Timing of births and juvenile mortality in the South American fur seal in Peru

In most species, synchronous, seasonal reproduction is usually associated with higher offspring survival in animals giving birth around the peak, relative to those breeding at the extremes of the reproductive season. In contrast, in the South American fur seal ( Arctocephalus australis ) at Punta San Juan, Peru, females pupping around the peak of births had a greater probability of losing their pups and/or tended to lose them at an earlier age than females breeding early or late in the season. However, the timing of breeding in this population varies little between years and is consistently synchronous. Individual females maintained their relative breeding times in consecutive years, regardless of whether or not they lost their pup the previous year. Thus, pup mortality seems to have no effect on the timing of reproduction in this population.
High breeding densities and the consequent high pup mortality in the S. American fur seal in Peru may have resulted from intense poaching outside protected areas and are of recent origin. The reproductive synchrony in this population could have originally evolved as a response to seasonal variations in food availability and weather conditions, differences in female or pup body condition, predation pressure, sexual selection and/or harassment avoidance, but at the present high density levels has become maladaptive.     

Sex‐specific reproductive investment of summer spawners of Illex argentinus in the southwest Atlantic

Energy investment in reproduction and somatic growth was investigated for summer spawners of the Argentinean shortfin squid Illex argentinus in the southwest Atlantic Ocean. Sampled squids were examined for morphometry and intensity of feeding behavior associated with reproductive maturation. Residuals generated from length‐weight relationships were analyzed to determine patterns of energy allocation between somatic and reproductive growth. Both females and males showed similar rates of increase for eviscerated body mass and digestive gland mass relative to mantle length, but the rate of increase for total reproductive organ weight relative to mantle length in females was three times that of males. For females, condition of somatic tissues deteriorated until the mature stage, but somatic condition improved after the onset of maturity. In males, there was no correlation between somatic condition and phases of reproductive maturity. Reproductive investment decreased as sexual maturation progressed for both females and males, with the lowest investment occurring at the functionally mature stage. Residual analysis indicated that female reproductive development was at the expense of body muscle growth during the immature and maturing stages, but energy invested in reproduction after onset of maturity was probably met by food intake. However, in males both reproductive maturation and somatic growth proceeded concurrently so that energy allocated to reproduction was related to food intake throughout the process of maturation. For both males and females, there was little evidence of trade‐offs between the digestive gland and reproductive growth, as no significant correlation was found between dorsal mantle length‐digestive gland weight residuals. The role of the digestive gland as an energy reserve for gonadal growth should be reconsidered. Additionally, feeding intensity by both males and females decreased after the onset of sexual maturity, but feeding never stopped completely, even during spawning.     

Lipid dynamics in the viperine snake, Natrix maura, from the Ebro Delta (NE Spain)

Body reserves play a major role in several aspects of vertebrate biology. Accurate identification and quantification of body reserves constitute a useful contribution to the better understanding of the energetic costs of reproduction and the implication of food availability in life history traits of organisms. In this study, lipid content in fat bodies, liver and muscle of the viperine snake (Natrix maura) were measured along an active season. Samples were collected monthly from a natural population of the Ebro Delta Natural Park (NE Spain). This methodology pointed out that lipids stored in fat bodies were the main energetic source during reproduction. In addition, lipids stored in liver appeared to be critical for vitellogenesis, while lipids stored in muscle turned out to be a supplementary energetic resource to fuel reproductive effort. Relationships between changes in body reserves and prey availability in canals of the Ebro Delta were also considered. In males, lipid reserves presented a positive correlation with food availability. On the contrary, lipid reserves in female viperine snakes decreased during vitellogenesis even though food availability increased in this period, which suggests a quick transfer of body lipids to clutch. In April, when rice fields of the Ebro Delta were dry and aquatic prey was scarce for viperine snakes, males and females presented a lower lipid content in fat bodies, liver and muscle than they did in other months, showing a clear link between prey availability and body reserves during food shortage. Thus, patterns of variation in fat levels indicated that Natrix maura is a capital breeder since it acquires resources in advance and stores them until they are invested during the reproductive period. Nevertheless, the shortage in April forces Natrix maura to turn into a facultative income breeder to fuel vitellogenesis. Finally, fat reserves in body components were compared with an estimate of body condition calculated by the residuals from the regression of body mass on body length. In male viperine snakes, the estimate of body condition was correlated with fat levels, revealing that this index is useful to measure condition in living individuals. On the contrary, body condition in females was not correlated with fat levels, which suggests that it is not appropriate to infer condition in female viperine snakes since it depends on the reproductive status of the individuals.     

