Gas exchange and habitat selection in the aquatic salamanders Necturus maculosus and Cryptobranchus alleganiensis

Summary The standard metabolic rate (SMR) and critical O₂ tension (P_c) of water-breathing mudpuppies and hellbenders were determined at 20° C using open-system respirometry. Both species are metabolic O₂ regulators, although the P_c of hellbenders (90 mmHg) is much higher than that of mudpuppies (40 mmHg). The SMR of the two species in water saturated with air was similar (19.5 and 20.0 l O₂/g·h for Cryptobranchus and Necturus, respectively) and not different from that of salamanders in general. Both species were able to survive for at least 5–11 days in severely hypoxic water (9–10 mmHg) by breathing air, indicating that the lungs are functional accessory respiratory structures.We conclude that hellbenders are restricted to relatively cool and flowing waters because of their limited gas exchange capabilities, particularly with regard to their limited aerobic scope for activity and slow recovery from exercise. Necturus maculosus is much more tolerant of hypoxia, but it is not known if they can inhabit areas were hypoxia is combined with hypercarbia.     

The Amphibian Chytrid Fungus, Batrachochytrium dendrobatidis, in Fully Aquatic Salamanders from Southeastern North America

Little is known about the impact that the pathogenic amphibian chytrid fungus, Batrachochytrium dendrobatidis (Bd), has on fully aquatic salamander species of the eastern United States. As a first step in determining the impacts of Bd on these species, we aimed to determine the prevalence of Bd in wild populations of fully aquatic salamanders in the genera Amphiuma, Necturus, Pseudobranchus, and Siren. We sampled a total of 98 salamanders, representing nine species from sites in Florida, Mississippi, and Louisiana. Overall, infection prevalence was found to be 0.34, with significant differences among genera but no clear geographic pattern. We also found evidence for seasonal variation, but additional sampling throughout the year is needed to clarify this pattern. The high rate of infection discovered in this study is consistent with studies of other amphibians from the southeastern United States. Coupled with previously published data on life histories and population densities, the results presented here suggest that fully aquatic salamanders may be serving as important vectors of Bd and the interaction between these species and Bd warrants additional research.     

Lung ventilation in salamanders and the evolution of vertebrate air-breathing mechanisms

Functional analysis of lung ventilation in salamanders combined with historical analysis of respiratory pumps provides new perspectives on the evolution of breathing mechanisms in vertebrates. Lung ventilation in the aquatic salamander Necturus maculosus was examined by means of cineradiography, measurement of buccal and pleuroperitoneal cavity pressures, and electromyography of hypaxial musculature. In deoxygenated water Necturus periodically rises to the surface, opens its mouth, expands its buccal cavity to draw in fresh air, exhales air from the lungs, closes its mouth, and then compresses its buccal cavity and pumps air into the lungs. Thus Necturus produces only two buccal movements per breath: one expansion and one compression. Necturus shares the use of this two-stroke buccal pump with lungfishes, frogs and other salamanders. The ubiquitous use of this system by basal sarcopterygians is evidence that a two-stroke buccal pump is the primitive lung ventilation mechanism for sarcopterygian vertebrates. In contrast, basal actinopterygian fishes use a four-stroke buccal pump. In these fishes the buccal cavity expands to fill with expired air, compresses to expel the pulmonary air, expands to fill with fresh air, and then compresses for a second time to pump air into the lungs. Whether the sarcopterygian two-stroke buccal pump and the actinopterygian four-stroke buccal pump arose independently, whether both are derived from a single, primitive osteichthyian breathing mechanism, or whether one might be the primitive pattern and the other derived, cannot be determined. Although Necturus and lungfishes both use a two-stroke buccal pump, they differ in their expiration mechanics. Unlike a lungfish (Protopterus), Necturus exhales by contracting a portion of its hypaxial trunk musculature (the m. Iransversus abdominis) to increase pleuroperitoneal pressure. The occurrence of this same expiratory mechanism in amniotes is evidence that the use of hypaxial musculature for expiration, but not for inspiration, is a primitive tetrapod feature. From this observation we hypothesize that aspiration breathing may have evolved in two stages: initially, from pure buccal pumping to the use of trunk musculature for exhalation but not for inspiration (as in Necturus); and secondarily, to the use of trunk musculature for both exhalation and inhalation by costal aspiration (as in amniotes).     

