Location of the ureteral openings in the cloacas of tinamous, some ratite birds, and crocodilians: a primitive character

Cloacas of 67 avian species, of both sexes, from various habitats and differing dietary habits, were examined macro- and microscopically to investigate possible variation in the location of the ureteral openings. Differing from most birds studied, in adult male Rhea americana and several tinamous species the ureters were found to open into the coprodeum. In these species the urodeum receives only the vas deferens or oviduct. Similarly, in crocodiles Caiman crocodilus yacare, but not in lizards Tropidurus montanus and snakes Crotalus durissus terrificus, the ureters empty into the coprodeum. This similarity between ancient birds (ratites and tinamous) and crocodiles may indicate a primitive character linking reptiles and birds. This unusual position of the ureteral orifice can represent an adaptation to facilitate urine collection into the coprodeum and large intestine. Another possibility is that this variation in ureter position is a male reproductive strategy to avoid the mixture of urine and semen in the cloaca. There were no evident correlations between the location of the ureteral openings and the birds' habitat, diet, or histology of the coprodeal mucosa. The occurrence of a phallus in eight species of birds was detected, as well as a peculiar vascularization related to the coprodeal epithelium of anseriformes. Together, these data add to the scarce information about the morphophysiology of the avian cloaca, and also contribute to clarify avian phylogenetic linkages.     

Does the ostrich (Struthio camelus) coprodeum have the electrophysiological properties and microstructure of other birds?

The ostrich is unique among birds in having complete separation of urine and faeces. The coprodeal epithelium is thus during dehydration exposed to a fluid 500 mOsm hyperosmotic to plasma. We have investigated whether the coprodeum is adapted like a mammalian bladder. The coprodeal epithelium was studied by electrophysiology in the Ussing chamber, and the anatomy by light microscopy and scanning electron microscopy. ELECTROPHYSIOLOGY: The short-circuit current (SCC) and open circuit electrical potential difference were recorded. The change induced by 0.1 mmol mucosal amiloride was recorded. An average basal SCC of 162+/-29 microA/cm(2) was observed, and a resistance of 297+/-34 Omega cm(2) calculated. These values are as observed in other avian coprodea. The resistance is much lower than in mammalian bladders (10000 Omega cm(2)). The amiloride-sensitive SCC, equal to net sodium absorption, was approximately 5 micromol/cm(2)h as observed in other avian species. ANATOMY: The mucosal membrane is composed of broad irregular folds with very short intestinal glands containing an unusually high proportion of goblet cells. CONCLUSION: The ostrich coprodeum is not adapted like a mammalian bladder. The abundance of goblet cells results in a copious secretion of mucus that establishes a thick unstirred layer giving effective osmotic protection.     

Ultrastructure and electrolyte transport of the epithelium of coprodeum,colon and the proctodeal diverticulum of Rhea americana

The structure and function of the lower intestinal tract of Rhea americana were characterized to evaluate the evolutionary relationship to other struthioniform and avian species. In 5 rheas the gross anatomy and the light and transmission electron microscopy were studied in parallel to in vitro electrophysiological measurements of ion transport. The mucosa in the colon was amplified with villi, often branched, and in the coprodeum with folds. In both tissues the epithelium was a monolayer composed of columnar absorptive cells, goblet cells and mitochondria-rich cells. Colon and coprodeum appeared to produce large amounts of mucus. The proctodeal diverticulum was rich in lymphoid tissue arranged into lobuli bursales, and it was concluded that this structure is a modified bursa of Fabricius. The sparse interlobular epithelium of the diverticulum resembled that of colon and coprodeum. Baseline short circuit currents (I_SC) averaged 114.5 ±13.8 µA/cm² in colon, 193.1 ± 30.3 µA/cm² in coprodeum and 60.4 ± 9.1 µA/cm² in the diverticulum. Amiloride sensitive Na⁺-transport amounted to 31, 88 and 38% of the baseline I_SC in these three tissues, respectively. In all tissues, there was also a modest, theophylline activated chloride secretion response, and ouabain, the Na⁺/K⁺-ATPase inhibitor, abolished most of the I_SC. The transepithelial resistance (TER) of the diverticulum was much higher than the other tissues. Upon dissection, urate from ureteral urine was observed in the lower third of the colon and to a lesser extent in the proctodeal diverticulum, indicating retrograde peristalsis of the urine. Thus, unlike the ostrich, there is no sphincter separating colon and coprodeum. On the other hand, a thick mucus layer was seen overlying the mucosa in both colon and coprodeum, as in the ostrich. This may help to prevent osmotic water loss, despite the presence of hyperosmotic urine (up to 800 mOsm) in the lower intestine. Both morphological and electrophysiological data from the rhea support the hypothesis that the rhea lower intestine contributes to post-renal modification of ureteral urine and to the regulation of osmotic balance, as also seen in domestic fowl and other avian species. The proctodeal diverticulum functions mainly as an immune organ, with only limited transport capability.     

