The demography of the coelacanth Latimeria chalumnae

Synopsis The very sparse data that are available on the abundance, population structure and biology of the coelacanth Latimeria chalumnae off Grand Comoro are summarised, and some simple numerical analyses are carried out to explore certain aspects of the population dynamics, particularly the age-profile of the population. The object has not been to provide estimates of key demographic parameters, such as mortality rates, but to propose various scenarios that are useful for comparison with real data as they become available. The analysis also makes it possible to reach some preliminary conclusions that are relevant to the management of the coelacanth population. For instance, it appears that the catch rate of coelacanths by artisanal fishermen may have a negligible effect on coelacanth survivorship, and it is more likely that population size and structure are determined by natural mortality rates and birth rates. It is suggested that predation is the main cause of natural mortality and that the main predators of coelacanths are likely to be large sharks. Interference with the traditional patterns of the Comoran artisanal fishery may threaten the coelacanth. Several important gaps in our knowledge of coelacanth demography are identified.     

Function of the pseudomaxillary fold in the mouth opening of the coelacanth, <Emphasis Type="Italic">Latimeria chalumnae</Emphasis>

The oral region of a coelacanth, Latimeria chalumnae, from the Comoro Islands, was examined in fresh condition, and the complex folding of the membrane between the lower jaw and the skull, and a strut-like structure in the membrane were found. When the mouth opened, the pseudomaxillary folds covered the lateral walls of the mouth. The movement of the membrane is inferred from the observation of the specimen. Coelacanths developed a way of covering the lateral sides of mouth that is different from that of other fishes; especially the pseudomaxillary fold above the strut is unique to coelacanths, but the pseudomaxillary fold below the strut is similar to the lower lip fold of the lepidosirenoid lungfish.     

Extremely slow rate of evolution in the HOX cluster revealed by comparison between Tanzanian and Indonesian coelacanths

Coelacanths are known as "living fossils" because their morphology has changed very little from that in the fossil record. To elucidate why coelacanths have evolved so slowly is thus of primary importance in evolutionary biology. In the present study, we determined the entire sequence of the HOX cluster of the Tanzanian coelacanth (Latimeria chalumnae) and compared it with that of the Indonesian coelacanth (L. menadoensis), which was available in the literature. The most intriguing result was the extremely small genetic divergence between the two coelacanths. The synonymous divergence of the HOX coding region between the two coelacanths was estimated to be 0.07%, which is ~11-fold smaller than that of human-chimp. When we applied the estimated divergence time of the two coelacanths of 6 million years ago (MYA) and 30 MYA, which were proposed in independent mitochondrial DNA analyses, the synonymous substitution rate of the coelacanth HOX cluster was estimated to be ~11-fold and 56-fold smaller than that of human-chimp, respectively. Thus, the present study implies that the reduction of the nucleotide substitution rate in coelacanth HOX genes may account for the conservation of coelacanth morphology during evolution.     

An inventory of all known specimens of the coelacanth Latimeria chalumnae,with comments on trends in the catches

Synopsis A list is presented of all known specimens of the coelacanth Latimeria chalumnae based on a survey of the literature and of museum, aquarium and university holdings. Details are given of the date, place, time and depth of capture, the name and age of the fisherman, the length, weight and sex of the fish, the first literature record, the method of preservation and the present location of specimens, if known. A new number is assigned to each specimen. At least 172 coelacanths are known to have been caught since 1938. The first coelacanth was caught off South Africa but all properly documented, subsequent specimens have been caught off the islands of Grand Comoro and Anjouan in the Comoros. An appeal is made to the Comoran authorities for each specimen that is caught to be made available for scientific study. Museum authorities are also encouraged to allow their specimens to be X-rayed or dissected so that vital information can be obtained on fecundity, foetal nutrition and dietary preferences. It is essential for the coelacanth conservation effort that this inventory is maintained by the Coelacanth Conservation Council.     

CoelacanthLatimeria chalumnae aggregates in caves: first observations on their resting habitat and social behavior

Synopsis We report first observations on social behavior ofLatimeria chalumnae in its natural environment at around 200 m depth in the Comoro Archipelago, Western Indian Ocean. Coelacanths aggregate in small nonaggressive groups in sheltered lava-caves. They live in open site-attached social systems with individuals occupying large overlapping home ranges of more than 8 km coastline, some for a period of at least 2 years.     