Respiratory,cardiovascular, and hemolymph acid‐base changes in the amphibious crab,Cardisoma guanhumi,during immersion and emersion

The cardiorespiratory and hemolymph acid base status of bimodal breathing crabs, Cardisoma guanhumi, was monitored during the transition from breathing air to breathing water. Upon immersion, oxygen uptake (MO₂) decreased by half. Ventilatory frequency (fsc) increased more than 5 fold, causing a decrease in hemolymph carbon dioxide partial pressure (PCO₂). This was nearly fully compensated for by a gradual decrease in hemolymph bicarbonate concentration ([HCO₃ ^‐]) over 96 hours post‐immersion. After one to two weeks of immersion, when crabs were removed from the water, oxygen uptake initially increased, but eventually returned to the initial immersed value. Heart rate was unchanged but fsc slowed dramatically. The decreased ventilation resulted in a buildup of hemolymph PCO₂, causing a respiratory acidosis that was slowly compensated for by increased hemolymph [HCO₃ ^‐]. C. guanhumi appears to be a truly amphibious crab with respiratory and acid‐base adaptations found in both fully aquatic and fully terrestrial species.     

Cutaneous gas exchange in snakes

Summary Cutaneous aquatic gas exchange and pulmonary gas exchange have been compared in an aquatic snakeAchrochordus javanicus and the terrestrial snakeConstrictor constrictor.Gas exchange was measured by closed respirometry with the snakes in air and in water with access to air. Frequency of air breathing, tidal volumes and total lung volumes were also compared in the two species. All measurements were done at 20–22 ° C.The aquaticAchrochordus showed long periods of apnea in submerged condition interrupted by short periods of breathing activity at the surface. Average frequency of air breathing activity was 2.6 times per hour. Breathing in constrictor was more frequent but irregular with an average frequency of 143 breaths per hour.Total lung volume was 66±31 ml/kg body weight and 72.5±59 ml/kg body weight inAchrochordus andConstrictor, respectively. Tidal volumes were 41.5±4.4 ml/kg body weight and 29.5±14.8 ml/kg body weight, largest inAchrochordus. Constrictor had the highest total O₂ uptake ( ) correlating with a higher activity. Total gas exchange ratio (R _E ) was 0.69 forConstrictor and 0.77 forAchrochordus. InConstrictor air breathing accounted for 97% of the total whereas 21% of the CO₂ exchange was aquatic. Corresponding figures forAchrochordus were 92% of total by air breathing with as much as 33% of the CO₂ elimination as aquatic gas exchange.The results demonstrate that the trend among early air breathing vertebrates (fishes and amphibians) of a conservative evolution of CO₂ elimination by air breathing also extends to snakes.Significantly the cutaneous exchange component was highest in the more aquatic species.The results are discussed in relation to recent reports of a higher than alleged role of the skin of reptiles in evaporative water loss.This study was supported by grant HE 12071 from the National Institutes of Health in the U. S. A.     

Respiratory gas exchange in the airbreathing fish,Synbranchus marmoratus

Synopsis The partitioning of O₂ uptake between aquatic and aerial gas exchange and its dependence on ambient water PO₂ was studied in the facultative air breathing teleost Synbranchus marmoratus, after acclimation to well aerated water and after acute and chronic exposure to hypoxic water. O₂ uptake was also studied following acute air exposure and after prolonged entrapment in soil. Breathing rates during water and air breathing in response to reduced water PO₂ and tidal volume during air breathing were also studied. S. marmoratus satisfies its O₂ requirement by water breathing alone until water PO₂ falls below 30–50 mm Hg (switching PO₂) depending on the acclimation history. Below the switching PO₂, air breathing is adopted while active water breathing stops. The O₂ uptake varied little for all groups when the principal mode of gas exchange changed at the switching PO₂. The highest O₂ uptake prevailed when the fish employed the mode of gas exchange in operation during the acclimation period (i.e. water breathing for normoxia-acclimated, air breathing for hypoxic-acclimated).Acclimation to chronic hypoxia gave a much higher switching PO₂ 55 mm Hg) than for the other groups (about 30 mm Hg). S. marmoratus maintained its O₂ uptake when acutely exposed to air. When entrapped in soil in an aestivating state, the O₂ uptake was reduced to 25% of that in water or during acute air exposure. The overall gas exchange ratio for air breathing was very low (R_E 0.1).Branchial water pumping increased with lowering of water PO₂. The rate of air breathing was independent of water PO₂.The findings are discussed in the light of the ecophysiological conditions confronting S. marmoratus.     

