Macrophagous Predators Research Articles

Growth and feeding ecology of coniform conodonts.

Conodonts were the first vertebrates to develop mineralized dental tools, known as elements. Recent research suggests that conodonts were macrophagous predators and/or scavengers but we do not know how this feeding habit emerged in the earliest coniform conodonts, since most studies focus on the derived, ‘complex’ conodonts. Previous modelling of element position and mechanical properties indicate they were capable of food processing. A direct test would be provided through evidence of in vivo element crown tissue damage or through in vivo incorporated chemical proxies for a shift in their trophic position during ontogeny. Here we focus on coniform elements from two conodont taxa, the phylogenetically primitive Proconodontus muelleri Miller, 1969 from the late Cambrian and the more derived Panderodus equicostatus Rhodes, 1954 from the Silurian. Proposing that this extremely small sample is, however, representative for these taxa, we aim to describe in detail the growth of an element from each of these taxa in order to the test the following hypotheses: (1) Panderodus and Proconodontus processed hard food, which led to damage of their elements consistent with prey capture function; and (2) both genera shifted towards higher trophic levels during ontogeny. We employed backscatter electron (BSE) imaging, energy-dispersive X-ray spectroscopy (EDX) and synchrotron radiation X-ray tomographic microscopy (SRXTM) to identify growth increments, wear and damage surfaces, and the Sr/Ca ratio in bioapatite as a proxy for the trophic position. Using these data, we can identify whether they exhibit determinate or indeterminate growth and whether both species followed linear or allometric growth dynamics. Growth increments (27 in Pa. equicostatus and 58 in Pr. muelleri) were formed in bundles of 4–7 increments in Pa. equicostatus and 7–9 in Pr. muelleri. We interpret the bundles as analogous to Retzius periodicity in vertebrate teeth. Based on applied optimal resource allocation models, internal periodicity might explain indeterminate growth in both species. They also allow us to interpret the almost linear growth of both individuals as an indicator that there was no size-dependent increase in mortality in the ecosystems where they lived e.g., as would be the case in the presence of larger predators. Our findings show that periodic growth was present in early conodonts and preceded tissue repair in response to wear and damage. We found no microwear and the Sr/Ca ratio, and therefore the trophic position, did not change substantially during the lifetimes of either individual. Trophic ecology of coniform conodonts differed from the predatory and/or scavenger lifestyle documented for “complex” conodonts. We propose that conodonts adapted their life histories to top-down controlled ecosystems during the Nekton Revolution.

Non-ammocoete larvae of Palaeozoic stem lampreys.

Ammocoetes-the filter-feeding larvae of modern lampreys-have long influenced hypotheses of vertebrate ancestry1-7. The life history of modern lampreys, whichdevelop from a superficially amphioxus-like ammocoete to a specialized predatory adult, appears to recapitulate widely accepted scenarios of vertebrate origin. However, no direct evidence has validated the evolutionary antiquity of ammocoetes, and their status as models of primitivevertebrate anatomy is uncertain. Here we report larval and juvenile forms of four stem lampreys from the Palaeozoic era (Hardistiella, Mayomyzon, Pipiscius, and Priscomyzon), including a hatchling-to-adult growth series of the genus Priscomyzon from Late Devonian Gondwana. Larvae of all four genera lack the defining traits of ammocoetes. They instead display features that are otherwise unique to adult modern lampreys, including prominent eyes, a cusped feeding apparatus, and posteriorly united branchial baskets. Notably, phylogenetic analyses find that these non-ammocoete larvae occur in at least three independent lineages of stem lamprey. This distribution strongly implies that ammocoetes are specializations of modern-lamprey life history rather than relics of vertebrate ancestry. These phylogenetic insights also suggest that the last common ancestor of hagfishes and lampreys was a macrophagous predator that did not have a filter-feeding larval phase. Thus, the armoured 'ostracoderms' that populate the cyclostome and gnathostome stems might serve as better proxies than living cyclostomes for the last common ancestor of all living vertebrates.

Disparate compound eyes of Cambrian radiodonts reveal their developmental growth mode and diverse visual ecology.

Radiodonts are nektonic stem-group euarthropods that played various trophic roles in Paleozoic marine ecosystems, but information on their vision is limited. Optical details exist only in one species from the Cambrian Emu Bay Shale of Australia, here assigned to Anomalocaris aff. canadensis We identify another type of radiodont compound eye from this deposit, belonging to 'Anomalocaris' briggsi This ≤4-cm sessile eye has >13,000 lenses and a dorsally oriented acute zone. In both taxa, lenses were added marginally and increased in size and number throughout development, as in many crown-group euarthropods. Both species' eyes conform to their inferred lifestyles: The macrophagous predator A. aff. canadensis has acute stalked eyes (>24,000 lenses each) adapted for hunting in well-lit waters, whereas the suspension-feeding 'A.' briggsi could detect plankton in dim down-welling light. Radiodont eyes further demonstrate the group's anatomical and ecological diversity and reinforce the crucial role of vision in early animal ecosystems.