A newly recognized fossil coelacanth highlights the early morphological diversification of the clade

Previously considered an actinopterygian or an osteichthyan incertae sedis, the Devonian (Givetian-Frasnian) Holopterygius nudus is reinterpreted as a coelacanth. This genus is among the oldest coelacanths known from articulated remains, but its eel-like morphology marks a considerable departure from the conventional coelacanth body plan. A cladistic analysis places Holopterygius as the sister taxon of the Carboniferous (Serpukhovian) genus Allenypterus. Despite the specialized morphology of these genera, they occupy a surprisingly basal position in coelacanth phylogeny; only Diplocercides and Miguashaia are further removed from the crown. A morphometric analysis reveals that coelacanths were anatomically disparate early in their history. Conflicts between this result and those of previous studies challenge the adequacy of systematic character sets for describing historical patterns of morphological variety. Coelacanths have long had an iconic place in the study of vertebrate evolution for their apparent anatomical conservatism over geological time, but Holopterygius provides clear evidence for rapid morphological evolution early in the history of this clade.     

Enzymes of the coelacanth Latimeria chalumnae evidenced by starch gel electrophoresis

Synopsis The coelacanth, Latimeria chalumnae, is often referred to as a living relic. The opportunity to examine its biochemical molecular structure was sought in an effort to define the degree of its genetic variability. The coelacanth is thought to live only in a small area around the Comoro Islands in the Western Indian Ocean. The scenario presented suggests that the coelacanth may have lost genetic variability as a result of genetic drift within a small population. The narrow geographic range of the coelacanth suggests adjustment to a relatively limited environment. The loss of specific alleles through genetic drift can reduce the ability of a fish population to adapt to changes in environmental conditions. The coelacanth needs strong conservation measures to be taken to curtail the capture of specimens and for the protection of its limited natural habitat.     

Oldest coelacanth, from the Early Devonian of Australia

Coelacanths are well-known sarcopterygian (lobe-finned) fishes, which together with lungfishes are the closest extant relatives of land vertebrates (tetrapods). Coelacanths have both living representatives and a rich fossil record, but lack fossils older than the late Middle Devonian (385-390 Myr ago), conflicting with current phylogenies implying coelacanths diverged from other sarcopterygians in the earliest Devonian (410-415 Myr ago). Here, we report the discovery of a new coelacanth from the Early Devonian of Australia (407-409 Myr ago), which fills in the approximately 20 Myr 'ghost range' between previous coelacanth records and the predicted origin of the group. This taxon is based on a single lower jaw bone, the dentary, which is deep and short in form and possesses a dentary sensory pore, otherwise seen in Carboniferous and younger taxa.     

Interspecies Insertion Polymorphism Analysis Reveals Recent Activity of Transposable Elements in Extant Coelacanths

Coelacanths are lobe-finned fish represented by two extant species, Latimeria chalumnae in South Africa and Comoros and L. menadoensis in Indonesia. Due to their intermediate phylogenetic position between ray-finned fish and tetrapods in the vertebrate lineage, they are of great interest from an evolutionary point of view. In addition, extant specimens look similar to 300 million-year-old fossils; because of their apparent slowly evolving morphology, coelacanths have been often described as « living fossils ». As an underlying cause of such a morphological stasis, several authors have proposed a slow evolution of the coelacanth genome. Accordingly, sequencing of the L. chalumnae genome has revealed a globally low substitution rate for protein-coding regions compared to other vertebrates. However, genome and gene evolution can also be influenced by transposable elements, which form a major and dynamic part of vertebrate genomes through their ability to move, duplicate and recombine. In this work, we have searched for evidence of transposition activity in coelacanth genomes through the comparative analysis of orthologous genomic regions from both Latimeria species. Comparison of 5.7 Mb (0.2%) of the L. chalumnae genome with orthologous Bacterial Artificial Chromosome clones from L. menadoensis allowed the identification of 27 species-specific transposable element insertions, with a strong relative contribution of CR1 non-LTR retrotransposons. Species-specific homologous recombination between the long terminal repeats of a new coelacanth endogenous retrovirus was also detected. Our analysis suggests that transposon activity is responsible for at least 0.6% of genome divergence between both Latimeria species. Taken together, this study demonstrates that coelacanth genomes are not evolutionary inert: they contain recently active transposable elements, which have significantly contributed to post-speciation genome divergence in Latimeria.     

