Hydrodynamic performance of the flippers of large‐bodied cetaceans in relation to locomotor ecology

Cetaceans evolved flippers that are unique in both size and shape probably due to selection pressures associated with foraging and body size. Flippers function as control surfaces for maneuverability and stability. Flippers of cetaceans and engineered hydrofoils are similar with streamlined cross‐sections and wing‐like planforms, which affect lift, drag and hydrodynamic efficiency. Scale models of the flippers from large‐bodied (body length > 6 m) cetaceans (fin whale, killer whale, sperm whale) were constructed from computed tomography (CT) scans of flippers. Flipper planforms were highly tapered for the fin whale, a rounded, paddle‐like design for the killer whale, and a square geometry for the sperm whale. Hydrodynamic properties of the models at varying angles of attack (?40º to 40^o) were determined in a water tunnel with a multi‐axis load cell. The flippers were found to have hydrodynamic characteristics similar to engineered wings. Differences in flipper morphology of large‐bodied cetaceans and their hydrodynamic performance are associated with the requirements of aquatic locomotion involved with ecology of the whales. The flippers of the killer whale provided the greatest maneuverability, whereas the flippers of the fin whale had low drag for lunging and the flippers of the sperm whale provided lift for diving.     

Hydrodynamic flow control in marine mammals

The ability to control the flow of water around the body dictates the performance of marine mammals in the aquatic environment. Morphological specializations of marine mammals afford mechanisms for passive flow control. Aside from the design of the body, which minimizes drag, the morphology of the appendages provides hydrodynamic advantages with respect to drag, lift, thrust, and stall. The flukes of cetaceans and sirenians and flippers of pinnipeds possess geometries with flexibility, which enhance thrust production for high efficiency swimming. The pectoral flippers provide hydrodynamic lift for maneuvering. The design of the flippers is constrained by performance associated with stall. Delay of stall can be accomplished passively by modification of the flipper leading edge. Such a design is exhibited by the leading edge tubercles on the flippers of humpback whales (Megaptera novaeangliae). These novel morphological structures induce a spanwise flow field of separated vortices alternating with regions of accelerated flow. The coupled flow regions maintain areas of attached flow and delay stall to high angles of attack. The delay of stall permits enhanced turning performance with respect to both agility and maneuverability. The morphological features of marine mammals for flow control can be utilized in the biomimetic design of engineered structures for increased power production and increased efficiency.     

TURNING MANEUVERS IN STELLER SEA LIONS (EUMATOPIAS JUBATUS)

Steller sea lions are highly maneuverable marine mammals (expressed as minimum turning radius). Video recordings of turns ( n = 195) are analyzed from kinematic measurements for three captive animals. Speed-time plots of 180° turns have a typical "V-shape." The sea lions decelerated during the first half of the turn, reached a minimum speed in the middle of the curved trajectory and reaccelerated by adduction of the pectoral flippers. The initial deceleration was greater than that for passive gliding due to pectoral flipper braking and/or change in body contour from a stiff, straight streamlined form. Centripetal force and thrust were determined from the body acceleration. Most thrust was produced during the power phase of the pectoral flipper stroke cycle. Contrary to previous findings on otariids, little or no thrust was generated during initial abduction of the pectoral flippers and during the final drag-based paddling phase of the stroke cycle. Peak thrust force at the center of gravity occurs halfway through the power phase and the centripetal force is maximal at the beginning of the power stroke. Performance is modulated by changes in the duration and intensity of movements without changing their sequence. Turning radius, maximum velocity, maximum acceleration and turning duration were 0.3 body lengths, 3.5 m/s, 5 m/s², and 1.6 s, respectively. The relative maneuverability based on velocity and length specific minimum turning radius is comparable to other otariids, superior to cetaceans but inferior to many fish.     