Convergent Evolution of Swimming Adaptations in Modern Whales Revealed by a Large Macrophagous Dolphin from the Oligocene of South Carolina

Modern whales and dolphins are superbly adapted for marine life, with tail flukes being a key innovation shared by all extant species. Some dolphins can exceed speeds of 50km/h, a feat accomplished by thrusting the flukes while adjusting attack angle with their flippers [1]. These movements are driven by robust axial musculature anchored to a relatively rigid torso consisting of numerous short vertebrae, and controlled by hydrofoil-like flippers [2-7]. Eocene skeletons of whales illustrate the transition from semiaquatic to aquatic locomotion, including development of a fusiform body and reduction of hindlimbs [8-11], but the rarity of Oligocene whale skeletons [12, 13] has hampered efforts to understand the evolution of fluke-powered, but forelimb-controlled, locomotion. We report a nearly complete skeleton of the extinct large dolphin Ankylorhiza tiedemani comb. n. from the Oligocene of South Carolina, previously known only from a partial rostrum. Its forelimb is intermediate in morphology between stem cetaceans and extant taxa, whereas its axial skeleton displays incipient rigidity at the base of the tail with a flexible lumbar region. The position of Ankylorhiza near the base of the odontocete radiation implies that several postcranial specializations of extant cetaceans, including a shortened humerus, narrow peduncle, and loss of radial tuberosity, evolved convergently in odontocetes and mysticetes. Craniodental morphology, tooth wear, torso vertebral morphology, and body size all suggest that Ankylorhiza was a macrophagous predator that could swim relatively fast, indicating that it was one of the few extinct cetaceans to occupy a niche similar to that of killer whales.

Calcium stable isotopes place Devonian conodonts as first level consumers

Conodont animals are an extinct group of jawless vertebrates whose hard parts, also known as conodont elements, represent the earliest evidence of a mineralised skeleton in the vertebrate lineage. Conodont elements are interpreted as parts of a feeding apparatus, which together with the presence of eyes and microwear patterns, support the controversial hypothesis that conodont animals were macrophagous predators and/or scavengers. Here, we explore the trophic position of five conodont genera (Palmatolepis, Polygnathus, Ancyrodella, Ancyrognathus and Icriodus) from five contemporary Late Devonian sites distributed worldwide (France, Morocco, Vietnam and Australia) by means of calcium (Ca) stable isotope compositions. The seawater Ca isotope composition was calibrated using contemporary Late Devonian brachiopod isotopic values. By comparison with extant marine trophic chain composed of cartilaginous fish, conodont Ca isotope compositions are indicative of a zooplanktivore-primary piscivore niche, with no genus-specific trophic pattern. The question of active predation or scavenging cannot be resolved definitively but our results strongly suggest that Late Devonian conodonts were first level consumers.

Observations and Experiments on the Biology and Life History of Riseriellus occultus (Heteronemertea: Lineidae).

Studies on the biology and life history of nemerteans are scarce, mostly because these animals are nocturnal. In order to broaden the knowledge base on the life history of nemerteans as a prerequisite for comparative analyses, we studied a population of Riseriellus occultus (Heteronemertea: Lineidae) inhabiting the rocky intertidal in southern Brittany near Concarneau (France) for more than 10 years. Our studies show that R. occultus is an iteroparous, perennial species exclusively inhabiting rocky shore crevices that result from onionskin weathering of the granite. From September through October R. occultus reproduces by external fertilization and develops via a planktonic pilidium larva, which, under laboratory conditions, metamorphoses after about six weeks. Adults of R. occultus are nocturnal macrophagous predators that preferentially feed on the gastropods Gibbula umbilicalis and Patella species, but also consume the bivalve Mytilus edulis. Since R. occultus devours the snail inside the shell, we fixed individuals while feeding, and serially sectioned them. Reconstruction of the sections shows that R. occultus swallows the entire soft body and finally detaches the columellar muscle from the shell. Estimates on the density of R. occultus inside the rock crevices provide evidence for clustered distribution and locally high abundance on the rocky shore. These data strongly suggest that R. occultus affects the structure of the rocky shore gastropod community. Although our data are still fragmentary with respect to the ecology of this species and its role in the local food web, our knowledge has grown to such extent that R. occultus can now be regarded as one of the few well characterized nemertean species.