A fiftieth anniversary reflection on the living coelacanth,Latimeria chalumnae: some new interpretations of its natural history and conservation status

Synopsis It all started about 400 million years ago, when representatives of a group of fish-like fleshy-finned creatures appeared in the fossil record (or was it through a childhood dream shared by all of us that we would one day study the coelacanth?). Many of the coelacanth's characters placed them close to the ancestry of terrestrial vertebrates. About 70 million years ago they disappeared from the fossil record. The discovery in 1938 of the first living coelacanth, in 1952 of the second and until now over 200 specimens parallels in excitement an encounter with a live dinosaur on a weekend walk, and in significance even more than that. For this year's 50th anniversary of the famous discovery of the first living coelacanth, we retraced the routes and visited the main actors of this zoological drama. New insights into coelacanth natural history were facilitated by novel interpretation of earlier data and our expeditions to the Comoro Islands, retracing the route of the second specimen, measuring unrecorded specimens, interviewing fishermen and describing their fishing crafts, and taking part in recent events on land and water near the only known habitat of the living coelacanth. Entry into this habitat and observations from the research submersible GEO opened up a new era in coelacanth research. Past studies of preserved specimens, which were caught as an incidental bycatch, were supplemented for the first time by studies of free-living coelacanths in their natural habitat. The first film footage taken from the submersible revealed the entirely unfishlike movements of this creature. Its mode of locomotion is a combination of flying and gliding, interspersed with head stands and belly-up drifts which appear to defy gravity. The narrow range of habitat in which the coelacanth has been encountered has led us to realize how vulnerable this ancient relict is. The members of our expeditions therefore cooperated in establishing an international organisation to coordinate efforts to conserve the coelacanth. Editorial     

Abundance and habitat choice as an indication of colonisation history in Comoro Land birds

Louette, M., Stevens, J. & Neri, F. 2000. Abundance and habitat choice as an indication of colonisation history in Comoro Land birds. Ostrich 71 (1 & 2): 345–346.

We measured bird abundance and habitat parameters by point transect counts. Bird abundance is different for a number of species among the four Comoro islands (Louette, M. et al. 1993. Oiseau 63: 115–126), and among sites on a given island (especially in the case of forest birds, such as on Grand Comoro, an island with great variation in habitat).     

Mitogenomic analysis for coelacanths (Latimeria chalumnae) caught in Tanzania

In recent years, a large number of individuals of the species Latimeria chalumnae, one of the living fossil coelacanths, have been landed off the coast of Tanzania. Although L. chalumnae specimens have also been landed at other localities in the western Indian Ocean, so far, viable populations of this species have been identified only at two localities, Comoros and South Africa. Therefore, the recent active catch off Tanzania suggests a new habitat for L. chalumnae. To examine the genetic background of the Tanzanian fish, we analyzed complete mtDNA sequences of two Tanzanian individuals (Kigombe-9 and Songo Mnara-1) collected from the north and south coasts of Tanzania. Using the recently reported criteria for six haplotypes established in a population genetic study for coelacanths living in the western Indian Ocean [Schartl, M., Hornung, U., Hissman, K., Schauer, J., Fricke, H., 2005. Relatedness among east African coelacanths. Nature 435, 901.], we characterized Songo Mnara-1 as haplotype 1 and Kigombe-9 as haplotype 5. We suggest that the Songo Mnara specimen is a member of the Comoran group, but was swept away by the South Equatorial current. The individual from Kigombe may be a member of an undiscovered population that exists near the boundary between Tanzania and Kenya. Further analysis using more than 19 individuals recently captured off the north coast of Tanzania will reveal whether a new population exists there. Our sequence data suggest additional variable sites in the mtDNA sequence that may define the population structure of coelacanths in the western Indian Ocean and also raise the possibility that the previously published Comoran coelacanth mtDNA sequence contains several critical errors including base changes and indels.     