Hypoxic Reptiles: Blood Gases, Body Temperature and Control of Breathing

Our contribution to this symposium is a review of recent modelsand experimental cdata on oxygen homeostasis in vertebrateswith normal intracardiac shunts; i.e., amphibians and reptiles.We focus on the interactions among hemoglobin function, bodytemperature regulation, and cardiovascular shunts under normalconditions (i.e., breathing fresh air at or near sea level)and during external hypoxia (e.g., altitude, burrows) and internalhypoxia (e.g., anemia, hemorrhage). Mathematical models andexperimental data suggest that animals with venous admixturefrom cardiovascular shunts will show biphasic arterial and mixedvenous Po₂ responses to warming; i.e., first increasing andthen, as the dissociation curve shifts too far to the right,decreasing. This has implications for many physiological functionsincluding oxygen consumption by tissues, control of breathing,as well as preferred body temperature and its regulation. Wepresent some of the recent experiments that have explored theseimplications.     

The scaling of locomotor performance in predator-prey encounters: from fish to killer whales.

During predator-prey encounters, a high locomotor performance in unsteady manoeuvres (i.e. acceleration, turning) is desirable for both predators and prey. While speed increases with size in fish and other aquatic vertebrates in continuous swimming, the speed achieved within a given time, a relevant parameter in predator-prey encounters, is size independent. In addition, most parameters indicating high performance in unsteady swimming decrease with size. Both theoretical considerations and data on acceleration suggest a decrease with body size. Small turning radii and high turning rates are indices of maneuverability in space and in time, respectively. Maneuverability decreases with body length, as minimum turning radii and maximum turning rates increase and decrease with body length, respectively. In addition, the scaling of linear performance in fish locomotion may be modulated by turning behaviour, which is an essential component of the escape response. In angelfish, for example, the speed of large fish is inversely related to their turning angle, i.e. fish escaping at large turning angles show lower speed than fish escaping at small turning angles. The scaling of unsteady locomotor performance makes it difficult for large aquatic vertebrates to capture elusive prey by using whole-body attacks, since the overall maneuverability and acceleration of small prey is likely to be superior to that of large predators. Feeding strategies in vertebrate predators can be related to the predator-prey length ratios. At prey-predator ratios higher than approximately 10(-2), vertebrate predators are particulate feeders, while at smaller ratios, they tend to be filter feeders. At intermediate ratios, large aquatic predators may use a variety of feeding methods that aid, or do not involve, whole body attacks. Among these are bubble curtains used by humpback whales to trap fish schools, and tail-slapping of fish by delphinids. Tail slapping by killer whales is discussed as an example of these strategies. The speed and acceleration achieved by the flukes of killer whales during tail slaps are higher and comparable, respectively, to those that can be expected in their prey, making tail-slapping an effective predator behaviour.     

Potential Abiotic and Biotic Impacts of Zebra Mussels on the Inland Waters of North America

SYNOPSIS. The expansion of zebra mussel distribution into inlandwaterways of North America portends significant abiotic andbiotic changes mediated either directly or indirectly by Dreissena.Dreissena fouls a wide array of submerged substrates includingrock surfaces, macrophytes, native molluscs, canal and dockwalls, and watercraft and motor outdrives. Fouling of waterintake pipes and associated installations can severely impairwater delivery to hydroelectric, municipal and industrial users,necessitating proactive or reactive control measures. Musselsincrease water clarity by removing suspended clay, silt, bacteria,phytoplankton, and small zooplankton. Clear water phases associatedwith Dreissena grazing may exceed in magnitude and durationthose generated by zooplankton grazing. Enhanced water clarityincreases light transmittance and growth of benthic plants.Some benthic invertebrates {e.g., unionid molluscs) are adverselyaffected by Dreissena, whereas others, including amphipod crustaceans,exploit structure associated with or wastes generated by zebramussels. Dreissena is exploited by a host of predators, mostnotably waterfowl, fish and crayfish. Waterfowl predators thatconsume contaminated Dreissena have elevated concentrationsof organic pesticides and polychlorinated biphenyl compounds.Invasion of shallow lakes and ponds by Dreissena may divertproduction and biomass from pelagic to benthic foodwebs, shiftingecosystems to an alternative state.     

