Effect of type of otolith and preparation technique on age estimation of larval and juvenile spot (Leiostomus xanthurus)

Otoliths of larval and juvenile fish provide a record of age, size, growth, and development (Campana and Neilson, 1985; Thorrold and Hare, 2002). However, determining the time of first increment formation in otoliths (Campana, 2001) and assessing the accuracy (deviation from real age) and precision (repeatability of increment counts from the same otolith) of increment counts are prerequisites for using otoliths to study the life history of fish (Campana and Moksness, 1991). For most fish species, first increment deposition occurs either at hatching, a day after hatching, or after first feeding and yolksac absorption (Jones, 1986; Thorrold and Hare, 2002). Increment deposition before hatching also occurs (Barkmann and Beck, 1976; Radtke and Dean, 1982). If first increment deposition does not occur at hatching, the standard procedure is to add a predetermined number to increment counts to estimate fish age (Campana and Neilson, 1985).

Age validation of quillback rockfish (Sebastes maliger) using bomb radiocarbon

Rockfishes (Sebastes spp.) support one of the most economically important f isheries of the Pacific Northwest and it is essential for sustainable management that age estimation procedures be validated for these species. Atmospheric testing of thermonuclear devices during the 1950s and 1960s created a global radiocarbon (14C) signal in the ocean environment that scientists have identified as a useful tracer and chronological marker in natural systems. In this study, we first demonstrated that fewer samples are necessary for age validation using the bomb-generated 14C signal by emphasizing the utility of the time-specific marker created by the initial rise of bomb-14C. Second, the bomb-generated 14C signal retained in fish otoliths was used to validate the age and age estimation method of the quillback rockfish (Sebastes maliger) in the waters of southeast Alaska. Radiocarbon values from the first year’s growth of quillback rockfish otoliths were plotted against estimated birth year to produce a 14C time series spanning 1950 to 1985. The initial rise in bomb-14C from prebomb levels (~ –90‰) occurred in 1959 [±1 year] and 14C levels rose relatively rapidly to peak Δ14C values in 1967 (+105.4‰) and subsequently declined through the end of the time series in 1985 (+15.4‰). The agreement between the year of initial rise of 14C levels from the quillback rockfish time series and the chronology determined for the waters of southeast Alaska from yelloweye rockfish (S. ruberrimus) otoliths validated the aging method for the quillback rockfish. The concordance of the entire quillback rockfish 14C time series with the yelloweye rockfish time series demonstrated the effectiveness of this age validation technique, confirmed the longevity of the quillback rockfish up to a minimum of 43 years, and strongly confirms higher age estimates of up

Age, growth and mortality of the toadfish, Halobatrachus didactylus (Schneider, 1801) (Pisces: Batrachoididae), in the Bay of Cádiz (southwestern Spain)

Age, growth and mortality of the toadfish, Halobatrachus didactylus, were determined by examination of the whole sagittal otoliths of fish sampled in the Bay of Cádiz (southwestern Spain) from March 1999 to March 2000. A total of 844 specimens (425 males, 416 females, and 3 of indeterminate sex), ranging from 95 to 470 mm in total length were examined. Eighty-nine percent of the otoliths could be read allowing an age estimation. The opaque zone was formed between April and May coincident with the maximum reproductive peak, while the translucent zone formed mainly in summer-fall (June to December). Maximum ages for males and females were 12 and 10 years, respectively. The samples were dominated by 2- to 6-year-old specimens. Males matured at an age of approximately 2 years and females at 3 years. Fish total length and otolith radius were closely related. The von Bertalanffy growth curve was used to describe growth. The parameters were derived from back-calculated length-at-age. Significant differences in the growth parameters were found between sexes. Although the growth analysis revealed that this species is slow-growing, males reached larger sizes than females. Females appeared to experience higher natural mortality rates than males.