The Mitochondrial Genome of Erpobdella octoculata (Hirudinida: Arhynchobdellida), as the First Representative for the Suborder Erpobdelliformes

Erpobdella octoculata Linnaeus, 1758 (Hirudinea: Arhynchobdellida: Erpobdellidae) is a very common and morphologically variable macrophagous predators of aquatic invertebrates. Here we determined the complete mitochondrial DNA (mtDNA) sequence of this species, as the first representative of the suborder Erpobdelliformes. This genome is 14,407 bp in length with an A+T content of 71.55%, containing 37 typical animal mitochondrial genes and a non-coding region (NCR). All genes were arranged in the same order as most reported annelid. A remarkable strand bias found for all protein coding genes (PCGs) was positive AT-skew and negative GC-skew. The models of secondary structures for the two rRNA genes of this species were predicted. rrnL consisted of six domains and 46 helices, while three domains and 34 helices for rrnS. The secondary structures of some transfer RNA genes were relatively special. Bayesian phylogenetic and maximum likelihood analyses, based on concatenated nucleotide sequences of 12 PCGs, placed E. octoculata within the suborder Hirudiniformes, and the relationships depicted within that order were not quite in agreement with previous morphological classifications. The complete mtDNA sequence of E. octoculata provides important genetic markers for identification, ecological and evolutionary studies of leeches.

The complete mitochondrial genome sequence of Erpobdella octoculata (Hirudinea: Arhynchobdellida: Erpobdellidae)

Erpobdella octoculata (Linnaeus, 1758; Hirudinea: Arhynchobdellida: Erpobdellidae) is a very common and morphologically variable macrophagous predators of aquatic invertebrates. Here we determined the complete mitochondrial DNA (mtDNA) sequence of this species, as the first representative of the suborder Erpobdelliformes. This genome is 14,407 bp in length with an A + T content of 71.55%, containing 37 typical animal mitochondrial genes and a non-coding region (NCR). It has high AT content and the same gene arrangement pattern as those of typical annelids. The complete mtDNA sequence of E. octoculata provides useful genetic markers for identification, ecological and evolutionary studies of leeches.

A bizarre new toothed mysticete (Cetacea) from Australia and the early evolution of baleen whales

Extant baleen whales (Cetacea, Mysticeti) are all large filter-feeding marine mammals that lack teeth as adults, instead possessing baleen, and feed on small marine animals in bulk. The early evolution of these superlative mammals, and their unique feeding method, has hitherto remained enigmatic. Here, I report a new toothed mysticete from the Late Oligocene of Australia that is more archaic than any previously described. Unlike all other mysticetes, this new whale was small, had enormous eyes and lacked derived adaptations for bulk filter-feeding. Several morphological features suggest that this mysticete was a macrophagous predator, being convergent on some Mesozoic marine reptiles and the extant leopard seal (Hydrurga leptonyx). It thus refutes the notions that all stem mysticetes were filter-feeders, and that the origins and initial radiation of mysticetes was linked to the evolution of filter-feeding. Mysticetes evidently radiated into a variety of disparate forms and feeding ecologies before the evolution of baleen or filter-feeding. The phylogenetic context of the new whale indicates that basal mysticetes were macrophagous predators that did not employ filter-feeding or echolocation, and that the evolution of characters associated with bulk filter-feeding was gradual.

Origins and Early Evolution of Predation

Predation, in the broad sense of an organism killing another organism for nutritional purposes, is probably as old as life itself and has originated many times during the history of life. Although little of the beginnings is caught in the fossil record, observations in the rock record and theoretical considerations suggest that predation played a crucial role in some of the major transitions in evolution. The origin of eukaryotic cells, poorly constrained to about 2.7 Ga by geochemical evidence, was most likely the ultimate result of predation among prokaryotes. Multicellularity (or syncytiality), as a means of acquiring larger size, is visible in the fossil record soon after 2 Ga and is likely to have been mainly a response to selective pressure from predation among protists. The appearance of mobile predators on bacteria and protists may date back as far as 2 Ga or it may be not much older than the Cambrian explosion, or about 600 Ma. The combined indications from the decline of stromatolites and the diversification of acritarchs, however, suggest that such predation may have begun around 1 Ga. The Cambrian explosion, culminating around 550 Ma, represents the transition from simple, mostly microbial, ecosystems to ones with complex food webs and second- and higher-order consumers. Macrophagous predators were involved from the beginning, but it is not clear whether they originated in the plankton or in the benthos. Although predation was a decisive selective force in the Cambrian explosion, it was a shaper rather than a trigger of this evolutionary event.

