Deep-water Habitats Research Articles

Shark and ray diversity in the Tropical America (Neotropics)-an examination of environmental and historical factors affecting diversity.

We present the first comprehensive review of the present and past shark and ray diversity in marine waters of Tropical America, examining the patterns of distribution in the Eastern Central Pacific (EP) and Western Central Atlantic (WA) realms. We identified the major regions of diversity and of endemism, and explored the relations to physical variables. We found a strong relationship between shark and ray diversity with area and coastal length of each province. The Tropical Northwestern Atlantic Province is characterized by high diversity and greater occurrence of endemic species, suggesting this province as the hotspot of sharks and rays in Tropical America. The historical background for the current biogeography is explored and analyzed. Referential data from 67 geological units in 17 countries, from both shallow and deep-water habitats, across five time-clusters from the Miocene to the Pleistocene were studied. New data include 20 new assemblages from six countries. The most diverse Neogene and extant groups of shark and ray are Carcharhiniformes and Myliobatiformes, respectively. The differentiation between Pacific and Atlantic faunas goes to at least the middle Miocene, probably related with the increasing closure of the Central American Seaway acting as a barrier. The highest faunal similarity between the assemblages from the EP and the WA at the early Miocene could be related to the lack of a barrier back then, but increased sampling is needed to substantiate this hypothesis.

The first record of Sirenia (Mammalia) from the early Oligocene of the Paratethys

Sirenians have been extensively recorded from the Mediterranean and west European localities but there are only few finds to the east of it, from the area covered by the Paratethys. For the early Oligocene, to our knowledge, there are no published records of sirenians from inner seas of the Old World. Here we report a specimen of Dugongidae indet., consisting of two partial vertebrae and 12 fragments of ribs, collected in a manganese ore mine in Ukraine and dated as the earliest Oligocene (33–32 Ma). The specimens, as preserved, did not differ in morphology and size from ‘Halitherium schinzii’ and therefore can belong to Kaupitherium, at present the single early Oligocene genus recorded from Europe. However, its vertebral and rib anatomy is not specific for Kaupitherium, so we identify it only by family level. The marks of scavenging on a rib possibly are due to gastropod or bivalve mollusks. The sea, as suggested from biotic data, had a temperate or subtropical climate, relatively cold waters and high diversity of pelagic and deep-water habitats. Thus, the onset of the Oligocene was a period when sirenians could enter temperate inner Eurasian waters, a marginal area in their worldwide dispersal.

From top to bottom: Do Lake Trout diversify along a depth gradient in Great Bear Lake, NT, Canada?

Depth is usually considered the main driver of Lake Trout intraspecific diversity across lakes in North America. Given that Great Bear Lake is one of the largest and deepest freshwater systems in North America, we predicted that Lake Trout intraspecific diversity to be organized along a depth axis within this system. Thus, we investigated whether a deep-water morph of Lake Trout co-existed with four shallow-water morphs previously described in Great Bear Lake. Morphology, neutral genetic variation, isotopic niches, and life-history traits of Lake Trout across depths (0–150 m) were compared among morphs. Due to the propensity of Lake Trout with high levels of morphological diversity to occupy multiple habitat niches, a novel multivariate grouping method using a suite of composite variables was applied in addition to two other commonly used grouping methods to classify individuals. Depth alone did not explain Lake Trout diversity in Great Bear Lake; a distinct fifth deep-water morph was not found. Rather, Lake Trout diversity followed an ecological continuum, with some evidence for adaptation to local conditions in deep-water habitat. Overall, trout caught from deep-water showed low levels of genetic and phenotypic differentiation from shallow-water trout, and displayed higher lipid content (C:N ratio) and occupied a higher trophic level that suggested an potential increase of piscivory (including cannibalism) than the previously described four morphs. Why phenotypic divergence between shallow- and deep-water Lake Trout was low is unknown, especially when the potential for phenotypic variation should be high in deep and large Great Bear Lake. Given that variation in complexity of freshwater environments has dramatic consequences for divergence, variation in the complexity in Great Bear Lake (i.e., shallow being more complex than deep), may explain the observed dichotomy in the expression of intraspecific phenotypic diversity between shallow- vs. deep-water habitats. The ambiguity surrounding mechanisms driving divergence of Lake Trout in Great Bear Lake should be seen as reflective of the highly variable nature of ecological opportunity and divergent natural selection itself.