Gross and fine anatomy of the respiratory vasculature of the mudskipper,Periophthalmodon schlosseri (Gobiidae: Oxudercinae)

To illustrate vascular modification accompanying transition from aquatic to amphibious life in gobies, we investigated the respiratory vasculatures of the gills and the bucco‐opercular cavities in one of the most terrestrially‐adapted mudskippers, Periophthalmodon schlosseri, using the corrosion casting technique. The vascular system of Pn. schlosseri retains the typical fish configuration with a serial connection of the gills and the systemic circuits, suggesting a lack of separation of O₂‐poor systemic venous blood and O₂‐rich effluent blood from the air‐breathing surfaces. The gills appear to play a limited role in gas exchange, as evidenced from the sparsely‐spaced short filaments and the modification of secondary lamellar vasculature into five to eight parallel channels that are larger than red blood cell size, unlike the extensive sinusoidal system seen in purely water‐breathing fishes. In contrast, the epithelia of the bucco‐opercular chamber, branchial arches, and leading edge of the filaments are extensively laden with capillaries having a short (<10 μm) diffusion distance, which strongly demonstrate the principal respiratory function of these surfaces. These capillaries form spiral coils of three to five turns as they approach the epithelial surface. The respiratory capillaries of the bucco‐opercular chamber are supplied by efferent blood from the gills and drained by the systemic venous pathway. We also compared the degree of capillarization in the bucco‐opercular epithelia of Pn. schlosseri with that of the three related intertidal‐burrowing gobies (aquatic, non‐air‐breathing Acanthogobius hasta; aquatic, facultative air‐breathing Odontamblyopus lacepedii; amphibious air‐breathing Periophthalmus modestus) through histological analysis. The comparison revealed a clear trend of wider distribution of denser capillary networks in these epithelia with increasing reliance on air breathing, consistent with the highest aerial respiratory capacity of Pn. schlosseri among the four species. J. Morphol. 2011. © 2011 Wiley‐Liss, Inc.     

CO2‐metabolism in early tetrapods revisited: inferences from osteological correlates of gills,skin and lung ventilation in the fossil record

One of the most important physiological changes during the conquest of land by vertebrates was the increasing reliance on lung breathing, with the concomitant decrease in importance of gill breathing. The main problem involved here was to cope with the excessive accumulation of CO₂ in the body and to avoid respiratory acidosis. In the past, several often mutually contradicting hypotheses of CO₂‐elimination via skin, lungs and gills in early tetrapods have been proposed, based on theoretical physiological considerations and comparison with extant air‐breathing fishes and amphibians. This study proposes a revised scenario of CO₂‐elimination in early tetrapods based on fossil evidence, that is recently identified osteological correlates of gills, skin structure and mode of lung ventilation. In stem tetrapods of the Devonian and Carboniferous, O₂‐uptake via the lungs by buccal pumping was decoupled from CO₂‐release via internal gills, and the rather gas‐impermeable skin played a minor role in gaseous exchange. The two main lineages of crown‐group tetrapods, the amphibian and amniote lineage, used different strategies of CO₂‐elimination. As in stem tetrapods, O₂‐uptake and CO₂‐release remained always largely decoupled in temnospondyls, which ventilated their lungs via buccal pumping and relied mainly on their internal gills for CO₂‐release. Temnospondyls were not able to reduce their internal gills before their skin became more gas permeable and their body size was reduced, to shift from internal gills to the skin as the major site of CO₂‐elimination, a pattern that is retained in most lissamphibians. In contrast, internal gills were lost very early in stem amniote evolution. This was associated with the evolution of the more effective aspiration pump that allowed the elimination of the bulk of CO₂ via the lungs, leading to a coupled O₂‐uptake and CO₂‐loss in stem amniotes and later in amniotes.     