Pond tadpoles with generalized morphology: is it time to reconsider their functional roles in aquatic communities?

With rare exceptions, anuran larvae have traditionally been considered to occupy lower trophic levels in aquatic communities where they function as microphagous suspension feeders. This view is being challenged by studies showing that tadpoles with generalized morphology often function as macrophagous predators. Here, we review the literature concerning macrophagy by tadpoles and provide two additional examples involving generalized tadpoles. In the first, we demonstrate with laboratory and field experiments that wood frog (Rana sylvatica) tadpoles are major predators of macroinvertebrates in ponds. In the second, we show that green frog (R. clamitans) tadpoles can cause catastrophic reproductive failure of the wood frog via egg predation. These results and data from other studies challenge the assumption that generalized tadpoles function as filter-feeding omnivores, and question the general applicability of community organization models which assume that predation risk increases with pond permanence. We suggest that predation risk is greater in temporary ponds than in more permanent ponds for many organisms that are vulnerable to predation by tadpoles. This being so, a conditional model based upon interactions that are species-specific, life-stage-specific, and context-dependent may better explain community organization along hydrological gradients than models which assume that temporary ponds have few or no predators.

Upper Placentian—Branchian series of mainland Nova Scotia (middle-upper Lower Cambrian): Faunas, paleoenvironments, and stratigraphic revision

Lithostratigraphy and depositional and epeirogenic history of the upper Placentian Series (Cuslett-Fosters Point Formations of the Bonavista Group) and Branchian Series (Brigus Formation) are identical in the northern Antigonish Highlands; Cape Breton Island; and eastern Placentia Bay, southeastern Newfoundland. Preliminary evidence suggests that the lower Middle Cambrian is present in the field area. A unified, uppermost Precambrian–Lower Cambrian, formation- and member-level nomenclature is appropriate to Avalonian North America, and the stratigraphic nomenclature of southeastern Newfoundland is applied in northern mainland Nova Scotia.Latest Placentian shoaling and deposition of a peritidal carbonate lithosome and unconformable onlap of the trilobite-bearing Branchian Series occurred in shallow Avalonian shale basins from eastern Massachusetts to central England.Uppermost Placentian Series faunas are very diverse in the Fosters Point Formation. Limited similarities with the South Australian Lower Cambrian are indicated by the presence ofCamenellasp. cf.C. reticulosa, Conotheca australiensis, andHyptiothecasp., but these forms do not contribute to highly resolved correlation.Twenty-eight taxa are illustrated from the upper Placentian and Branchian Series.Caveacus rectusn. gen. and sp., a phosphatic problematicum, is limited to the upper Placentian Series. The oldest, skeletalized, macrophagous predators are the Pseudoconodontida and the later appearing Protoconodontida (n. orders). The Pseudoconodontida includes the Protohertzinacea n. superfamily and Strictocorniculacea n. superfamily (with the Rhombocorniculidae and Strictocorniculidae n. families).Strictocorniculum vanallerumn. gen. and sp. is described. The tommotiid family Sunnaginiidae emend. includesEccentrotheca, Sunnaginia, Kulparina, andJayceia deltiformisn. gen. and sp.

Breeding Habitat Segregation of Wood Frogs and American Toads: The Role of Interspecific Tadpole Predation and Adult Choice

In western North Carolina, adult wood frogs and American toads often occur syntopically but segregate with respect to breeding habitat. American toads arrive at breeding ponds 4-10 wk after wood frogs and typically select breeding sites that lack wood frog tadpoles. In an experiment conducted in seminatural ponds, adult toads completely avoided ovipositing in ponds with wood frog tadpoles. Additional experiments strongly suggest that predation of wood frog tadpoles on toad eggs and larvae is the primary reason for the evolution of adult avoidance behavior in toads. In one set of experiments, toad eggs and hatchlings suffered complete mortality within two days after being placed in artificial ponds containing natural densities of wood frog tadpoles. Similar mortality occurred when eggs and hatchlings were introduced into natural breeding sites. In one experiment in which 2000 toad hatchlings were added to natural ponds, all were killed and consumed within 45 min by wood frog tadpoles. Although tadpoles of most anurans are considered to be microphagous suspension feeders that occupy lower trophic levels of food chains, a growing body of evidence indicates that many tadpoles can temporarily shift feeding modes to become upper level, macrophagous predators. Because facultative predators such as wood frog tadpoles often reach far higher densities in ponds than do top predators, their role in structuring aquatic communities via opportunistic predation may be more important than previously realized.