Lymph Node Structure. An Ontogenetic Explanation for Divergence in Eutheria, Metatheria, and Prototheria

The development of the mesenteric lymph node was studied inthe polyprotodont marsupial, Marmosa mitis. A primary mesenchymalradiation from the mesenteric arterial investment into the surroundingmesenteric lymph space yielded reticulum having lymphoid affinity.A secondary radiation yielded the supporting connective tissues,i.e., the node's hilum and trabeculae. This secondary radiationis limited to eutherian and metatherian mammals; its absencein prototheria explains both the absence of complex lymph nodesin these mammals and the singular follicles found in their lymphvessels. The mesenchymal source indicates that complex lymphnodes, like splenic follicles, derive from mesenchyme associatedwith portions of the arterial tree. The ensuing evolution fromsource mesenchyme to mature nodal configuration is describedin a tentative histogenetic scheme.     

Stability versus maneuverability in aquatic locomotion

The dictionary definition of stability as "Firmly established,not easily to be changed" immediately indicates the conflictbetween stability and maneuverability in aquatic locomotion.The present paper addresses several issues resulting from theseopposing requirements. Classical stability theory for bodiesmoving in fluids is based on developments in submarine and airshipmotions. These have lateral symmetry, in common with most animals.This enables the separation of the equations of motion intotwo sets of 3 each. The vertical (longitudinal) set, which includesmotions in the axial (surge), normal (heave) and pitching directions,can thus be separated from the lateral-horizontal plane whichincludes yaw, roll and sideslip motions. This has been founduseful in the past for longitudinal stability studies basedon coasting configurations but is not applicable to the analysisof turning, fast starts and vigorous swimming, where the lateralsymmetry of the fish body is broken by bending motions. Thepresent paper will also examine some of the aspects of the stabilityvs. maneuverability tradeoff for these asymmetric motions. Ananalysis of the conditions under which the separation of equationsof motions into vertical and horizontal planes is justified,and a definition of the equations to be used in cases wherethis separation is not accurate enough is presented.     

Differences in Endogenous Levels of Gibberellin Activity in Male and Female Partners of Two Dioecious Tree Species

It was found that in two dioecious species, the carob (Ceratoniasiliqua) and the date palm (Phoenix dactylifera) growing intheir natural Mediterrancan habitat, the female partner invariablymanifested higher endogenous GN-like activity. While this trendwas consistent, marked seasonal fluctuations in both free andglycosidic bound-GN were observedhigher levels being associatedwith essentially vegetative spring growth and the converse associatedwith anthesis. It is postulated that while a clear correlationexists between female growth and high GN activity in these species,the switch from vegetative to generative growth in both themale and female may be associated with a lower relative proportionof endogenous GN in a specific multi-hormonal complex presumablyrequired for flower induction.     

The effect of intra- and interspecific aggression on patch residence time in Negev Desert gerbils: a competing risk analysis

We observed patch-use behavior by two gerbil species in a fieldsetting and investigated how aggression and intrinsic decision-makinginteract to influence patch residence times. Results were interpretedby using a competing risk analysis model, which uniquely enabledus to estimate the intrinsic patch-leaving decisions independentlyof external interruptions of foraging bouts by aggression. Theexperiment was conducted in two 1-ha field enclosures completelysurrounded by rodent-proof fences and included allopatric (onlyGerbillus andersoni allenbyi) and sympatric (G. a. allenbyiand G. pyramidum) treatments. We predicted that increased foodpatch quality (i.e., habitat quality) should decrease intrinsicpatch-leaving rates and increase rates of aggressive interactionsinvolving the forager feeding in the patch (i.e., the occupantindividual). We also anticipated that increasing populationdensity should result in an increase in the rate of aggressiveinteractions involving the occupant individual. Our resultssupported the first two predictions, indicating a trade-offbetween foraging and aggression. However, the third predictionwas realized only for G. a. allenbyi in allopatry. Furthermore,in allopatry, occupant G. a. allenbyi individuals with highcompetitive ranks were involved in aggressive interactions atlower rates than those with low competitive ranks. However,in sympatry, patch-use behavior of occupant G. a. allenbyi individualswas mainly influenced by aggressive behavior of G. pyramidum,which did not respond to their competitive rank. Thus, it shouldpay less for G. a. allenbyi to be aggressive in sympatric populations.The observed reduction in intraspecific aggression among individualG. a. allenbyi in the presence of G. pyramidum supports thisassertion. We suggest that this reduction likely weakens thenegative effect of intra- and interspecific density on the percapita growth rate of G. a. allenbyi. Because this would changethe slope of the isocline of G. a. allenbyi, it could be animportant mechanism promoting coexistence when habitat selectionis constrained.     