Differences in renal-cloacal function between Crocodylus porosus and Alligator mississippiensis have implications for crocodilian evolution

Major electrolytes and nitrogenous excretory products were analysed in the blood plasma, ureteral urine and cloacal urine of juvenile Alligator mississippiensis and Crocodylus porosus in fresh and hypoosmotic salt water (206 mosmol · l⁻¹). Both species coped well with saline water, showing little (Alligator) or no (Crocodylus) change in plasma composition. Comparisons of renal-cloacal function point to major differences in their osmoregulatory physiology. The cloaca of C. porosus is a very active osmoregulatory organ in salt and fresh water, contributing to water conservation and NaCl excretion through the lingual salt glands. In contrast, the cloaca of Alligator has little impact on the composition of excreted urine. It seems likely that A.␣mississippiensis is largely constrained to a renal response to osmotic and ionic stress while C. porosus is able to call on a more complex mix of renal response, post-renal modification of urine in the cloaca, and excretion of excess NaCl through the salt glands. The results support the idea that there are deep-seated differences in the osmoregulatory physiology of alligatorids and crocodylids (Eusuchia), an understanding of which should provide valuable insights into their evolution and zoogeography. Accepted: 7 September 1996     

An analogue computer simulation of cloacal resorption of salt and water from ureteral urine in birds

The transmural flow of NaCl and water occurring during the retrograde flow of ureteral urine into the coprodeum and large intestine of birds has been simulated by analogue computation. The purpose was to estimate whether a fraction of the urine (water) which in the dehydrated state is hyperosmotic to plasma can, in spite of this, be absorbed from the narrow space between the epithelium and the central faeces core. The values of urine flow, urine osmolality, osmotic permeability, net NaCl absorption rate, and solute-linked water flow determined by in vivo perfusion studies in the domestic fowl were used in the calculation. The cloacal sojourn of ureteral urine was found to result in a net water gain but at the expense of a hyperosmotic NaCl absorption. The model was further used to evaluate the quantitative influence of the system's parameters upon the fractional water absorption. This was found very sensitive to the urine osmolality, moderately sensitive to the urine flow and NaCl absorption rate and almost unaffected by the osmotic permeability of the coprodeum and large intestine within a reasonable physiological range. The change of the epithelial transport parameters from the normally hydrated to the dehydrated state resulted in a marked increase in water absorption.     

Sodium excretion rates and renal responses to acute salt loading in the European starling

Summary Renal clearance studies were performed in European starlings (Sturnus vulgaris) in order to determine the extent of ureteral sodium excretion under control conditions and during an acute, hyperosmotic salt stress. These experiments also estimated the contribution of the lower intestine (colon and cloaca) to postrenal solute reabsorption by making both cloacal and ureteral urine collections in the same birds. A comparison of ureteral vs cloacal excretion rates found significantly higher sodium (9.09±1.30 vs 1.03±0.38 Eq·kg^–1·min^–1) and chloride (4.15±0.56 vs 1.00±0.38 Eq·kg^–1·min^–1) excretion rates during the ureteral collections. Fractional excretion of sodium was also significantly higher during ureteral collections, but this value did not exceed 1% of the filtered sodium load during either collection series. Urine flow rate was significantly higher during cloacal collections, suggesting osmotic back-flux of water across the cloacal wall. Infusion of a 1M NaCl solution resulted in rapid increases in glomerular filtration rate (GFR), urine flow rate, and urine osmolality. Fractional sodium and water reabsorption decreased by 11% and 4%, respectively. Glomerular counts and size distribution profiles, measured by in vivo alcian blue labelling, provided no evidence for a reduction in the number of filtering glomeruli during hyperosmotic saline loading. We conclude that renal sodium excretion rates for the starling are similar to those seen in other avian species and in mammals. These studies also provide direct evidence for postrenal modification of urine in this species, even under conditions of continuous flow. Acute hyperosmotic salt stress can, under some conditions, cause increased rather than decreased GFR, indicating multiple regulatory pathways. Finally, there was no evidence in these studies for glomerular shutdown in response to salt loading.     