Age and growth of the estuarine dolphin (Sotalia guianensis) (Cetacea, Delphinidae) on the Paraná Coast, southern Brazil

Teeth of 71 estuarine dolphins (Sotalia guianensis) incidentally caught on the coast of Paraná State, southern Brazil, were used to estimate age. The oldest male and female dolphins were 29 and 30 years, respectively. The mean distance from the neonatal line to the end of the first growth layer group (GLG) was 622.4 ±19.1 μm (n=48). One or two accessory layers were observed between the neonatal line and the end of the first GLG. One of the accessory layers, which was not always present, was located at a mean of 248.9 ±32.6 μm (n=25) from the neonatal line, and its interpretation remains uncertain.The other layer, located at a mean of 419.6 ±44.6 μm (n=54) from the neonatal line, was always present and was first observed between 6.7 and 10.3 months of age. This accessory layer could be a record of weaning in this dolphin. Although no differences in age estimates were observed between teeth sectioned in the anterior-posterior and buccal-lingual planes, we recommend sectioning the teeth in the buccal-lingual plane in order to obtain on-center sections more easily. We also recommend not using teeth from the most anterior part of the mandibles for age estimation. The number of GLGs counted in those teeth was 50% less than the number of GLGs counted in the teeth from the median part of the mandible of the same animal. Although no significant difference (P>0.05) was found between the total lengths of adult male and female estuarine dolphins, we observed that males exhibited a second growth spurt around five years of age. This growth spurt would require that separate growth curves be calculated for the sexes. The asymptotic length (TL∞), k, and t0 obtained by the von Bertalanffy growth model were 177.3 cm, 0.66, and –1.23, respectively, for females and 159.6 cm, 2.02, and –0.38, respectively, for males up to five years, and 186.4 cm, 0.53 and –1.40, respectively, for males older than five years. The total weight (TW)/total length (TL) equations obtained for male and female estuarine dolphins were TW = 3.156 × 10−6 × TL 3.2836 (r=0.96), and TW = 8.974 × 10−5 × TL 2.6182 (r=0.95), respectively.

Age and growth of Hawaiian seaturtles (Chelonia mydas): an analysis based on skeletochronology

Skeletochronological data on growth changes in humerus diameter were used to estimate the age of Hawaiian green seaturtles ranging from 28.7 to 96.0 cm straight carapace length. Two age estimation methods, correction factor and spline integration, were compared, giving age estimates ranging from 4.1 to 34.6 and from 3.3 to 49.4 yr, respectively, for the sample data. Mean growth rates of Hawaiian green seaturtles are 4–5 cm/yr in early juveniles, decline to a relatively constant rate of about 2 cm/yr by age 10 yr, then decline again to less than 1 cm/yr as turtles near age 30 yr. On average, age estimates from the two techniques differed by just a few years for juvenile turtles, but by wider margins for mature turtles. The spline-integration method models the curvilinear relationship between humerus diameter and the width of periosteal growth increments within the humerus, and offers several advantages over the correction-factor approach.

Bomb radiocarbon and tag-recapture dating of sandbar shark (Carcharhinus plumbeus)

The sandbar shark (Carcharhinus plumbeus) was the cornerstone species of western North Atlantic and Gulf of Mexico large coastal shark fisheries until 2008 when they were allocated to a research-only fishery. Despite decades of fishing on this species, important life history parameters, such as age and growth, have not been well known. Some validated age and growth information exists for sandbar shark, but more comprehensive life history information is needed. The complementary application of bomb radiocarbon and tag-recapture dating was used in this study to determine valid age-estimation criteria and longevity estimates for this species. These two methods indicated that current age interpretations based on counts of growth bands in vertebrae are accurate to 10 or 12 years. Beyond these years, we could not determine with certainty when such an underestimation of age begins; however, bomb radiocarbon and tag-recapture data indicated that large adult sharks were considerably older than the estimates derived from counts of growth bands. Three adult sandbar sharks were 20 to 26 years old based on bomb radiocarbon results and were a 5- to 11-year increase over the previous age estimates for these sharks. In support of these findings, the tag-recapture data provided results that were consistent with bomb radiocarbon dating and further supported a longevity that exceeds 30 years for this species.