Silver coelacanths from Spain are not proofs of a Pre-scientific Discovery

In 1964 and 1965, two silver fish pendants depicting coelacanths and thought to be religious votive from Spanish churches were purchased on the art market in Spain, dated between the 17th and late 19th century, long before the discovery for science of the living coelacanth in 1938. It was speculated that they originated either from the Mediterranean, subtropical Atlantic, or subtropical Pacific (de Sylva 1966, Bruton 1985, Anonymous 1993, Greenwell 1994, Raynold & Mangiacopra 1995). Some authors believe that the silver fish pendants represent a new species of coelacanthiforms (Raynold & Mangiacopra 1995), others claimed the pendants to be only a wishful fantasy of scientists and not coelacanths at all (Thomson 1991). However, new studies revealed that the silver fish artefacts are indeed coelacanths but were in fact produced more recently with the Comorian coelacanth, Latimeria chalumnae, as a model.     

First results from satellite-linked archival tagging of porbeagle shark, Lamna nasus: Area fidelity, wider-scale movements and plasticity in diel depth changes

Understanding the habitat preferences of large marine vertebrates has only recently become tractable with the widespread availability of satellite telemetry for monitoring movements and behaviour. For many species with low population abundances, however, little progress has been made in identifying space use patterns. The endothermic porbeagle shark, Lamna nasus, has declined in the North Atlantic due to severe fishing pressure, with little evidence of recovery. One potential factor exacerbating population decline is area fidelity to coastal waters where fisheries are intensive. We tested for short-term area fidelity by attaching pop-up satellite-linked archival transmitters to four porbeagles in summer 2007, resulting in 175 days total tracking time covering an estimated 10,256 km distance. Throughout July and August the sharks occupied localised areas (8,602 – 90,153 km²) within the Celtic Sea, between the south-west UK, south-west Wales and southern Ireland. Only one shark was tracked into the autumn, when it moved into deep water off the continental shelf, then north towards colder latitudes. Sharks occupied a broad vertical depth range (0 – 552 m) and water temperatures (9° - 19 °C). Dives were made frequently from the surface to near the seabed in shelf areas, however, in shelf edge habitats extended periods of time were spent at depths > 300 m. Porbeagles showed considerable plasticity in diel depth changes within and between individuals and as a function of habitat type. In addition to no obvious day-night difference in depth occupation, some sharks showed reverse diel vertical migration (DVM) (dawn ascent – dusk descent) in well-mixed coastal waters whereas normal DVM (dawn descent – dusk ascent) characterised movements into deeper, thermally well-stratified waters. The variable behaviours may reflect the need for different search strategies depending on habitat and prey types encountered. These results show porbeagles are potentially vulnerable to fisheries throughout the summer when they aggregate, and that large scale movement across national boundaries identifies the need for international conservation measures.     

Coelacanths: a human responsibility

The living coelacanth Latimeria chalumnae has a unique position in world biodiversity which raises important questions about conservation and ethics. Some relevant details of coelacanth biology are summarized, including those obtained by direct observation from submersibles. The importance of the coelacanth for evolutionary theory and palaeontology is shown to be paralleled in cultural, literary and artistic areas of human heritage. Threats to the Comoran coelacanths from artisanal fishing are described and conservation measures discussed in relation to local customs and economies as well as the promotion of tourism to spread a new awareness and concern for coelacanths worldwide.     

Population ecology ofPimelodella kronei,troglobitic catfish from Southeastern Brazil (Siluriformes,Pimelodiae)

Synopsis Pimelodella kronei, a cave derivative of the pimelodid catfishP. transitoria, has been found in five caves of the Ribeira River valley, SE Brazil. I investigated the ecology of the most conspicuous population, that from Areias Caves, by capture-mark-recapture. During the two-year study, 170 specimens ofP. kronei were marked, 81 were recaptured at least once, multiple recaptures were frequent. These blind catfishes tended to be recaptured in the same site where they were marked, suggesting a sedentary life and possibly territoriality. The home range was 200 m along the stream, or more where food was scarcer.P. kronei population size in 800 m of permanent stream was 150–200 individuals.P. transitoria also inhabits the cave, at a density ten times lower. The growth rate ofP. kronei was less than 0.1 cm × month^–1, and longevity was 10–15 years. These data, and evidences of an infrequent reproduction, indicate a K-selected life cycle, probably as a consequence of food scarcity.     

