进一步分析发现黄东海群体间的遗传连通性与空间距离呈显著的负相关,而与浮游幼体期呈正相关。因此,根据本研究结果,建议未来保护区网络建设应针对黄、东海生态区分别构建网络,在网络设计时应考虑保护对象的扩散能力及保护区间的空间距离。本研究不仅为构建涵盖黄东海的海洋保护区网络提供理论基础,...     

Horizontal Transfer of Matter by a Cave-Dwelling Mysid

Abstract. The role of a cave-dwelling mysid population as a matter carrier was evaluated in a cave of the Medes Is. (NW Mediterranean) during 1988–89. Hemimysis spehmcola (Ledoyer , 1963) is a gregarious mysid whose swarms migrate daily from the inner end of the cave - where they remain during the day–to the exterior where they feed during the night. Fecal pellet composition, pellet egestion and pellet decomposition were measured in order to evaluate the transfer of matter by mysids. Based on the strong daily behavioural rhythms of these mysids. special importance was attached to the sampling frequency (every 2 h). In order to assess seasonal variability, four daily cycles were evaluated within a year. In situ incubations were carried out to determine changes on the egestion rates, morphology, weight and composition (AA. C: N ratios) of pellets in the course of a day. The number of pellets deposited inside the cave was estimated using containers distributed along the cave bottom by SCUBA divers. Fecal pellets showed an amorphous composition, whereby diatom frustules, dinoflagellate loricae and coccolithoporids were very scarce. About 25% of the body weight were estimated to be egested daily as fecal pellets, suggesting a detritivorous feeding habit. The population oscillated seasonally between 1 and 12 millions. Individuals egested between 1.6 and 3.5 pellets a day into the cave, each averaging from 9.4 to 11.9 μg DW, 0.5 to 1.0 μg C and 0.046 to 0.27 μg N. Therefore, the population carried daily about 20 407 g DW POM. 2–21 g C and 0.5 2.7 g N from outside to inside the cave. Pellets decomposed very quickly; between 20 to 50% of both C and N were released from pellets in less than 2 h after egestion. Oxidation of pellets theoretically consumes an amount of oxygen which agrees with the high BOD values previously reported for this cave. Marine caves are generally viewed as strictly oligotrophic systems; dense mysid populations, however, could strongly modify the trophic relationships in marine caves.     

Habitat Selection and Ecology of Two Speleophilic Gobiid Fishes(Pisces: Gobiidae) from the Western Mediterranean Sea

The life habits and microhabitat selection of speleophilic gobiid fishes were investigated on the Balearic Island of Ibiza (western Mediterranean Sea). Corcyrogobius liechtensteini (Kolombatović, 1891) was the most frequent species within submarine crevices, cavities and caves. Didogobius splechtnai Ahnelt & Patzner, 1995 co-occurred with C. liechtensteini in several cases but showed a clear spatial separation based on a different microhabitat preference. The microhabitat selection of both species was confirmed by habitat-choice experiments in anaquarium. Corcyrogobius liechtensteini was frequent in depths between 5 m and 25 m and attained abundances of 14.2 individuals · m⁻². This species inhabited the ceilings and upper parts of the walls within the caves. D. splechtnai was mainly found between 7 m and 11 m and showed abundances of approximately 2 indivi-duals · m⁻². This species occupied the fine sediment bottoms in the innermost parts of crevices and caves. The growth and the maximum age of both species were determined by length-frequency distributions. The overall sex ratio from catches of three different years are given for C. liechtensteini .