6-Keto-prostaglandin F1 alpha is not added to urine during passage through the rat urinary bladder in vivo

Studies were conducted to determine whether prostaglandins are added to the urine during its passage through the rat urinary bladder in vivo. Control rats and rats with chronic streptozotocin-induced diabetes were anesthetized with Inactin, 100 mg/kg i.p., and urine was collected simultaneously from both kidneys. Urine from the left kidney was collected directly from the renal pelvis via a ureteral cannula, while urine from the right kidney was collected via a cannula in the urinary bladder. Prostaglandins in the urine were measured by radioimmunoassay. No difference in urinary concentration or rate of excretion of 6-keto-PGF1 alpha or PGE2 was seen between ureteral urine and bladder urine from either normal or diabetic rats. The results of this study indicate that in vivo there is no intralumenal addition of either 6-keto-PGF1 alpha or PGE2 to the urine by the ureteral bladder of rats.     

Osmoregulatory and metabolic costs of salt excretion in the Rufous-collared sparrow Zonotrichia capensis

Recent experiments on shorebirds have demonstrated that maintaining an active osmoregulatory machinery is energetically expensive. This may, in part, explain diet and habitat selection in birds with salt glands. However little is known about the osmoregulatory costs in birds lacking functional salt glands. In these birds, osmotic work is done almost exclusively by the kidneys. We investigated the osmoregulatory cost in a bird species lacking functional salt glands, the passerine Zonotrichia capensis. After 20 days of acclimation to fresh water (FW) and salt water (200 mM NaCl, SW), SW birds tended to be heavier than FW birds. However, this difference was not statistically significant. Total basal metabolic rate was higher in SW birds as compared with FW birds. Renal and heart masses were also higher in the SW group. We also found greater medullary development and an increase in urine osmolality in the SW group. In spite of Z. capensis' ability to tolerate a moderate salt load in the laboratory, we hypothesize that increased cost of maintenance produced by salt consumption may significantly affect energy budget, dietary, and habitat choices in the field.     

The development of the bladder trigone, the center of the anti-reflux mechanism

The urinary tract is an outflow system that conducts urine from the kidneys to the bladder via the ureters that propel urine to the bladder via peristalsis. Once in the bladder, the ureteral valve, a mechanism that is not well understood, prevents backflow of urine to the kidney that can cause severe damage and induce end-stage renal disease. The upper and lower urinary tract compartments form independently, connecting at mid-gestation when the ureters move from their primary insertion site in the Wolffian ducts to the trigone, a muscular structure comprising the bladder floor just above the urethra. Precise connections between the ureters and the trigone are crucial for proper function of the ureteral valve mechanism; however, the developmental events underlying these connections and trigone formation are not well understood. According to established models, the trigone develops independently of the bladder, from the ureters, Wolffian ducts or a combination of both; however, these models have not been tested experimentally. Using the Cre-lox recombination system in lineage studies in mice, we find, unexpectedly, that the trigone is formed mostly from bladder smooth muscle with a more minor contribution from the ureter, and that trigone formation depends at least in part on intercalation of ureteral and bladder muscle. These studies suggest that urinary tract development occurs differently than previously thought, providing new insights into the mechanisms underlying normal and abnormal development.     

Regulation of salt gland, gut and kidney interactions

Marine birds can drink seawater because their cephalic 'salt' glands secrete a sodium chloride (NaCl) solution more concentrated than seawater. Salt gland secretion generates osmotically free water that sustains their other physiological processes. Acclimation to saline induces interstitial water and Na move into cells. When the bird drinks seawater, Na enters the plasma from the gut and plasma osmolality (Osm(pl)) increases. This induces water to move out cells expanding the extracellular fluid volume (ECFV). Both increases in Osm(pl) and ECFV stimulate salt gland secretion. The augmented intracellular fluid content should allow more rapid expansion of ECFV in response to elevated Osm(pl) and facilitate activation of salt gland secretion. To fully utilize the potential of the salt glands, intestinally absorbed NaCl must be reabsorbed by the kidneys. Thus, Na uptake at gut and renal levels may constrain extrarenal NaCl secretion. High NaCl intake elevates plasma aldosterone concentration of Pekin ducks and aldosterone stimulates intestinal and renal water and sodium uptake. High NaCl intake induces lengthening of the small intestine of adult Mallards, especially males. High NaCl intake has little effect on glomerular filtration rate or tubular sodium Na uptake of birds with competent salt glands. Relative to body mass, kidney mass and glomerular filtration rate (GFR) are greater in birds with salt glands than in birds that do not have them. Birds with salt glands do not change GFR, when they drink saline. Thus, their renal filtrate contains excess Na that is, in some species, almost completely renally reabsorbed and excreted in a more concentrated salt gland secretion. Na reabsorption by kidneys of other species, like mallards is less complete and their salt glands make less concentrated secretion. Such species may reflux urine into the hindgut, where additional Na may also be reabsorbed for extrarenal secretion. During exposure to saline, marine birds maintain elevated aldosterone levels despite high Na intake. Marine birds are excellent examples of physiological plasticity.     