Morphological variation of hypaxial musculature in salamanders (Lissamphibia: Caudata)

Despite the acknowledged importance of the locomotory and respiratory functions associated with hypaxial musculature in salamanders, variation in gross morphology of this musculature has not been documented or evaluated within a phylogenetic or ecological context. In this study, we characterize and quantify the morphological variation of lateral hypaxial muscles using phylogenetically and ecologically diverse salamander species from eight families: Ambystomatidae (Ambystoma tigrinum), Amphiumidae (Amphiuma tridactylum), Cryptobranchidae (Cryptobranchus alleganiensis), Dicamptodontidae (Dicamptodon sp.), Plethodontidae (Gyrinophilus porphyriticus), Proteidae (Necturus maculosus), Salamandridae (Pachytriton sp.), and Sirenidae (Siren lacertina). For the lateral hypaxial musculature, we document 1) the presence or absence of muscle layers, 2) the muscle fiber angles of layers at mid‐trunk, and 3) the relative dorsoventral positions and cross‐sectional areas of muscle layers. Combinations of two, three, or four layers are observed. However, all species retain at least two layers with opposing fiber angles. The number of layers and the presence or absence of layers vary within species (Necturus maculosus and Siren lacertina), within genera (e.g., Triturus), and within families. No phylogenetic pattern in the number of layers can be detected with a family‐level phylogeny. Fiber angle variation of hypaxial muscles is considerable: fiber angles of the M. obliquus externus range from 20–80°; M. obliquus internus, 14–34°; M. transversus abdominis, 58–80° (acute angles measured relative to the horizontal septum). Hypaxial musculature comprises 17–37% of total trunk cross‐sectional area. Aquatic salamanders show relatively larger total cross‐sectional hypaxial area than salamanders that are primarily terrestrial. J. Morphol. 241:153–164, 1999. © 1999 Wiley‐Liss, Inc.     

Energetics of chemolithoautotrophy in the hydrothermal system of Vulcano Island, southern Italy

The hydrothermal system at Vulcano, Aeolian Islands (Italy), is home to a wide variety of thermophilic, chemolithoautotrophic archaea and bacteria. As observed in laboratory growth studies, these organisms may use an array of terminal electron acceptors (TEAs), including O₂, , Fe(III), , elemental sulphur and CO₂; electron donors include H₂, , Fe²⁺, H₂S and CH₄. Concentrations of inorganic aqueous species and gases were measured in 10 hydrothermal fluids from seeps, wells and vents on Vulcano. These data were combined with standard Gibbs free energies () to calculate overall Gibbs free energies (ΔG_r) of 90 redox reactions that involve 16 inorganic N‐, S‐, C‐, Fe‐, H‐ and O‐bearing compounds. It is shown that oxidation reactions with O₂ as the TEA release significantly more energy (normalized per electron transferred) than most anaerobic oxidation reactions, but the energy yield is comparable or even higher for several reactions in which , or Fe(III) serves as the TEA. For example, the oxidation of CH₄ to CO₂ coupled to the reduction of Fe(III) in magnetite to Fe²⁺ releases between 94 and 123 kJ/mol e^?, depending on the site. By comparison, the aerobic oxidation of H₂ or reduced inorganic N‐, S‐, C‐ and Fe‐bearing compounds generally yields between 70 and 100 kJ/mol e^?. It is further shown that the energy yield from the reduction of elemental sulphur to H₂S is relatively low (8–19 kJ/mol e^?) despite being a very common metabolism among thermophiles. In addition, for many of the 90 reactions evaluated at each of the 10 sites, values of ΔG_r tend to cluster with differences < 20 kJ/mol e^?. However, large differences in ΔG_r (up to ～ 60 kJ/mol e^?) are observed in Fe redox reactions, due largely to considerable variations in Fe²⁺, H⁺ and H₂ concentrations. In fact, at the sites investigated, most variations in ΔG_r arise from differences in composition and not in temperature.     

Tree size frequency distributions, plant density, age and community disturbance

We show that explicit mathematical and biological relationships exist among the scaling exponents and the allometric constants (α and β, respectively) of log–log linear tree‐community size frequency distributions, plant density N_T, and minimum, maximum and average stem diameters (D_min, D_max, and , respectively). As individuals grow in size and D_max increases, N_T is predicted to decrease as reflected by a decrease in the numerical value of α and an increase in the value of β. Our derivations further show that N_T decreases as increases even if D_min or D_max remain unchanged. Because D_max and the age of the largest individuals in a community are correlated, albeit weakly, we argue that the interdependent relationships among the numerical values of α, β, N_T, and shed light on the extent to which communities have experienced recent global disturbance. These predicted relationships receive strong statistical support using two large datasets spanning a broad spectrum of tree‐dominated communities.     