Age, growth and reproduction of Aphanius iberus (Cuv. & Val., 1846) in the lower reaches of the Guadalquivir river (south-west Spain)

SUMMARY. 1. Age, growth and reproduction in an Atlantic population of the small fish, Aphanius iberus (Cuv. & Val., 1846), from the estuary of the Guadalquivir river were studied.
2. The growth period was from April to September and the annulus on the scales was formed in April.
3. The life-span was short with a winter age-structure of 95.9% 0+ fish, 3.9% 1+ and 0.2% 2+.
4. The fecundity of this stock, before the beginning of reproduction, is represented by the formula: Fec=0.907 TL (cm)^4.099. During the reproductive period 1+ females contained more, smaller eggs g⁻¹ of gonad than 0+ fish.
5. Males matured before females, but females were larger than males. The overall sex ratio did not differ from 1:1 but males dominated significantly during most of the spawning period.
6. Aphanius iberus spawned intermittently during the reproductive period. From April to July, 1+ group specimens (7–12 months old) reproduced. Their offspring (0+ group; 3–4 months old) reproduced from July to September. Very few of the 1+ group specimens survived to spawn the following year.
7. Mean gonadosomatic indices were much higher in the 1+ group than in the 0+ group and in females compared to males.
8. The loss of condition and the disappearance of the 1 + group during spawning, may indicate the cost of a prolonged high level of reproductive effort.
9. In this productive environment, the life history of Aphanius iberus is similar to that of Gambusia affinis (Baird & Girard, 1853) and Fundulus heteroclitus (Linnaeus) found in the same area.     

Is negative density‐dependent reproduction regulated by density‐induced stress in root voles? Two field experiments

Density dependence in reproduction plays an important role in stabilizing population dynamics via immediate negative feedback from population density to reproductive output. Although previous studies have shown that negative density‐dependent reproduction is associated with strong spacing behavior and social interaction between individuals, the proximal mechanism for generating negative density‐dependent reproduction remains unclear. In this study, we investigated the effects of density‐induced stress on reproduction in root voles. Enclosed founder populations were established by introducing 6 (low density) and 30 (high density) adults per sex into per enclosure (four enclosures per density in total) during the breeding season from April to July 2012 and from May to August 2015. Fecal corticosterone metabolite (FCM) levels, reproductive traits (recruitment rate and the proportion of reproductively active individuals), and founder population numbers were measured following repeated live trapping in both years. The number of founders was negatively associated with recruitment rates and the proportion of reproductively active individuals, displaying a negative density‐dependent reproduction. FCM level was positively associated with the number of founders. The number of founder females directly affected the proportion of reproductive females, and directly and indirectly through their FCM levels affected the recruitment rate; the effect of the number of male founders on the proportion of reproductive males was mediated by their FCM level. Our results showed that density‐induced stress negatively affected reproductive traits and that density‐induced stress is one ecological factor generating negative density‐dependent reproduction.     

Analysis of the cycle of reproduction of Sardina pilchardus (Walbaum, 1792) off the Moroccan Atlantic coast

This work focuses on aspects of reproductive biology of Sardina pilchardus from the Atlantic coast of Morocco. The mean values of batch fecundity estimated for the species is 23150(+/-1301) oocytes for a mean size of 19.5(+/-0.49) cm, the mean relative fecundity being 346(+/-7.34) oocytes per gram of female without ovary. Batch fecundity increases with total length and body weight without ovary. Sizes at first sexual maturity (L50) are reached for males and females at 15.8(+/-0.29) cm and 15.8(+/-0.35) cm, respectively. The spawning period for the population extends between October and July and the spawning peak occurs from October to February. However, the small sardines (14.5-17 cm) in their first reproduction spawn between November and June, whereas larger fish (17.5-25 cm) spawn between October and July. The factor of condition (K) increased in summer during the sexual resting phase. It is weak in winter during the period of reproduction. Regarding, the sex ratio, there was no significant difference in the number of males and females.     