The prevalence and implications of copepod behavioral responses to oceanographic gradients and biological patchiness

Several species and developmental stages of calanoid copepodswere tested for responses to environmental cues in a laboratoryapparatus that mimicked conditions commonly associated withpatches of food in the ocean. All species responded to the presenceof phytoplankton by feeding. All species responded by increasingproportional residence time in one, but not both, of the treatmentsdefined by gradients of velocity or density. Most species increasedswimming speed and frequency of turning in response to the presenceof chemical exudates or gradients of velocity. Only one species,Eurytemora affinis, increased proportional time of residencein response to gradients in density of the water. Responsesof E. affinis to combined cues did not definitively demonstratea hierarchical use of different cues as previously observedfor Temora longicornis and Acartia tonsa. A simple foragingsimulation was developed to assess the applicability in thefield of the behavioral results observed in the laboratory.These simulations suggest that observed fine-scale behaviorscould lead to copepod aggregations observed in situ. The presentstudy demonstrates that behavioral response to cues associatedwith fine-scale oceanographic gradients and biological patchinessis functionally important and prevalent among copepods and likelyhas significant impacts on larger-scale distributional patterns.     

Laboratory studies on the larval growth and development of Nyctiphanes capensis (Euphausiacea)

Larvae of Nyctiphanes capensis Hansen were reared in the laboratoryunder different trophic conditions (i.e., algal, animal andmixed diets) from the second calyptopis stage to the first juvenilestage. It is demonstrated that diet is important in determininggrowth rates and development of the larvae. The highest growthrates were achieved on diets of Artemia nauplii mixed with eitherthe flagellate Tetraselmis or Pseudodiaptomus nauplii. Poorerdietary conditions such as with the diatom Phaeodactylum yieldedslower growth rates where additional moults were needed to completefurcilia development. Under favourable trophic conditions theintermoult period was 4–6 days and was not age dependent.Three dominant pathways of pleopod development were found inhealthy individuals.     

Predation impact of Cyclops vicinus on the rotifer community in Lake Constance in spring

The predation impact of Cyclops vicinus on rotifers was studiedunder near-natural conditions in small enclosures to evaluatewhether copepod predation is responsible for the decline ofrotifers in Lake Constance in spring. Cyclops vicinus fed selectivelyon Synchaeta spp.; Keratella and Polyarthra spp. were not selectedfor. Predation rates increased with prey density up to a maximumof 37 Synchueta day^–1 at a density of 1.6 x 10⁶ Synchaetam^–2, i.e. at -1200 Synchaeta l^–1. Calculation ofcropping rates suggests that Cyclops alone can control the abundanceof Synchaeta in spring, i.e. that mainly Cyclops is responsiblefor the decline of Synchaeta species in Lake Constance in May.     

Spatio-temporal structure of tidally mixed coastal waters: variability and heterogeneity

In the tidally mixed coastal waters of the eastern English Channel,the vertical properties of temperature, salinity, in vivo fluorescence,turbidity and light were investigated along an inshore-offshoretransect, sampled in different tidal conditions. The verticaldistribution of these parameters was then characterized in termsof mean value, variability (i.e. standard deviation) and heterogeneity(i.e. fractal dimension), regarded as a quantitative characterizationof the structure of the vertical variability of both physicaland biological parameters. These different estimates were thensubjected to multivariate spatio-temporal analysis and showeda very specific spatio-temporal structure suggesting a differentialcontrol of the vertical properties of water masses associatedwith both inshore-offshore gradient and tidal advection. Inparticular, the fractal dimension (i.e. heterogeneity) of invivo fluorescence is higher during ebb for offshore waters,suggesting a density dependence of the phytoplankton biomassstructure. In contrast, the vertical variability of fluorescenceis higher during flood for inshore waters, leading to an inverserelationship between variability and heterogeneity of the verticaldistribution of phytoplankton biomass.     