Biological traits and life table of the exotic Harmonia axyridis compared with Hippodamia variegata,and Adalia bipunctata (Col., Coccinellidae)

Abstract: As part of an environmental risk assessment study of exotic natural enemies used in inundative biological control, life‐history characteristics of Harmonia axyridis (Pallas), Hippodamia variegata (Goeze) and Adalia bipunctata (L.) (Col., Coccinellidae) were quantified under laboratory conditions at 25°C on Myzus persicae (Sulzer) as prey. Comparative studies showed significant differences among pre‐adult development times: H. axyridis developed slower ( = 19.8 days) than H. variegata ( = 18.1 days) and A. bipunctata ( = 18.4 days). Differences were also evident in the duration of egg, larval and pupal stages. No measurable differences among the three species were found for fecundity, oviposition rate and adult longevity. Harmonia axyridis exhibited the longest pre‐oviposition ( = 7.4 days) and interoviposition ( = 3.6 days) periods and the shortest oviposition period ( = 13.7 days). The Bieri model was used to describe age‐specific fecundity for the three species of coccinellids. The intrinsic rate of increase (r_m), net reproductive rate (R₀) and mean generation time (T) were higher for H. variegata (r_m = 0.114, R₀ = 52.75, T = 41.88 days) than for H. axyridis (r_m = 0.089, R₀ = 26.27, T = 38.81 days) or A. bipunctata (r_m = 0.081, R₀ = 18.49, T = 40.06 days). Our findings show that the biological traits of H. axyridis do not seem to be factors that may contribute to the invasiveness of this coccinellid.     

Renal transport of various organic anions inNecturus

Summary The renal handling of several organic anions has been investigated in the salamanderNecturus maculosus. Uptake ofP-aminohippurate (PAH) by theNecturus kidney slice was inhibited by iodoacetamide (IA), ouabain, removal of sodium from the medium (Table 1) and by probenecid, N-(4-azido-2-nitrophenyl)-2-aminoethyl sulfonate (NAP-taurine), and 4-acetamido-4-isothiocyano-2,2-disulfonic stilbene (SITS) (Fig. 4), all of which also inhibit PAH uptake in the mammalian kidney. The sulfonic anions, NAP-taurine and phenol red (PSP), were not actively taken up by the slice (Table 1), and PSP had no effect on PAH uptake. NAP-taurine, on the other hand, competitively inhibited PAH uptake (Fig. 5) withK _i=2.2×10^–5 M. The maximum inhibitory effect was 50% of control uptake at NAP-taurine concentrations greater than 10^–5 M, whereas a maximum of 80% inhibition was obtained with probenecid (Fig. 4). These results support the idea that theNecturus kidney is not able to actively secrete sulfonic anions. The interaction of NAP-taurine with the PAH uptake system may be due to structural features other than the sulfonic acid moiety. There appear to be two components of PAH uptake, one which is sensitive to NAP-taurine, and one which is not.Abbreviations PAH p-aminohippurate - PSP phenolsulfophthalein (phenol red) - NAP taurine N-(4-azido-2-nitrophenyl)-2-aminoethyl sulfonate - SITS 4-acctamido-4-isothiocyano-2,2-disulfonic stilbene - S/M slice to medium ratio:IA iodoacctamide     