Is bigger better? The relationship between size and reproduction in female Asian elephants

The limited availability of resources is predicted to impose trade‐offs between growth, reproduction and self‐maintenance in animals. However, although some studies have shown that early reproduction suppresses growth, reproduction positively correlates with size in others. We use detailed records from a large population of semi‐captive elephants in Myanmar to assess the relationships between size (height and weight), reproduction and survival in female Asian elephants, a species characterized by slow, costly life history. Although female height gain during the growth period overlapped little with reproductive onset in the population, there was large variation in age at first reproduction and only 81% of final weight had been reached by peak age of reproduction at the population level (19 years). Those females beginning reproduction early tended to be taller and lighter later in life, although these trends were not significant. We found that taller females were more likely to have reproduced by a given age, but such effects diminished with age, suggesting there may be a size threshold to reproduction which is especially important in young females. Because size was not linked with female survival during reproductive ages, the diminishing effect of height on reproduction with age is unlikely to be due to biased survival of larger females. We conclude that although reproduction may not always impose significant costs on growth, height may be a limiting factor to reproduction in young female Asian elephants, which could have important implications considering their birth rates are low and peak reproduction is young – 19 years in this population.     

Energy versus risk: costs of reproduction in free‐ranging pythons in tropical Australia

Abstract Data from a 12‐year field study have allowed us to quantify ‘costs of reproduction’ in a natural population of water pythons (Liasis fuscus) in tropical Australia. Both sexes of pythons cease feeding during the reproductive season. For males, this involves fasting for a 6 week period. Adult males lose weight rapidly over this period (approximately 17% of their body mass) but regain condition in the following months, and do not experience reduced survival. In contrast, reproductive adult females cease feeding for 3 months, lose an average of 44% of their body mass over this period, and experience increased mortality. A causal link between reproductive output and reduced female survival is supported by (i) a decrease in survival rates at female maturation; (ii) a correlation between survival rates and frequency of reproduction, in a comparison among different size classes of adult pythons; and (iii) a lowered survival rate for females that allocated more energy to reproduction. Hence, both sexes experience substantial energy costs of reproduction, but a relatively higher energy cost translates into a survival cost only in females. Such non‐linearities in the relationship between energy costs and survival costs may be widespread, and challenge the value of simple energy‐based measures of 'reproductive effort’.     

Condition, growth and reproduction in female red lechwe (Kobus leche leche Gray 1850)

Information on condition, growth and reproduction was collected from a sample of 155 female lechwe in the Linyanti Swamp, northern Botswana. Condition varied significantly with age, reproductive state and season and indications were found of relationships between body mass and the attainment of puberty in young females and body mass and the fertility of adult females. There were also indications that the population was under nutritional stress because of high water levels and that this had caused reductions in the growth rate and fertility of young females. It is suggested that the sensitivity of young females to adverse environmental conditions could be exploited to optimize population monitoring for conservation purposes by using them as an indicator class, rather than monitoring the entire population. Because of relationships between condition, growth and reproduction, it is also suggested that long-term monitoring of condition should form part of any effort to understand population dynamics.     

Reproductive biology of the broad‐snout [Chondrostoma holmwoodii (Boulenger, 1896)] in Tahtalı Dam Lake, İzmir,Turkey

The study aimed to determine the reproductive characteristics of the broad‐snout Chondrostoma holmwoodii population (spawning season, number of eggs and first age of sexual maturity) in order to assess the population status in the Tahtal? Dam Lake reservoir. A total of 1208 specimens were examined of which 72.1% were females and 27.9% males (female:male ratio = 1 : 0.40); the percentage of males, however, declined in the higher age classes (IV and V). The monthly determined gonadosomatic index indicated that the peak reproduction period of the species is in April and early May. The study covered age classes I–V, with specimens at age II reaching maturity. The total number of eggs per female varied from 8000 to 41 700, the mean number of eggs increasing proportionally with age of the spawner. Egg diameters also increased during the 5‐month ripening period from December to April.     