Validation and interpretation of annual skeletal marks in loggerhead (Caretta caretta) and Kemp’s ridley (Lepidochelys kempii) sea turtles

Numerous studies have applied skeletochronology to sea turtle species. Because many of the studies have lacked validation, the application of this technique to sea turtle age estimation has been called into question. To address this concern, we obtained humeri from 13 known-age Kemp’s ridley (Lepidochelys kempii) and two loggerhead (Caretta caretta) sea turtles for the purposes of examining the growth marks and comparing growth mark counts to actual age. We found evidence for annual deposition of growth marks in both these species. Corroborative results were found in Kemp’s ridley sea turtles from a comparison of death date and amount of bone growth following the completion of the last growth mark (n=76). Formation of the lines of arrested growth in Kemp’s ridley sea turtles consistently occurred in the spring for animals that strand dead along the mid- and south U.S. Atlantic coast. For both Kemp’s ridley and loggerhead sea turtles, we also found a proportional allometry between bone growth (humerus dimensions) and somatic growth (straight carapace length), indicating that size-at-age and growth rates can be estimated from dimensions of early growth marks. These results validate skeletochronology as a method for estimating age in Kemp’s ridley and loggerhead sea turtles from the southeast United States.

Estimation of the abundance of yellowfin tuna, Thunnus albacares, by age groups and regions within the Eastern Pacific Ocean

ENGLISH: Monthly estimates of the abundance of yellowfin tuna by age groups and regions within the eastern Pacific Ocean during 1970-1988 are made, using purse-seine catch rates, length-frequency samples, and results from cohort analysis. The numbers of individuals caught of each age group in each logged purse-seine set are estimated, using the tonnage from that set and length-frequency distribution from the "nearest" length-frequency sample(s). Nearest refers to the closest length frequency sample(s) to the purse-seine set in time, distance, and set type (dolphin associated, floating object associated, skipjack associated, none of these, and some combinations). Catch rates are initially calculated as the estimated number of individuals of the age group caught per hour of searching. Then, to remove the effects of set type and vessel speed, they are standardized, using separate weiznted generalized linear models for each age group. The standardized catch rates at the center of each 2.5 0 quadrangle-month are estimated, using locally-weighted least-squares regressions on latitude, longitude and date, and then combined into larger regions. Catch rates within these regions are converted to numbers of yellowfin, using the mean age composition from cohort analysis. The variances of the abundance estimates within regions are large for 0-, 1-, and 5-year-olds, but small for 1.5- to 4-year-olds, except during periods of low fishing activity. Mean annual catch rate estimates for the entire eastern Pacific Ocean are significantly positively correlated with mean abundance estimates from cohort analysis for age groups ranging from 1.5 to 4 years old. Catch-rate indices of abundance by age are expected to be useful in conjunction with data on reproductive biology to estimate total egg production within regions. The estimates may also be useful in understanding geographic and temporal variations in age-specific availability to purse seiners, as well as age-specific movements. SPANISH: Se calculan estimaciones mensuales de la abundancia del atún aleta amarilla por grupos de edad y regiones en el Océano Pacífico oriental durante 1970-1988, usando tasas de captura cerquera, muestras de frecuencia de talla, y los resultados del análisis de cohortes. Se estima el número de individuos capturados de cada grupo de edad en cada lance cerquero registrado, usando el tonelaje del lance en cuestión y la distribución de frecuencia de talla de la(s) muestra(s) de frecuencia de talla "más cercana/s)," "Más cercana" significa la(s) muestra(s) de frecuencia de talla más parecida(s) al lance cerquero en cuanto a fecha, distancia, y tipo de lance (asociado con delfines, con objeto flotante, con barrilete, con ninguno de éstos, y algunas combinaciones). Se calculan inicialmente las tasas de captura como el número estimado de individuos del grupo de edad capturado por hora de búsqueda. A continuación, para eliminar los efectos del tipo de lance y la velocidad del barco, se estandardizan dichas tasas, usando un modelo lineal generalizado ponderado, para cada grupo por separado. Se estima la tasa de captura estandardizada al centro de cada cuadrángulo de 2.5°-mes, usando regresiones de mínimos cuadrados ponderados localmente por latitud, longitud, y fecha, y entonces combinándolas en regiones mayores. Se convierten las tasas de captura dentro de estas regiones en números de aletas amarillas individuales, usando el número promedio por edad proveniente del análisis de cohortes. Las varianzas de las estimaciones de la abundancia dentro de las regiones son grandes para los peces de O, 1, Y5 años de edad, pero pequeñas para aquellos de entre 1.5 Y4 años de edad, excepto durante períodos de poca actividad pesquera. Las estimaciones de la tasa de captura media anual para todo el Océano Pacífico oriental están correlacionadas positivamente de forma significativa con las estimaciones de la abundancia media del análisis de las cohortes para los grupos de edad de entre 1.5 y 4 años. Se espera que los índices de abundancia por edad basados en las tasas de captura sean útiles, en conjunto con datos de la biología reproductiva, para estimar la producción total de huevos por regiones. Las estimaciones podrían asimismo ser útiles para la comprensión de las variaciones geográficas y temporales de la disponibilidad específica por edad a los barcos cerqueros, y también las migraciones específicas por edad. (PDF contains 35 pages.)