Modelling evolutionary processes in small populations: not as ideal as you think

Evolutionary processes are routinely modelled using ‘ideal’ Wright–Fisher populations of constant size N in which each individual has an equal expectation of reproductive success. In a hypothetical ideal population, variance in reproductive success (V_k) is binomial and effective population size (N_e) = N. However, in any actual implementation of the Wright–Fisher model (e.g., in a computer), V_k is a random variable and its realized value in any given replicate generation () only rarely equals the binomial variance. Realized effective size () thus also varies randomly in modelled ideal populations, and the consequences of this have not been adequately explored in the literature. Analytical and numerical results show that random variation in  and  can seriously distort analyses that evaluate precision or otherwise depend on the assumption that  is constant. We derive analytical expressions for Var(V_k) [4(2N – 1)(N – 1)/N³] and Var(N_e) [N(N – 1)/(2N – 1) ≈ N/2] in modelled ideal populations and show that, for a genetic metric G = f(N_e), Var(?) has two components: Var_Gene (due to variance across replicate samples of genes, given a specific ) and Var_Demo (due to variance in ). Var(?) is higher than it would be with constant N_e = N, as implicitly assumed by many standard models. We illustrate this with empirical examples based on F (standardized variance of allele frequency) and r² (a measure of linkage disequilibrium). Results demonstrate that in computer models that track multilocus genotypes, methods of replication and data analysis can strongly affect consequences of variation in . These effects are more important when sampling error is small (large numbers of individuals, loci and alleles) and with relatively small populations (frequently modelled by those interested in conservation).     

Respiration of the air breathing fishPiabucina festae

Summary Piabucina festae, a Central American stream fish, breathes air frequently, even in air saturated water, however, is not an obligate air breather. Without access to air, it can maintain routine by aquatic respiration down to aP _wO₂ of about 70 Torr which is its critical O₂ tension (P _cO₂, Fig. 5). Aerial respiration averages 10% of total in air saturated water and 70% in hypoxic water (Fig. 4). At lowP _wO₂ air breathing is more frequent (Fig. 1), and more O₂ is utilized from each air breath (Table 3), and tidal volume may increase (Fig. 7). Vascularized respiratory compartments or cells (Fig. 6), located in the second chamber of the physostomus gas bladder, function for aerial respiration. In ventilation air is gulped and forced through a large pneumatic duct into the gas bladder, excess gas is then released through opercula. Inspiration always precedes expiration and tidal volume is small, keeping gas bladderP _{O 2} low (Table 4). Major differences in the air breathing physiology ofP. festae and other species are its higherP _cO₂, a low aerial in normoxic water, even though air gulps are frequent, and its pattern of inhalation prior to expiration. The interrelationship and optimization of the three gas bladder functions (buoyancy, sound reception, and air breathing) inP. festae is discussed. Aerial respiration may have evolved secondarily to the gas bladder's function in buoyancy control.     

Estimating parameters in a land-surface model by applying nonlinear inversion to eddy covariance flux measurements from eight FLUXNET sites

Flux measurements from eight global FLUXNET sites were used to estimate parameters in a process‐based, land‐surface model (CSIRO Biosphere Model (CBM), using nonlinear parameter estimation techniques. The parameters examined were the maximum photosynthetic carboxylation rate () the potential photosynthetic electron transport rate (j_{max, 25}) of the leaf at the top of the canopy, and basal soil respiration (r_{s, 25}), all at a reference temperature of 25°C. Eddy covariance measurements used in the analysis were from four evergreen forests, three deciduous forests and an oak‐grass savanna. Optimal estimates of model parameters were obtained by minimizing the weighted differences between the observed and predicted flux densities of latent heat, sensible heat and net ecosystem CO₂ exchange for each year. Values of maximum carboxylation rates obtained from the flux measurements were in good agreement with independent estimates from leaf gas exchange measurements at all evergreen forest sites. A seasonally varying and j_{max, 25} in CBM yielded better predictions of net ecosystem CO₂ exchange than a constant and j_{max, 25} for all three deciduous forests and one savanna site. Differences in the seasonal variation of and j_{max, 25} among the three deciduous forests are related to leaf phenology. At the tree‐grass savanna site, seasonal variation of and j_{max, 25} was affected by interactions between soil water and temperature, resulting in and j_{max, 25} reaching maximal values before the onset of summer drought at canopy scale. Optimizing the photosynthetic parameters in the model allowed CBM to predict quite well the fluxes of water vapor and CO₂ but sensible heat fluxes were systematically underestimated by up to 75 W m⁻².     