Distribution and foraging patterns of common loons on Lake Michigan with implications for exposure to type E avian botulism

Abstract Common loons (Gavia immer) staging on the Great Lakes during fall migration are at risk to episodic outbreaks of type E botulism. Information on distribution, foraging patterns, and exposure routes of loons are needed for understanding the physical and ecological factors that contribute to avian botulism outbreaks. Aerial surveys were conducted to document the spatiotemporal distribution of common loons on Lake Michigan during falls 2011–2013. In addition, satellite telemetry and archival geolocator tags were used to determine the distribution and foraging patterns of individual common loons while using Lake Michigan during fall migration. Common loon distribution observed during aerial surveys and movements of individual radiomarked and/or geotagged loons suggest a seasonal pattern of use, with early fall use of Green Bay and northern Lake Michigan followed by a shift in distribution to southern Lake Michigan before moving on to wintering areas. Common loons tended to occupy offshore areas of Lake Michigan and, on average, spent the majority of daylight hours foraging. Dive depths were as deep as 60 m and dive characteristics suggested that loons were primarily foraging on benthic prey. A recent study concluded that round gobies (Neogobius melanostomus) are an important prey item of common loons and may be involved in transmission of botulinum neurotoxin type E. Loon distribution coincides with the distribution of dreissenid mussel biomass, an important food resource for round gobies. Our observations support speculation that energy transfer to higher trophic levels via gobies may occur in deep-water habitats, along with transfer of botulinum neurotoxin.

Adult Snapping Turtle (<i>Chelydra serpentina</i>) Feeding on Goldeneye Embryos of Pumpkinseed (<i>Lepomis gibbosus</i>) in Defended Nests

Rarely observed predatory behaviour of adult Snapping Turtles (Chelydra serpentina) was recorded using remote video technology. We observed turtles inspecting and, in one case, apparently feeding on goldeneye stage embryos (< 3 mm) from defended nests of Pumpkinseed (Lepomis gibbosus). This novel behaviour was limited to nests in a secluded bay and was not observed at nests located along exposed shorelines or on shallow shoals in the deep open water habitat of an inland oligotrophic lake. The benefit of feeding on small prey is likely enhanced by embryos being clustered in nests and by an abundance of sunfish nests. Low-cost and low-intrusion video technology provides excellent opportunities, even in aquatic systems, to document novel predator and prey behaviours.

Temperature and salinity influence the chemistry in the pre-hatch otolith region of capelin, Mallotus villosus, during lab and field egg incubation experiments

Identifying the natal origin of marine fish is important to understand connectivity and productivity among populations residing in different habitats. Capelin, Mallotus villosus, is a key marine forage fish species that spawns at both beach (warmer, less saline) and deep-water (15–40 m; cooler, more saline) habitats along the northeast Newfoundland coast. Currently, the contribution from each habitat to the spawning population is unknown. Previous research, where capelin eggs from known families were lab-reared, identified family-based otolith chemical signatures in the pre-hatch (embryonic) region of larvae. In this study, we investigated whether temperature and salinity influenced embryonic otolith chemistry to determine whether variation in environmental conditions would result in habitat-specific signatures, thereby overwhelming family-based signatures and allowing the identification of natal origin. Capelin eggs from many families were incubated together under controlled temperature (4, 8, 10 °C) and salinity (10, 20, 30 psu) treatments in the lab and uncontrolled conditions within each habitat in the field. Elemental concentrations (i.e., Sr, Ba, Mg, Mn) in the pre-hatch region of 1-day old larvae were quantified via LA ICP-MS. Elemental concentrations varied among individuals reared under identical conditions, likely due to family-based chemical signatures. Despite this variation, mean elemental concentrations differed when mean temperatures varied by ≥4 °C between treatments (lab) and rearing habitats (field), resulting in high treatment- and habitat-specific classification success (~73–88%) of individuals. In contrast, embryonic otolith chemistry did not vary consistently with salinity. Temperature differs consistently between rearing habitats and, thus, these findings suggest that when capelin rearing habitats differ by ≥4 °C within a year, chemical signatures in the embryonic otolith may be used to determine the natal origin of individuals.