Adaptive Evolution of the Hox Gene Family for Development in Bats and Dolphins

Bats and cetaceans (i.e., whales, dolphins, porpoises) are two kinds of mammals with unique locomotive styles and occupy novel niches. Bats are the only mammals capable of sustained flight in the sky, while cetaceans have returned to the aquatic environment and are specialized for swimming. Associated with these novel adaptations to their environment, various development changes have occurred to their body plans and associated structures. Given the importance of Hox genes in many aspects of embryonic development, we conducted an analysis of the coding regions of all Hox gene family members from bats (represented by Pteropus vampyrus, Pteropus alecto, Myotis lucifugus and Myotis davidii) and cetaceans (represented by Tursiops truncatus) for adaptive evolution using the available draft genome sequences. Differences in the selective pressures acting on many Hox genes in bats and cetaceans were found compared to other mammals. Positive selection, however, was not found to act on any of the Hox genes in the common ancestor of bats and only upon Hoxb9 in cetaceans. PCR amplification data from additional bat and cetacean species, and application of the branch-site test 2, showed that the Hoxb2 gene within bats had significant evidence of positive selection. Thus, our study, with genomic and newly sequenced Hox genes, identifies two candidate Hox genes that may be closely linked with developmental changes in bats and cetaceans, such as those associated with the pancreatic, neuronal, thymus shape and forelimb. In addition, the difference in our results from the genome-wide scan and newly sequenced data reveals that great care must be taken in interpreting results from draft genome data from a limited number of species, and deep genetic sampling of a particular clade is a powerful tool for generating complementary data to address this limitation.     

Bicapitate Nitzschia species: abundant nanoplankton in aggregates during November-December (l992) in the equatorial Pacific

From 12°S to 9°N-140°W on the benthic cruise ofthe US Joint Global Ocean Flux Studies (JGOFS), horizontal near-surfacenet tows (n = 18) and vertical net hauls (n = 15, 200-0 m) yieldedhigh relative abundances of small, bicapitate, intact Nitzschiavalves (average 59.0 and 56.0% of total diatoms, respectively),although >90% of them were too small to have been caughteffectively with the net used. This complex was much less common(average 3.2% of diatoms) in the surface sediment samples (n= 7, 12°S-9°N, below water depths of 4269-4991 m). Examinationunder the light microscope showed concentrations of cells associatedwith organic detritus, planar and cylindrical membranous structures,fecal pellets, and occasionally on other diatoms (e.g. Asterolampra),i.e. on or within particles large enough to be collected inthe net. Although Nitzschio bicapitato Cleve has been consideredto have a characteristic range of shapes, and has often beencited from open-ocean habitats in nano- and microplanktonicstudies, a scanning electron microscopic study revealed severalspecies in our material. Examples of larger and smaller speciesare given, and Nitzschia ikeanae G.Fryx. & H.Lee, sp. nov.,is described here. Ecologically, these observations indicatethat although some are grazed and have been seen aggregatedin fecal pellets, these small pennate diatoms can affix to andgrow on substrates, termed ‘pseudo-benthic habitats’in the open ocean, resulting in unexpected concentrations thathave implications for their life histories and sexual cycles,selection or avoidance by grazers, sinking on substrates, androles in the recycling of nutrients in near-surface waters.     