Evolutionary insights into the North American Necturus beyeri complex (Amphibia: Caudata) based on molecular genetic and morphological analyses

Necturus beyeri (Caudata: Proteidae), as conceived by some, contains paedomorphic salamanders distributed from the Ochlockonee drainage of Florida to the Angelina drainage of Texas. Because these salamanders differ in color pattern and karyotype across their geographic range, we performed a phylogeographic analysis that included representatives from all major drainages as well as of all congeners. The mitochondrially encoded ND2 gene was used to infer phylogenetic relationships using Bayesian inference. Morphometrics of head shape were analyzed and included as an independent data set. Our work suggests that Necturus comprises 11 lineages. A basal split within the genus separates an ancestor of two Atlantic Coastal Plain species (Necturus lewisi and Necturus punctatus) from the ancestor of nine distinct Gulf Coastal Plain lineages. One lineage is consistent with Necturus alabamensis, a species currently recognized in the Black Warrior drainage of Alabama. Two lineages comprise Necturus maculosus, as historically recognized, and six lineages comprise N. beyeri, as recognized by some, each of which occupies a unique drainage. Both of these species are demonstrated to be paraphyletic. Head morphometrics show the same patterns as the mtDNA. Overall, lineages within Necturus exhibit an east‐to‐west progression of appearance on the phylogenetic trees. This pattern corroborates biogeographic hypotheses based on previous karyological work. Within N. beyeri, this progression separates a pattern class of two eastern lineages lacking bold spotting and possessing relatively small mean body lengths from a pattern class of four western lineages possessing bold spotting and larger mean body sizes. Thus, the two eastern lineages of N. beyeri are similar in color pattern and body size to N. punctatus either through retention of the ancestral color pattern and size for the genus or through convergent selection in eastern streams of the Gulf Coastal Plain.     

INTERACTIONS BETWEEN NITRATE UPTAKE AND NITROGEN FIXATION IN CONTINUOUS CULTURES OF THE MARINE DIAZOTROPH TRICHODESMIUM (CYANOBACTERIA)1

Diazotrophic cyanobacteria can take up combined nitrogen (nitrate, ammonium, amino acids, dissolved organic nitrogen) from solution, but the interaction between N₂ fixation and uptake of combined nitrogen is not well understood. We studied the effects of combined nitrogen ) additions on N₂ fixation rates in the cyanobacterium Trichodesmium erythraeum (IMS‐101) maintained in continuous culture in an N‐free medium (YBCII) and a 12:12‐h light:dark cycle. We measured acetylene reduction rates, nutrient concentrations, and biomass throughout the 12 h of illumination after the addition of nitrate (0.5–20 μM) at the start of the light period. Compared with unamended controls, Trichodesmium showed strong inhibition of acetylene reduction (up to 70%) in the presence of , with apparent saturation of the inhibition effect at an initial concentration of approximately 10 μM. The inhibition of acetylene reduction persisted through much of the light period as concentration in the culture vessel decreased. Recovery of N₂ fixation was observed late in the light period in cultures amended with low concentrations of (<5 μM) when ambient concentrations had decreased to 0.3–0.4 μM in the culture vessel. Nitrate uptake accounted for as much as 86% of total N uptake and, at the higher treatment concentrations, more than made up for the observed decrease in N₂ fixation rates. We conclude that Trichodesmium can obtain significant quantities of N through uptake of nitrate and does so in preference to N₂ fixation when sufficient is available.     

Apparent Survival Estimates for Five Species of Tropical Birds in an Endangered Forest Habitat in Western Ecuador1

Estimates of annual survival are essential for addressing topics in evolutionary and conservation ecology. However, most demographic studies of land birds are based on north temperate species, and few robust estimates of survival based on mark–recapture statistics are available for continental South American birds. We used time‐since‐marking models to estimate apparent survival of adult birds from 7 yr of mist netting data in the Colonche Hills. This site is one of few remaining large tracts of premontane forest in southwestern Ecuador, and an area of high priority for avian conservation. Species with sufficient data for analysis included three hummingbirds (Adelomyia melanogenys, Heliodoxa jacula, Phaethornis baroni), a cotinga (Schiffornis turdinus), and a wren (Henicorhina leucophrys). Our parameter estimates had reduced precision because the number of recaptures was small. Probability of recapture was low in three species , and moderate in two others . Adelomyia and Phaethornis had moderate apparent survival (; probability that a bird neither died nor emigrated from our survey area in a given year). Adults of Adelomyia moved seasonally, and it is possible that permanent emigration from our survey area contributed to low estimates of apparent survival. The other three species had relatively high estimates of adult apparent survival ranging from in Heliodoxa and Schiffornis to a high of in Henicorhina.