Confirmation of diving and swimming behavior in the Sora (Porzana carolina)

ABSTRACT We first observed Sora (Porzana carolina) swimming and diving under water while capturing them with hand nets at night. Since that time, we have observed the behavior several times and documented it with photos and video. Rails are among the most elusive birds. Despite living in wetlands, water depth has often been discussed as a factor limiting habitat use by rails, with the assumption that when water levels are too deep, rails walk on top of floating or emergent vegetation. Here we present new photo and video evidence that confirms previous observations that Sora are adept at swimming and diving in deep water wetland environments, which should alter how we think about their ability to avoid predators and use deeper water habitats.

Seasonal changes in habitat suitability for adult shovelnose sturgeon in the lower Mississippi River

Shovelnose sturgeon Scaphirhynchus platorynchus are a large-river fish distributed throughout the Mississippi River basin, including the lower 1,533 km of the Mississippi River where riverine habitat has been and continues to be modified for navigation and is a potential site for development of instream hydrokinetic electric power generation. Information about habitat use and preference is essential to future conservation efforts. Shovelnose sturgeon have previously been found to select particular habitat types, and these selected habitats vary seasonally; although these past analyses do not consider the selected habitats in a landscape context. We used ecological niche factor analysis (ENFA) that uses distributions of telemetry locations and environmental variables to model habitat suitability in a landscape context. We recorded 333 locations of shovelnose sturgeon during July–December 2013 that included periods of relatively high and low river stages. The ENFA analysis indicated high-suitability locations were in or near deep water during both high and low river stages. During high river stages, high-suitability locations were near island tip habitat, deep water, and steep bottom slope and far from main channel habitat. During low stages, high-suitability locations were in or near deep water and main channel habitat and far from secondary channel and wing dike habitats. This landscape-scale analysis supports seasonal shifts in habitat use and provides insights that can be used to inform habitat conservation and management to benefit shovelnose sturgeon in the lower Mississippi River and possibly other large rivers.

Effect of colored light regimes on the stress response and RNA/DNA ratio of juvenile red sea bream, Pagrus major

We hypothesized that fish with red-sensitive retina would be stressed by red light and thus inhibited in somatic growth. Red sea bream (Pagrus major) juveniles (total length =3 cm) with red-sensitive retina were chosen to test this hypothesis. We examined the effect of different color lights (red with λmax 605 nm; green with λmax 540 nm; blue with λmax at 435 nm; and white with full spectrum) on unfed juveniles in laboratory tanks. Stress level was measured by the plasma cortisol and glucose concentrations, and nutritional status by muscle RNA/DNA ratio. Under red light, plasma cortisol and glucose, and muscle RNA/DNA were significantly higher than under green, blue, or white light. Our hypothesis was partly supported by previous findings on the effects of the color environment and spectral sensitivity of reared fishes. However, the levels of cortisol, glucose, and RNA/DNA in this study were low compared to published values. It seems that hatchery-bred juvenile red sea bream have adapted to red-rich surface light and are able to cope with the stress of living in surface floating cages which is so different from their deep-water habitats.