Age-specific resource investment strategies: evidence from female Richardson's ground squirrels (Spermophilus richardsonii)

To avoid a possible cost to their future survival and/or reproduction, individuals must balance their somatic and reproductive investments. The van Noordwijk and De Jong model of resource investment predicts that investments into reproduction and soma can vary among individuals of a population based on the variation in the total amount of energy that individuals acquire. With principal components analysis (PCA), we created two axes of life history for female Richardson's ground squirrels Spermophilus richardsonii : an index of total energy investment (PC1) and an index of investment tactic (PC2). Using these indices, we examined patterns of resource allocation to reproductive and somatic investments. Because yearling female Richardson's ground squirrels complete growth to adult size during pregnancy and early lactation, their somatic needs exceed those of older, fully grown females. Therefore, we predicted that yearlings would show more evidence of a tradeoff between reproductive and somatic investments compared with older females. Both yearling and older females invested four to five times more mass into their litters than into their own body mass. With increasing total investment, yearling females increased investment in both reproduction and themselves, whereas older females invested relatively more in reproduction than themselves. Regardless of age, females that emerged heavier from hibernation invested fewer resources into themselves and more into their litters. Variation in total energy investment and investment tactic indices was similar for yearling and older females. Contrary to our prediction, however, yearling females showed positive associations between reproductive and somatic investments, whereas older females exhibited showed no significant association between reproductive and somatic investments.     

Sociality and individual fitness in yellow-bellied marmots: insights from a long-term study (1962–2001)

Theoretical and empirical studies suggest that the age of first reproduction (the age at which reproduction begins) can have a substantial influence on population dynamics and individual fitness. Using complete survival and reproductive histories of 428 female yellow-bellied marmots (Marmota flaviventris) from a 40-year study (1962-2001), we investigated causes and fitness consequences of delayed maturity. Most females (86%) died without reproducing. The age of first reproduction of females that survived to reproduce at least once (n=60) ranged from 2 to 6 years. Females maturing later did not have a larger lifetime number of successful reproductive events or offspring production, nor did they experience improved survival. Females reproducing earlier had a higher fitness than those that delayed maturity. These results suggest that the net cost of early maturity was less than fitness benefits associated with early onset of reproduction, and that age of first reproduction in our study population is under substantial directional selection favoring early maturity. We conclude that female yellow-bellied marmots delay onset of reproduction not because of fitness benefits of foregoing reproduction at an earlier age, but due to the social suppression of reproduction by older, reproductive females, which enhances their own fitness to the detriment of the fitness of young females. Our results indicate that female yellow-bellied marmots that survive to reproduce may act to increase their own direct fitness, and that social suppression of reproduction of young females is a part of that strategy.     

Frequency and effort of reproduction in female Vipera aspis from a southern population

The frequency of reproduction of the asp viper (Vipera aspis, Viperidae) was studied in a population living along the coasts of central Italy. An annual reproductive cycle seemed to be the rule during the 5-year study period. Annual reproduction, high average mass of reproductive females, and large size of neonates, compared with other northern or continental populations, are presumably due to the particularly suitable climatic conditions of the area, as in most coastal habitats of the Mediterranean region. Such a scenario should influence the extent of the feeding period, which allows females, within a few months after parturition, to regain their previous body condition and reproduce again the following year.     

新热带区两种潮虫的生态特征(甲壳纲:等足目)

对体型大小不同的两种陆生等足类动物进行形态特征和行为特征研究,探讨环境变化对其繁殖特征和种群结构的影响。自2004年5月至2005年4月,在巴西Parque Estadual de Itapua进行野外采样,其中每月采集12个凋落物和土壤样品,实验室手拣法收集等足类,共获得2540只,同时进行个体性别鉴定与测量。结果表明,Atlantoscia floridana平均密度为114只Pm2,Balloniscus glaber平均密度为133只Pm2。A.floridana雌性个体全年可育,B.glaber雌性个体只在早春至初夏时段可育,其中春季这两种动物雌性可育个体数量高。2005年夏季干旱,该气候因子对两种动物种群影响存在差异,其中干旱使A.floridana种群减少,而B.glaber种群基本保持稳定,两种动物间形态结构不同是上述差异的主要原因,尤其是B.glaber,特殊的体型和腹肢肺的存在使其更能忍耐干燥。此外,B.glaber可能也依赖行为策略适应环境,例如挖掘活动,在不利气候条件下挖掘活动能够增加年轻个体生存的机会,也可能对成年个体起到保护作用。对于A.floridana来说,它具有连续繁殖、性成熟时间短和高雌性生育比率的特征,这些策略是对其幼体死亡率高的一种反应     

Reproductive Inhibition of Female Peromyscus leucopus: Female Competition and Behavioral Regulation