6-keto-prostaglandin F1α is not added to urine during passage through the rat urinary bladder

Studies were conducted to determine whether prostaglandins are added to the urine during its passage through the rat urinary blader . Control rats and rats with chronic streptozotocin-induced diabetes were anesthetized with Inactin, 100 mg/kg i.p., and urine was collected simultaneously from both kidneys. Urine from the left kidney was collected directly from the renal pelvis via a ureteral cannula, while urine from the right kidney was collected via a cannula in the urinary bladder. Prostaglandins in the urine were measured by radioimmunoassay. No difference in urinary concentration or rate of excretion of 6-keto-PGF_1α or PGE₂ was seen between ureteral urine and bladder urine from either normal or diabetic rats. The results of this study indicate that there is no intralumenal addition of either 6-keto-PGF_1α or PGE₂ to the urine by the ureteral bladder of rats.     

The functional capacities of the kidneys in hamsters of the genus Phodopus

Studies have been made on total osmolarity and concentration of Na and K in the urine, as well as on the level of diuresis in three species of the hamsters (Phodopus sungorus, Ph. campbelli, Ph. roborovskii) under the conditions of presence or absence of wet food and during water and salt loading. Mean value of osmotic concentration of the urine at free access to wet food varied within 2,600-3,500 mosmol, under the condition of dry-eating--within, 5,600--6,400 mosmol, which indicates the existence of a highly effective renal mechanism of water economy in the investigated species. At the same time, the capacity to maintain constant body weight in animals fed only air-dried food was noted only in separate specimens. In experiments with water load (30 microliters/g), the volume of fluid excreted within 4 hours amounted up to 87% of the load for Ph. sungorus, 70%--for Ph. campbelli and 43% for Ph. roborovskii. Studies on sodium and potassium contents of the urine under the conditions of dry-eating and salt loading indicate that the species investigated are capable of cation concentrating, yet they do not reveal any special mechanisms accounting for the excretion of excessive salts.     

Failure of Urinary Beta-Glucuronidase Activity to Localize the Site of Urinary Tract Infection

The differentiation of renal from bladder bacteriuria is difficult on clinical grounds alone. To evaluate the correlation between site of infection and urinary beta-glucuronidase activity, 46 patients with well documented recurrent bacteriuria were studied by bilateral ureteral catheterization. Urinary beta-glucuronidase activity was also determined in 46 control subjects. In general, asymptomatic patients with renal bacteriuria, either unilateral or bilateral, had levels of enzyme activity in their urine comparable to patients with infection confined to the bladder and to normals. Only 4 of 25 patients with renal bacteriuria had significant elevations of urinary beta-glucuronidase. After localization of infection, 9 of 10 patients treated with kanamycin, a potentially nephrotoxic drug, developed significant elevations of urinary beta-glucuronidase. The results of these studies indicate that determination of beta-glucuronidase activity in urine is not useful in predicting the site of infection in patients with bacteriuria but may find a role in screening for early nephrotoxicity.     

The functional morphology of the english sparrow cecum

As birds do not have a urinary bladder, the kidneys and lower gastrointestinal tract must function in concert to maintain fluid and electrolyte homeostasis. In birds, urine is conveyed to the cloaca, and moved by reverse peristalsis into the colon and digestive ceca. Digestive ceca have been well studied for non-passerine birds and have been shown to absorb substrates and water. The ceca of passerine birds have been suggested to be non-functional because of their small size. The present study was undertaken to examine the morphology and cytochemistry of the small ceca of the English sparrow (Passer domesticus). Three-dimensional reconstruction of the ceca from serially sectioned tissue showed these organs to have a central channel with a large number of side channels. Electron micrographs indicated that all of the channels are lined by epithelial cells with a very dense microvillus brush border as well as a region densely packed with mitochondria just below the brush border. Specific staining for Na(+), K(+)-ATPase indicated the enzyme to be localized to the brush border. Quantification of Na(+), K(+)-ATPase activity showed it to be comparable to the coprodeum of domestic fowl. The data suggest that the small ceca of passerine birds may function in fluid and electrolyte homeostasis.     