Reproductive features of the deep water hermit crab Sympagurus dimorphus (Anomura: Parapaguridae) inhabiting pseudoshells in the SW Atlantic Ocean

ABSTRACTWe herein provide information on reproductive aspects (the gonopore morphology of both sexes, the size of embryonic eggs and the fecundity) of the hermit crab Sympagurus dimorphus from deep waters of Argentina. Sampling was performed in different oceanographic and fishery campaigns between 2007 and 2010, at 38°01′–39°03′S, 55°–56°05′W, 85–150 m depth, in the SW Atlantic Ocean. Mean shield length (SL) of females was 9.41 ± 1.68 mm (range: 6.6–10.5 mm). Female gonopores measured 0.74 ± 0.10 mm (range: 0.45–0.90 mm), were oval in shape, and were more rounded and larger than male gonopores (0.51 ± 0.007, range: 0.4–0.63 mm). The egg volume for each of the three stages of embryonic development was 0.94 ± 0.20 mm3 for stage I, 1.14 ± 0.31 mm3 for stage II, and 2.07 ± 0.41 mm3 for stage III. Egg volume doubled from the initial to the final stage of the embryonic development. The fecundity (number of eggs) varied from 831 (6.62 mm SL) to 2246 (10 mm SL) for females carrying eggs in stages I+II, and between 158 (158 mm SL) and 1279 (9.4 mm SL) in stage III. The reproductive characteristics of the observed profile are intimately related to S. dimorphus use of a pseudoshell resource and a deep water habitat.

Chemical signatures in embryonic otoliths of capelin, Mallotus villosus: Influence of family and environmental conditions

Identifying the natal origin of marine fish species is important to understand the relative productivity and connectivity of local populations. Otolith chemistry can represent habitats occupied during early life history stages, thereby distinguishing individuals reared in different habitats. Maternal investment of trace elements into otoliths, however, may confound identifying an individual's natal origin. Capelin, Mallotus villosus, is a key forage fish species that spawns at both beach and deep-water (‘demersal’, 15–40m) habitats along the eastern Newfoundland coast. Currently, the relative productivity of these habitats to the spawning population is unknown. Previous research demonstrated habitat-specific chemical signatures of bulk samples of embryonic capelin otoliths could be identified. It is unclear, however, whether these habitat-specific signatures would be detectable in individuals due to confounding maternal effects. To investigate maternal investment, we tested whether family-based chemical signatures could be identified from otoliths of 1-d old larvae when eggs from known families (1 male+1 female) were reared under controlled temperature (10, 15°C) and salinity (10, 20, 30psu) treatments. Trace element concentrations in embryonic otoliths, particularly Ba, differed significantly among families, resulting in a high classification success of offspring into families (~83%). Classification success was similarly high into temperature treatments (~83%). A water spiking experiment, whereby enriched 137Ba was added to the water of incubating eggs, also revealed that trace elements from the ambient water are incorporated into developing otoliths during egg incubation. Together, these findings suggest that family-specific chemical signatures in embryonic capelin otoliths may be modified by ambient water chemistry and environmental conditions, particularly temperature, possibly allowing the natal origin of individual capelin to be identified.

Survival and habitat selection of Canada Geese during autumn and winter in metropolitan Chicago, USA

ABSTRACT Winter distribution and resource use of animals is driven by myriad interacting biotic and abiotic factors. Urban areas provide sanctuaries from hunting for game animals and may have thermal benefits during winter through reduced thermoregulatory costs. We deployed cellular GPS transmitters affixed to neck collars of 41 Canada Geese (Branta canadensis) in the Greater Chicago Metropolitan Area (GCMA) of northeastern Illinois, USA, to determine habitat selection and survival during autumn and winter. Canada Geese selected green spaces (59.8%) in greater proportion than available (14%), but they also regularly used industrial urban habitats such as rooftops and rail yards (11.3%), which has not been previously reported. Use of green spaces (−55.8%) decreased and use of industrial urban (+11.4%), riverine (+23.8%), and deep-water habitats (+140.7%) increased as temperatures dropped below the lower critical temperature for Canada Geese (i.e. the temperature at which increased thermoregulatory costs ar...

The genomic basis of cichlid fish adaptation within the deepwater "twilight zone" of Lake Malawi.