The ecology and conservation of the coelacanth Latimeria chalumnae

Synopsis Studies on the ecology of the living coelacanth, Latimeria chalumnae, are reviewed and assessed. Early predictions on the life history of the coelacanth have proved to be accurate but recent findings have improved our understanding of its habitat and feeding preferences, diel activity patterns and social behaviour. A history of coelacanth conservation reveals that there has been a sustained concern for the survival of this species which has eventually culminated in several effective conservation actions in recent years. The coelacanth is threatened by a number of socio-economic and biological factors, but international action directed at managing the fishery in the Comoros should ensure that the species survives. Recent observations on living coelacanths in their natural environment have greatly improved our knowledge of the behaviour and relative abundance of adults. Important priorities for future research include studies on the distribution and abundance of juveniles and breeding adults, both off the Comoros and elsewhere. The coelacanth is a highly specialised, precocial fish which occupies a unique place in biology. Co-ordinated international efforts should continue to be made to understand and conserve this remarkable fish.     

Population structure and movement patterns within an isolated and endangered population of the moth Dysauxes ancilla L. (Lepidoptera, Ctenuchidae): implications for conservation

Population structure and movement patterns were studied in a geographically isolated and endangered Swedish population of the moth Dysauxes ancilla. Mark-recapture experiments were conducted between 1993–97 and observations made of adult behaviour between 1993–99. The population size was estimated to be 2356 individuals and the average male residence time 2.6 days. Adults of D. ancilla were sedentary and all movements occurred within the breeding area, despite searches up to 0.5 km away. Mean (± S.D.) distance between two successive captures in males was 43 ± 36 m and the longest single move was 144 m. The moths showed a behavioural reluctance to enter heavily grazed meadows or dense vegetation at the edges of the breeding habitat. Since there are no suitable habitats in the surroundings, conservation efforts should focus on managing the continuity of the current breeding area. Further, the restoration of a former breeding habitat situated 1 km to the north should be considered as well as establishing connections with the extant population.     

Growth, Natural Mortality, Length–weight Relationship, Maximum Length and Length-at-first-maturity of the Coelacanth Latimeria chalumnae

Based on the re-interpretation of published data, the von Bertalanffy growth function parameters of the coelacanth, Latimeria chalumnae, are estimated as L=218cm total length, s.e. 23; K=0.059 (year^–1), s.e. 0.012; t₀=–3.3 (year), s.e. 0.5, corresponding to a life span of 48 years. The length–weight relationship of the form W=a*TLb, with wet weight (W) in g and total length (TL) in cm, is estimated as a=0.0278, b=2.89, r²=0.893, n=87, range=42.5–183cm TL. Using extreme value theory, the maximum length for female coelacanths is estimated as 199cm TL (95% confidence interval=175–223cm TL) and for males as 168cm TL (95% confidence interval 155–180cm TL). Based on data from seven females with embryos or mature eggs, the length-at-first-maturity for females is estimated to be about 150cm TL, corresponding to an age of about 16 years. Based on the value of t₀=–3.3 years and on the presence of three scale rings found in a newborn coelacanth, the period of embryogenesis lasts for about three years, the longest known in vertebrates. The natural mortality rate is estimated at M=0.12. Population food consumption is found to be 1.4 times the existing biomass per year, and gross food conversion efficiency indicates that only 10% of the consumed food is utilized for somatic growth.     

Early Growth Stages in Coelacanth Fishes

ALL known specimens of the Recent coelacanth fish, Latimeria, are large specimens (mostly more than 100 cm total length) and only one female with eggs has been recorded^1,2. Consequently, the ontogeny and the early growth stages of the Recent coelacanth are unknown. In the fossil record, one specimen of Holophagus (= Undina) from the Upper Jurassic of Solnhofen, southern Germany, has been recorded with two young coelacanths inside³. Watson has argued from this finding that the coelacanths are viviparous but it seems more reasonable to interpret this fossil specimen as a cannibal that had just swallowed two young of its own kind. This interpretation is favoured by the position of the two specimens and by the discovery of other fossil coelacanths containing large specimens of other kinds of fishes in their stomachs⁴.