Age- or length-based methods of growth estimation. What drives the choice?

Based upon a global comparison of over 400 fisheries, the Principal Components Analysis (PCA) methodology was used to identify factors affecting the choice of growth estimation methods. Of the six factors examined, the growth rate (K) and asymptotic length (L8) explained most of the variations. Financial resources, i.e., Gross National Product (GNP), and latitude were also important factors.

Estimation of year class abundance and mortality of yellowfin tuna in the Eastern Tropical Pacific

ENGLISH: Age composition of catch, and growth rate, of yellowfin tuna have been estimated by Hennemuth (1961a) and Davidoff (1963). The relative abundance and instantaneous total mortality rate of yellowfin tuna during 1954-1959 have been estimated by Hennenmuth (1961b). It is now possible to extend this work, because more data are available; these include data for 1951-1954, which were previously not available, and data for 1960-1962, which were collected subsequent to Hennemuth's (1961b) publication. In that publication, Hennemuth estimated the total instantaneous mortality rate (Z) during the entire time period a year class is present in the fishery following full recruitment. However, this method may lead to biased estimates of abundance, and hence mortality rates, because of both seasonal migrations into or out of specific fishing areas and possible seasonal differences in availability or vulnerability of the fish to the fishing gear. Schaefer, Chatwin and Broadhead (1961) and Joseph etl al. (1964) have indicated that seasonal migrations of yellowfin occur. A method of estimating mortality rates which is not biased by seasonal movements would be of value in computations of population dynamics. The method of analysis outlined and used in the present paper may obviate this bias by comparing the abundance of an individual yellowfin year class, following its period of maximum abundance, in an individual area during a specific quarter of the year with its abundance in the same area one year later. The method was suggested by Gulland (1955) and used by Chapman, Holt and Allen (1963) in assessing Antarctic whale stocks. This method, and the results of its use with data for yellowfin caught in the eastern tropical Pacific from 1951-1962 are described in this paper. SPANISH: La composición de edad de la captura, y la tasa de crecimiento del atún aleta amarilla, han sido estimadas por Hennemuth (1961a) y Davidoff (1963). Hennemuth (1961b), estimó la abundancia relativa y la tasa de mortalidad total instantánea del atún aleta amarilla durante 1954-1959. Se puede ampliar ahora, este trabajo, porque se dispone de más datos; éstos incluyen datos de 1951 1954, de los cuales no se disponía antes, y datos de 1960-1962 que fueron recolectados después de la publicación de Hennemuth (1961b). En esa obra, Hennemuth estimó la tasa de mortalidad total instantánea (Z) durante todo el período de tiempo en el cual una clase anual está presente en la pesquería, consecutiva al reclutamiento total. Sin embargo, este método puede conducir a estimaciones con bias (inclinación viciada) de abundancia, y de aquí las tasas de mortalidad, debidas tanto a migraciones estacionales dentro o fuera de las áreas determinadas de pesca, como a posibles diferencias estacionales en la disponibilidad y vulnerabilidad de los peces al equipo de pesca. Schaefer, Chatwin y Broadhead (1961) y Joseph et al. (1964) han indicado que ocurren migraciones estacionales de atún aleta amarilla. Un método para estimar las tasas de mortalidad el cual no tuviera bias debido a los movimientos estacionales, sería de valor en los cómputos de la dinámica de las poblaciones. El método de análisis delineado y usado en el presente estudio puede evitar este bias al comparar la abundancia de una clase anual individual de atún aleta amarilla, subsecuente a su período de abundancia máxima en un área individual, durante un trimestre específico del año, con su abundancia en la misma área un año más tarde. Este método fue sugerido por Gulland (1955) y empleado por Chapman, Holt y Allen (1963) en la declaración de los stocks de la ballena antártica. Este método y los resultados de su uso, en combinación con los datos del atún aleta amarilla capturado en el Pacífico oriental tropical desde 1951-1962, son descritos en este estudio.