Deepwater environments are characterized by low levels of available light at narrow spectra, great hydrostatic pressure, and low levels of dissolved oxygen—conditions predicted to exert highly specific selection pressures. In Lake Malawi over 800 cichlid species have evolved, and this adaptive radiation extends into the “twilight zone” below 50 m. We use population‐level RAD‐seq data to investigate whether four endemic deepwater species (Diplotaxodon spp.) have experienced divergent selection within this environment. We identify candidate genes including regulators of photoreceptor function, photopigments, lens morphology, and haemoglobin, many not previously implicated in cichlid adaptive radiations. Colocalization of functionally linked genes suggests coadapted “supergene” complexes. Comparisons of Diplotaxodon to the broader Lake Malawi radiation using genome resequencing data revealed functional substitutions and signatures of positive selection in candidate genes. Our data provide unique insights into genomic adaptation within deepwater habitats, and suggest genome‐level specialization for life at depth as an important process in cichlid radiation.

Eocene squalomorph sharks (Chondrichthyes, Elasmobranchii) from Antarctica

Rare remains of predominantly deep-water sharks of the families Hexanchidae, Squalidae, Dalatiidae, Centrophoridae, and Squatinidae are described from the Eocene La Meseta Formation, Seymour Island, Antarctic Peninsula, which has yielded the most abundant chondrichthyan assemblage from the Southern Hemisphere to date. Previously described representatives of Hexanchus sp., Squalus weltoni, Squalus woodburnei, Centrophorus sp., and Squatina sp. are confirmed and dental variations are documented. Although the teeth of Squatina sp. differ from other Palaeogene squatinid species, we refrain from introducing a new species. A new dalatiid taxon, Eodalatias austrinalis gen. et sp. nov. is described. This new material not only increases the diversity of Eocene Antarctic elasmobranchs but also allows assuming that favourable deep-water habitats were available in the Eocene Antarctic Ocean off Antarctica in the Eocene. The occurrences of deep-water inhabitants in shallow, near-coastal waters of the Antarctic Peninsula agrees well with extant distribution patterns.

New deep-water brachiopod resilient assemblage from the South-Iberian Palaeomargin (Western Tethys) and its significance for the brachiopod adaptive strategies around the Early Toarcian Mass Extinction Event

The Pliensbachian-Toarcian transition was a period of changes in long-term environmental conditions leading up to the Early Toarcian Mass Extinction Event (ETMEE), which resulted in a noticeable extinction and turnover in the marine biota. The westernmost Tethyan basins, especially the peri-Iberian platforms, provide an exceptional brachiopod record to better understand the adaptive strategies and the severe ecological effects of these faunas within the marine ecosystems. This event marks a critical interval in the evolutionary history of the Phylum Brachiopoda as two orders, the Athyridida and Spiriferinida, became extinct. Evolutionary patterns displayed by several taxa from these groups and some rhynchonellids typifying deep-water habitats are analyzed across this biotic crisis spanning several Mediterranean and NW-European basins. New work performed in La Cerradura section, a deep pelagic trough from the South-Iberian palaeomargin, reveals two new taxa (Koninckodonta sumuntanensis and Atychorhynchia falsiorigo) herein described. This newly documented fauna supports pre-extinction dwarfing and resilience in deep refugia linked to the ETMEE, and an episode of speciation which is interpreted in terms of a pre-extinction radiation. In the ETMEE repopulation phase an opportunistic strategy occurs typified by Soaresirhynchia bouchardi, and a case of homoplasy involving post-extinction pioneers (Elvis taxon) is detected. Similar adaptive strategies occurred associated with other mass extinctions such as the Permian/Triassic and the Cretaceous/Paleocene events, supporting a possible standard pattern in the response of the brachiopod fauna to such biotic crises and shedding light on the ecological effects of the mass extinction events.