Control of renal and extrarenal salt and water excretion by plasma angiotensin II in the kelp gull (Larus dominicanus)

Summary The osmoregulatory effects of intravenously (i.v.) administered angiotensin II (AII) at dose rates of 5, 15 and 45 ng · kg^–1 · min^–1 were examined in kelp gulls utilizing salt glands and/or kidneys as excretory organs.In birds given i.v. infusion of 1200 mOsmolal NaCl at 0.3 ml · min^–1 and utilizing only the salt glands to excrete the load, infusion of AII for 30 min consistently inhibited salt gland function in a dose-dependent manner.In birds given i.v. infusion of 500 mOsmolal NaCl at 0.72 ml · min^–1 and utilizing both salt glands and kidneys to excrete the load, each dose of AII given for 2 h inhibited salt gland function but stimulated the kidney, so that the overall outputs of salt and water were enhanced and showed significant (2P<0.01) positive correlations with plasma AII.In birds given i.v. infusion of 200 mOsmolal glucose at 0.5 ml · min^–1 and utilizing only the kidneys to excrete the load, low doses of AII (5 and 15 ng · kg^–1 · min^–1) caused renal salt and water retention, whereas a high dose (45 ng · kg^–1 · min^–1) stimulated salt and water output.The actions of plasma AII in kelp gulls support the concept that this hormone plays a vital role in avian osmoregulation, having effects on both salt gland and kidney function. Elevation of plasma AII consistently inhibits actively secreting salt glands, but its effects upon renal excretion depend primarily on the osmotic status as well as on the plasma AII concentration. In conditions of salt and volume loading doses of AII stimulate sodium and water excretion. With salt and volume depletion, the action of AII is bi-phasic with low doses promoting renal sodium and water retention but high circulating levels causing natriuresis and diuresis.     

Relative thickness of the renal medulla in birds

The intraspecific mean length of medullary cones in avian kidneys is analogous to medullary thickness in the mammalian kidney. Hence, relative medullary thickness (based upon kidney volume) can be calculated for birds as was done in mammals years ago. Comparative figures are given for 26 species from nine avian orders. The organizational pattern of cortex and medulla in the bird kidney is reviewed, and a simplified diagram of this relationship is presented. With some exceptions, urine concentrating ability and relative medullary thickness are directly proportional in mammals. Contrarily, no similar trend was evident in birds when current information on water economy was compared to relative medullary thickness in various species. There are a number of factors (such as the respective functional roles of reptilian and mammalian-type nephrons, interspecific variations in ion transport, etc.) which require study before the significance of relative thickness in the avian medulla can be evaluated more thoroughly.     

Kidney concentrating ability of a subterranean xeric rodent,the naked mole-rat (Heterocephalus glaber)

The naked mole-rat (Heterocephalus glaber) is a strictly subterranean mammal inhabiting the arid zones of north-east Africa. These animals have no access to free water and water balance thus might be facilitated by regulating renal water loss. The urinary concentrating ability of the naked mole-rat was determined using five dietary manipulations in which both water and salt content were altered. Control animals (n=7) received a high quality protein cereal mixed to a thin paste with water (1 g cereal: 85 g water). Water stress was induced by reducing the water content of the diet by either 50% (n=7) or 65% (n=7). Salt loading was facilitated by replacing the water with the same volume of either 0.9% salt (n=7) or 3.0% salt (n=4) solutions. Changes in body mass, food consumption and urine volume were measured daily. The effect of diet on osmolality and electrolyte concentrations of urine and plasma were determined on termination of the diet trials. Although energy intake was not reduced, naked mole-rats lost body weight with both water stress treatments. Urine volume voided per day decreased significantly with both water stress treatments (P<0.05), such that the most extreme water stress led to an 80% reduction in urine volume. Mildly salt-loaded animals gained weight, yet underwent a sodium diuresis, as indicated by a 1.3-fold increase in the daily volume of urine voided (P<0.05). Maximum urine concentration (1521±250 mmol·kg^-1) was achieved with mild water stress and was 4.6±0.9 times that of plasma. Neither further water stress nor salt loading further increased urine osmolality (P>0.05). The naked mole-rat exhibits a moderate kidney concentrating ability and cannot maintain plasma osmolality or body mass with either extreme water stress or salt loading. Although this species succesfully inhabits arid zones, survival in these areas is not facilitated by renal water conservation, but rather by their underground existence in a microhabitat where humidities are high and radiant heat loads low. In this milieu a moderate kidney concentrating ability is adequate.Abbreviations Bm body mass - ESL extreme salt load - EWS extreme water stress - MSL mild salt load - MWS mild water stress     