Validation of daily growth increments and estimation of growth rates of larval and early juvenile black skipjack, Euthynnus lineatus, using otoliths

ENGLISH: Increments in otoliths (sagittae) were examined, using light and scanning electron microscopy, to determine ages and estimate growth rates of larval and early-juvenile black skipjack, Euthynnus lineatus. Larvae and juveniles were collected between 1987 and 1989 from coastal waters of Panama in the eastern Pacific Ocean. Results from a laboratory experiment indicated that immersion for 6 and 12 hours in a 200 mg/L solution of tetracycline hydrochloride adequately marks otoliths and that increments are formed daily in the sagittae of postflexion larvae and early juveniles. Further, survival rates of tetracycline-treated fish were not significantly different from those of control fish. Growth rates were derived from length-age relationships of 218 field-collected specimens ranging in size from 5.7 to 20.3 mm SL. A growth rate of 0.70 mm/d was estimated from the weighted regression of standard length on age for all specimens. This rate lies within the range reported for larvae and early juveniles of other species of subtropical and tropical scombrids. Growth rates of postflexion larvae and early juveniles were not significantly different between the rainy season in July-August 1988 and the dry, upwelling season in January-February 1989. Growth was, however, significantly more variable for older individuals in July-August than in January-February, and may correspond, in part, to seasonal patchiness of prey. The growth rates of the otoliths relative to fish length were also not significantly different between seasons; however, the otoliths were larger relative to the lengths of fish collected in the rainy season, which may reflect slower growth during earlier larval stages. SPANISH: Se examinaron incrementos en otolitos (ságitas), usando microscopia de luz y de barrido electrónico, a fin de determinar la edad y estimar las tasas de crecimiento de barriletes negros, Euthynnus lineatus, larvales y juveniles tempranos. Entre 1987 y 1989 se capturaron larvas y juveniles en las aguas costeras de Panamá en el Océano Pacífico oriental. Los resultados de un experimento de laboratorio indicaron que una inmersión de 6 a 12 horas de duración en una solución de 200 mg/L de hidrocloro de tetraciclina marca los otolitos adecuadamente y que los incrementos se forman a diario en las ságitas de larvas en postflexión y juveniles tempranos. Además, las tasas de supervivencia de los peces tratados con tetraciclina no fueron significativamente diferentes a aquellas de los peces de control. Se calcularon las tasas de crecimiento a partir de las relaciones de talla-edad de 218 especímenes de TE entre 5.7 y 20.3 mm capturados en el mar. Se estimó.una tasa de crecimiento de 0.70 mm/día a partir de la regresión ponderada de talla estándar sobre edad para todos los especímenes. Esta tasa cae dentro del rango reportado para larvas y juveniles tempranos de otras especies de escómbridos subtropicales y tropicales. Las tasas de crecimiento de larvas en postflexión y juveniles tempranos no fueron significativamente diferentes entre la temporada de lluvias en julio-agosto de 1988 y la temporada de sequía y afloramiento en enero-febrero de 1989. Sin emoargo, el crecimiento fue significativamente más variable para los individuos de mayor edad en julio-agosto que en enero-febrero, y quizás corresponda parcialmente a la irregularidad temporal de la abundancia de presas. Las tasas de crecimiento de los otolitos en relación a la talla de los peces tampoco fueron significativamente diferentes entre temporadas; sin embargo, los otolitos eran más grandes en relación a la talla en peces capturados en la temporada de lluvias, lo cual podría reflejar crecimiento más lento durante las etapas larvales más tempranas. (PDF contains 42 pages.)

Production of the chironomids of the Uchinsk Reservoir. [Translation of: Methods for the estimation of production of aquatic animals. (Handbook and papers) (ed. G. G. Vinberg) p226-239. Minsk, Vysheishaya Shkola, 1968]

The method of E.V. Borutski was used for determining the production of chironomids, that is, the dynamics of the number and biomass of the larvae were analysed, their death, a calculation of emergence and the number of deposited egg layings was carried out. In addition to the method of Borutski, the authors also calculated the seasonal dynamics of the number of larvae of the younger age stages in the microbenthos.