Transient deep-water oxygenation in the early Cambrian Nanhua Basin, South China

Many late Neoproterozoic to early Cambrian fossils of multicellular eukaryotes, including those of benthic animals, are found preserved under anoxic and even euxinic bottom-water conditions, which is contradictory to the consensus that oxygen is essential to eukaryotes. To investigate this conundrum, we conducted an integrated study of iron speciation, redox-sensitive trace elements, and Mo isotopes (δ98Mo) on the black shale interval of the lower Cambrian Hetang Formation (∼535–521Ma) at Lantian, South China, in which benthic sponge fossils are abundant in the lower member (LM) but absent in the upper member (UM). Iron speciation data point to uniformly anoxic-ferruginous conditions in the LM and euxinic conditions in the UM, whereas the trace-element and δ98Mo data show greater secular variation in redox conditions. The LM shows higher mean trace element concentrations (Mo: 108ppm, U: 36ppm, V: 791ppm) and lower and more variable δ98Mo (+0.13 to +1.76‰) relative to the UM (Mo: 45ppm, U: 18ppm, V: 265ppm, δ98Mo: +1.59 to +1.67‰), and ratios of redox-sensitive trace element concentrations to total organic carbon are significantly more variable and higher on average in the LM relative to the UM. The appearance of sponge fossils and lower δ98Mo values correlate strongly with gray (i.e., lighter-colored) layers in the LM. These patterns can best be interpreted as recording mainly euxinic conditions throughout deposition of the study units, with more intense background euxinia in the LM relative to the UM, but also with frequent transient oxygenation events in the LM that do not appear in the UM. The transient oxygenation events of the LM led to the initial colonization of the deep Nanhua Basin by sponges, and the termination of these events in the UM caused sponge faunas to disappear until a general rise in O2 levels later in the Cambrian permitted their return to deeper-water habitats. Our study also illustrates that multiple geochemical and paleobiological proxies exhibit different responses in ‘poikiloredox’ environments (i.e., characterized by small-scale spatial and high-frequency temporal variations), which can lead to apparent contradictions between metazoan fossil occurrences and their inferred watermass redox conditions.

Water Quality Analysis in Wetlands Freshwater: Common Floodplain of Jijia-Prut Rivers

Water resources in the Jijia catchment basin are limited and often polluted. The catchment basin of Jijia is situated in northeastern Romania and it crosses the Moldavian Plain on the north-west-south-east direction. The purpose of the present study is to analyze 26 physico-chemical parameters providing the annual and multiannual water quality index. Two water-sampling points were selected: Jijia-Victoria [S.1] and Jijia-Opriseni [S.2]. The high values of nitrates are caused by the use of nitrogen-based chemical fertilizers and of manure. Contamination with nitrites (N-NO2-) and nitrates (N-NO3-) of wetlands and deepwater habitats in the floodplain of Jijia is still high because of agricultural and zootechnical activities. The phosphorus within freshwater habitats is a consequence of anthropogenic pressure: improper storage of animal waste and/or use of phosphates-based fertilizers. Global water quality index (WQi) shows that both monitoring stations are included in the Medium high class.

Demersal fish assemblages on seamounts and other rugged features in the northeastern Caribbean

Recent investigations of demersal fish communities in deepwater (>50m) habitats have considerably increased our knowledge of the factors that influence the assemblage structure of fishes across mesophotic to deep-sea depths. While different habitat types influence deepwater fish distribution, whether different types of rugged seafloor features provide functionally equivalent habitat for fishes is poorly understood. In the northeastern Caribbean, different types of rugged features (e.g., seamounts, banks, canyons) punctuate insular margins, and thus create a remarkable setting in which to compare demersal fish communities across various features. Concurrently, several water masses are vertically layered in the water column, creating strong stratification layers corresponding to specific abiotic conditions. In this study, we examined differences among fish assemblages across different features (e.g., seamount, canyon, bank/ridge) and water masses at depths ranging from 98 to 4060m in the northeastern Caribbean. We conducted 26 remotely operated vehicle dives across 18 sites, identifying 156 species of which 42% of had not been previously recorded from particular depths or localities in the region. While rarefaction curves indicated fewer species at seamounts than at other features in the NE Caribbean, assemblage structure was similar among the different types of features. Thus, similar to seamount studies in other regions, seamounts in the Anegada Passage do not harbor distinct communities from other types of rugged features. Species assemblages, however, differed among depths, with zonation generally corresponding to water mass boundaries in the region. High species turnover occurred at depths <1200m, and may be driven by changes in water mass characteristics including temperature (4.8–24.4°C) and dissolved oxygen (2.2–9.5mg per l). Our study suggests the importance of water masses in influencing community structure of benthic fauna, while considerably adding to the knowledge of mesophotic and deep-sea fish biogeography.