栽培大豆和野生大豆耐盐性及离子效应的比较

以国际上常用的耐盐大豆（Glycine max L.)品种Lee68为对照,在发芽期和苗期两个阶段,利用发芽指数、指害指数和耐盐系数等指标对一年生具盐腺野生大豆（Glycine soja L.)和部分栽培大豆（Glycine max L.)及某些野生大豆品系或品种的耐盐性进行了比较,讨论了耐盐指标的可行性。从离子效应方面比较了Na^ 和Cl^-对大豆发芽率的影响,并对具盐腺野生大豆的耐盐机理进行了初步分析。结果表明,大豆品种的耐盐性在发芽期和苗期无一致相关性。轻度等渗胁迫下,Na^ 对种子发芽率的抑制作用大于Cl^-,而重度等渗胁迫下则相反。通过减少由根系吸收的Na^ 、Cl^-向叶片的运输,维持叶片中较高含量的K^ ,减轻盐离子毒害,可能是具盐腺野生大事耐盐的主要生理机制之一。     

The ability of rufous hummingbirds Selasphorus rufus to dilute and concentrate urine

Most terrestrial animals face the challenge of having to conserve water in a desiccating environment. Not surprisingly, the ability to produce concentrated urine has been relatively well studied in birds. Nectar-feeding birds are unusual among terrestrial animals in that they often ingest and excrete prodigious water volumes to obtain adequate energy. Thus, they confront the unusual challenge of having to conserve electrolytes. The diluting abilities of birds and the renal mechanisms that may correlate with them have been relatively neglected. To elucidate diluting and concentrating abilities in nectar-feeding birds, we fed rufous hummingbirds Selasphorus rufus an electrolyte-free nectar and a nectar containing a range of NaCl concentrations. Hummingbirds had a spectacular (and possibly unique) diluting ability: when fed on electrolyte-free food they produced excreta containing less than 0.5 mM l⁻¹ each of sodium and potassium. Hummingbirds also had a poor concentrating ability, retaining sodium and chloride when their food (0.632 M l⁻¹ sucrose) contained more than 35 mM l⁻¹ of NaCl. The kidneys of hummingbirds do not appear to be suited for concentrating urine, and possibly contain structural features that give them a unique diluting ability compared with those of birds that do not feed on nectar.     

Inter- and intraspecific variation in the use of marine food resources by three Cinclodes (Furnariidae, Aves) species: carbon isotopes and osmoregulatory physiology

The avian genus Cinclodes (Furnariidae) includes species that inhabit both inland and marine shores. We compared the carbon isotopic composition and osmoregulatory capacities of field caught individuals of three Cinclodes species in Chile. Cinclodes nigrofumosus is a resident of coastal shores, whereas C. oustaleti and C. patagonicus inhabit both coastal and inland environments. The tissues of C. nigrofumosus exhibited distinctively marine delta(13)C values, whereas those of C. oustaleti and C. patagonicus were intermediate between marine and terrestrial values. The differences in carbon isotopic composition among these three species were paralleled by differences in osmoregulatory characteristics. The species that carbon isotopes revealed to be strictly marine, C. nigrofumosus, had relatively larger kidneys with a higher fraction of total renal tissue occupied by medullary cones than its congeners C. oustaleti and C. patagonicus. Cinclodes nigrofumosus individuals also produced more concentrated urine. In addition to interspecific differences in osmoregulation, we found intraspecific differences. Cinclodes nigrofumosus collected at an arid site with limited or no available fresh water exhibited larger kidneys and higher relative medullary thickness than individuals collected at a mesic site. Cinclodes nigrofumosus, like all passerines, lacks functional salt glands. This species appears to be unique among passerines in its ability to live in extreme arid coastal environments and to cope with a marine diet that imposes high osmotic loads.