A-SCALA: an age-structured statistical catch-at-length analysis for assessing tuna stocks in the eastern tropical Pacific Ocean

English: We describe an age-structured statistical catch-at-length analysis (A-SCALA) based on the MULTIFAN-CL model of Fournier et al. (1998). The analysis is applied independently to both the yellowfin and the bigeye tuna populations of the eastern Pacific Ocean (EPO). We model the populations from 1975 to 1999, based on quarterly time steps. Only a single stock for each species is assumed for each analysis, but multiple fisheries that are spatially separate are modeled to allow for spatial differences in catchability and selectivity. The analysis allows for error in the effort-fishing mortality relationship, temporal trends in catchability, temporal variation in recruitment, relationships between the environment and recruitment and between the environment and catchability, and differences in selectivity and catchability among fisheries. The model is fit to total catch data and proportional catch-at-length data conditioned on effort. The A-SCALA method is a statistical approach, and therefore recognizes that the data collected from the fishery do not perfectly represent the population. Also, there is uncertainty in our knowledge about the dynamics of the system and uncertainty about how the observed data relate to the real population. The use of likelihood functions allow us to model the uncertainty in the data collected from the population, and the inclusion of estimable process error allows us to model the uncertainties in the dynamics of the system. The statistical approach allows for the calculation of confidence intervals and the testing of hypotheses. We use a Bayesian version of the maximum likelihood framework that includes distributional constraints on temporal variation in recruitment, the effort-fishing mortality relationship, and catchability. Curvature penalties for selectivity parameters and penalties on extreme fishing mortality rates are also included in the objective function. The mode of the joint posterior distribution is used as an estimate of the model parameters. Confidence intervals are calculated using the normal approximation method. It should be noted that the estimation method includes constraints and priors and therefore the confidence intervals are different from traditionally calculated confidence intervals. Management reference points are calculated, and forward projections are carried out to provide advice for making management decisions for the yellowfin and bigeye populations. Spanish: Describimos un análisis estadístico de captura a talla estructurado por edad, A-SCALA (del inglés age-structured statistical catch-at-length analysis), basado en el modelo MULTIFAN- CL de Fournier et al. (1998). Se aplica el análisis independientemente a las poblaciones de atunes aleta amarilla y patudo del Océano Pacífico oriental (OPO). Modelamos las poblaciones de 1975 a 1999, en pasos trimestrales. Se supone solamente una sola población para cada especie para cada análisis, pero se modelan pesquerías múltiples espacialmente separadas para tomar en cuenta diferencias espaciales en la capturabilidad y selectividad. El análisis toma en cuenta error en la relación esfuerzo-mortalidad por pesca, tendencias temporales en la capturabilidad, variación temporal en el reclutamiento, relaciones entre el medio ambiente y el reclutamiento y entre el medio ambiente y la capturabilidad, y diferencias en selectividad y capturabilidad entre pesquerías. Se ajusta el modelo a datos de captura total y a datos de captura a talla proporcional condicionados sobre esfuerzo. El método A-SCALA es un enfoque estadístico, y reconoce por lo tanto que los datos obtenidos de la pesca no representan la población perfectamente. Además, hay incertidumbre en nuestros conocimientos de la dinámica del sistema e incertidumbre sobre la relación entre los datos observados y la población real. El uso de funciones de verosimilitud nos permite modelar la incertidumbre en los datos obtenidos de la población, y la inclusión de un error de proceso estimable nos permite modelar las incertidumbres en la dinámica del sistema. El enfoque estadístico permite calcular intervalos de confianza y comprobar hipótesis. Usamos una versión bayesiana del marco de verosimilitud máxima que incluye constreñimientos distribucionales sobre la variación temporal en el reclutamiento, la relación esfuerzo-mortalidad por pesca, y la capturabilidad. Se incluyen también en la función objetivo penalidades por curvatura para los parámetros de selectividad y penalidades por tasas extremas de mortalidad por pesca. Se usa la moda de la distribución posterior conjunta como estimación de los parámetros del modelo. Se calculan los intervalos de confianza usando el método de aproximación normal. Cabe destacar que el método de estimación incluye constreñimientos y distribuciones previas y por lo tanto los intervalos de confianza son diferentes de los intervalos de confianza calculados de forma tradicional. Se calculan puntos de referencia para el ordenamiento, y se realizan proyecciones a futuro para asesorar la toma de decisiones para el ordenamiento de las poblaciones de aleta amarilla y patudo.