Temperature-based spawning habitat selection by capelin (Mallotus villosus) in Newfoundland

Abstract The location of reproduction is a key life history trait, as it influences the bio-physical conditions that offspring experience and, thus, fitness. Capelin Mallotus villosus is a small (&amp;lt; 200 mm), short-lived forage fish that spawns in two habitats in coastal Newfoundland: warm beaches and cool, deep water (15–40 m). From 2009 to 2014, we investigated temperature-dependent spawning habitat use by quantifying hourly temperature at spawning sites during July within each habitat along with population-level spawning site use. Capelin did not spawn at sites with temperatures &amp;lt;2 °C or consistently &amp;gt;12 °C, supporting the estimated suitable spawning temperature range (2–12 °C). Spawning typically occurred exclusively at beaches early in July when deep-water habitat was too cold (i.e. &amp;lt;2 °C), and then switched to deep-water habitat later in July when beaches became too warm (i.e. &amp;gt;12 °C). Spawning overlapped for 1–3 d in both habitats when temperatures were within suitable ranges (2011, 2013), but capelin also spawned exclusively in one habitat when temperatures remained suitable in the other. The latter suggests that other factors influence spawning habitat selection, such as conspecific egg densities and other environmental conditions. Overall, the flexible use of spawning habitats, along with wide tolerance ranges of capelin eggs to environmental conditions, are likely key adaptations to maximize fitness and stabilize population dynamics of this important forage fish species in an unpredictable environment.

Squamation and ecology of thelodonts.

Thelodonts are an enigmatic group of Paleozoic jawless vertebrates that have been well studied from taxonomical, biostratigraphic and paleogeographic points of view, although our knowledge of their ecology and mode of life is still scant. Their bodies were covered by micrometric scales whose morphology, histology and the developmental process are extremely similar to those of extant sharks. Based on these similarities and on the well-recognized relationship between squamation and ecology in sharks, here we explore the ecological diversity and lifestyles of thelodonts. For this we use classic morphometrics and discriminant analysis to characterize the squamation patterns of a significant number of extant shark species whose ecology is well known. Multivariate analyses have defined a characteristic squamation pattern for each ecological group, thus establishing a comparative framework for inferring lifestyles in thelodonts. We then use this information to study the squamation of the currently described 147 species of thelodonts, known from both articulated and disarticulated remains. Discriminant analysis has allowed recognizing squamation patterns comparable to those of sharks and links them to specific ecological groups. Our results suggest a remarkable ecological diversity in thelodonts. A large number of them were probably demersal species inhabiting hard substrates, within caves and crevices in rocky environments or reefs, taking advantage of the flexibility provided by their micromeric squamations. Contrary to classical interpretations, only few thelodonts were placed among demersal species inhabiting sandy and muddy substrates. Schooling species with defensive scales against ectoparasites could be also abundant suggesting that social interactions and pressure of ectoparasites were present in vertebrates as early the Silurian. The presence of species showing scales suggestive of low to moderate speed and a lifestyle presumably associated with open water environments indicates adaptation of thelodonts to deep water habitats. Scale morphology suggests that some other thelodonts were strong-swimming pelagic species, most of them radiating during the Early Devonian in association with the Nekton Revolution.