On the reorganization of fitness during evolutionary transitions in individuality

The basic problem in an evolutionary transition is to understandhow a group of individuals becomes a new kind of individual,possessing the property of heritable variation in fitness atthe new level of organization. During an evolutionary transition,for example, from single cells to multicellular organisms, thenew higher-level evolutionary unit (multicellular organism)gains its emergent properties by virtue of the interactionsamong lower-level units (cells). We see the formation of cooperativeinteractions among lower-level units as a necessary step inevolutionary transitions; only cooperation transfers fitnessfrom lower levels (costs to group members) to higher levels(benefits to the group). As cooperation creates new levels offitness, it creates the opportunity for conflict between levelsas deleterious mutants arise and spread within the group. Fundamentalto the emergence of a new higher-level unit is the mediationof conflict among lower-level units in favor of the higher-levelunit. The acquisition of heritable variation in fitness at thenew level, via conflict mediation, requires the reorganizationof the basic components of fitness (survival and reproduction)and life-properties (such as immortality and totipotency) aswell as the co-option of lower-level processes for new functionsat the higher level. The way in which the conflicts associatedwith the transition in individuality have been mediated, andfitness and general life-traits have been re-organized, caninfluence the potential for further evolution (i.e., evolvability)of the newly emerged evolutionary individual. We use the volvocaleangreen algal group as a model-system to understand evolutionarytransitions in individuality and to apply and test the theoreticalprinciples presented above. Lastly, we discuss how the differentnotions of individuality stem from the basic properties of fitnessin a multilevel selection context.     

Body size clines in sceloporus lizards: proximate mechanisms and demographic constraints

Although most species of animals examined to date exhibit Bergmann'sclines in body size, squamates tend to exhibit opposing patterns.Squamates might exhibit reversed Bergmann's clines because theytend to behaviorally regulate their body temperature effectively;the outcome of this thermoregulation is that warmer environmentsenable longer daily and annual durations of activity than coolerenvironments. Lizards of the genus Sceloporus provide an opportunityto understand the factors that give rise to contrasting thermalclines in body size because S. undulatus exhibits a standardBergmann's cline whereas S. graciosus exhibits a reverse Bergmann'scline. Interestingly, rapid growth by individuals of both speciesinvolves adjustments of physiological processes that enablemore efficient use of food. Patterns of adult body size arelikely the evolutionary consequence of variation in juvenilesurvivorship among populations. In S. undulatus, delayed maturationat a relatively large body size is exhibited in cooler environmentswhere juveniles experience higher survivorship, resulting ina Bergmann's cline. In S. graciosus, high juvenile survivorshipis not consistently found in cooler environments, resultingin no cline or a reversed Bergmann's cline, i.e., geographicpatterns in body size aren't necessarily produced by naturalselection. Thus, discerning the mechanistic links between thethermal physiology of an organism and environment-specific ratesof mortality will be critical to understanding the evolutionof body size in relation to environmental temperature.     

Control and Consequences of Alternative Developmental Modes in a Poecilogonous Polychaete

SYNOPSIS. The poecilogonous polychaete Streblospio benedicti(Webster) exhibits both planktotrophic and lecithotrophic modesof larval development. The alternative trophic modes are associatedwith differences in age and size at maturation, offspring number,size and energetic investment, larval planktonic period, morphologyand survivorship. This paper reviews a decade of research intothe control and consequences of the traits associated with planktotrophyand lecithotrophy in S. benedicti. The dominant control on reproductiveand developmental characters is genetic. Significant additivegenetic variance has been detected for egg diameter, fecundity,larval planktonic period and aspects of larval morphology. However,environmental factors such as temperature, food quality andphotoperiod, and intrinsic factors such as maternal age, exertconsiderable influence on non-trophic developmental traits (e.g.,offspring number, size and energy content). Demographic consequencesof development mode are reviewed for field and laboratory demesof S. benedicti dominated by individuals exhibiting either planktotrophyor lecithotrophy. Similar population size structure, fluctuationsin abundance, P: B ratios, and estimated population growth ratesare achieved through trade-offs between survivorship and fecundity. Development mode may best be viewed as a complex set of traitsthat are intimately linked developmentally and evolutionarilyto other aspects of an organism's life history. Greater insightinto the control and consequences of development mode shouldresult from further investigation of these linkages