Maturational delay of young female mice as the result of exposureto grouped female odors and reproductive inhibition as the resultof exposure to isolated adult females have both been observed(Drickamer, 1974; Skryja, 1978). Each has the potential to reducethe growth rate of populations. Reductions in a female's reproductionfacilitated by social stimulation from other females, whileeffective in reducing population growth, may in the case, ofmaturational delay and reproductive inhibition be an epiphenomenonor exadaptation of selection for improved relative reproductivesuccess in the females possessing these abilities. The ultimateoutcome of these selective processes may be the buffering ofpopulation numbers, but the selective forces may operate tomaximize a female's relative reproductive success. A females'relative reproductive success can be maximized by either increasingher own reproduction or decreasing the reproductive output ofother females. A body of evidence exists to suggest that inPeromyscus mamculatus and Peromyscus leucopus, females are physiologicallyconstrained and unable to increase their own reproduction. Ifthe assumption of physiological restraint is correct, then themost effective way for females to maximize their relative reproductivesuccess is to reduce the reproductive output of their competitors.In this paper, maturational delay and reproductive inhibitionas they affect both the adult female and young females are discussed.Examination of these effects reveals that while they can beeffective in population regulation, their main function andthe selective process that produced them is at the level ofindividual reproduction.     

Unpredictable perturbation reduces breeding propensity regardless of pre‐laying reproductive readiness in a partial capital breeder

Theoretically, individuals of migratory species should optimize reproductive investment based on a combination of timing of and body condition at arrival on the breeding grounds. A minimum threshold body mass is required to initiate reproduction, and the timing of reaching this threshold is critical because of the trade‐off between delaying breeding to gain in condition against the declining value of offspring with later reproductive timing. Long‐lived species have the flexibility within their life history to skip reproduction in a given year if they are unable to achieve this theoretical mass threshold. Although the decision to breed or not is an important parameter influencing population dynamics, the mechanisms underlying this decision are poorly understood. Here, we mimicked an unpredictable environmental perturbation that induced a reduction in body mass of Arctic pre‐breeding (before the laying period) female common eiders Somateria mollissima; a long‐lived migratory seaduck, while controlling for individual variation in the pre‐laying physiological reproductive readiness via vitellogenin (VTG) – a yolk‐targeted lipoprotein. Our aim was to causally determine the interaction between body condition and pre‐laying reproductive readiness (VTG) on breeding propensity by experimentally reducing body mass in treatment females. We first demonstrated that arrival body condition was a key driver of breeding propensity. Secondly, we found that treatment and VTG levels interacted to influence breeding propensity, indicating that our experimental manipulation, mimicking an unpredictable food shortage, reduced breeding propensity, regardless of the degree of pre‐laying physiological reproductive readiness (i.e. timing of ovarian follicles recruitment). Our experiment demonstrates that momentary environmental perturbations during the pre‐breeding period can strongly affect the decision to breed, a key parameter driving population dynamics.     

高原鼢鼠繁殖特性与其栖息草地质量的关系

高原鼢鼠种群密度动态变化与其繁殖特性密切相关,而繁殖特性又与其栖息环境有关。为了解高原鼢鼠繁殖特性与其栖息草地质量的关系,本文在祁连山东段选择两个不同高原鼢鼠种群密度分布区,在其繁殖期调查各密度区雌雄个体不同月份的繁殖特性、草地生物量、植物组成、地下根系重量及根系可溶性糖含量、土壤紧实度和水分变化。通过多因素方差分析和独立样本T检验研究繁殖特性与草地质量的关系。结果显示：在高原鼢鼠繁殖期,5月为繁殖高峰期,6月进入繁殖末期;不同种群密度区之间个体繁殖强度无显著性差异（P>0.05）;雌性繁殖强度在繁殖高峰期无显著性差异（P>0.05）,而繁殖末期存在显著差异（P<0.05）;高原鼢鼠性比在繁殖高峰期表现为低密度区大于高密度区,而在繁殖初期和繁殖末期为高密度区大于低密度区;两个种群密度区之间,草地植物组成、草地地上生物量、可利用草地生物量和地下生物量无显著性差异（P>0.05）,而0-30 cm土层根系根可溶性糖含量存在显著性差异（P<0.05）;除高原鼢鼠采食深度（0-20 cm）外,不同密度区土壤紧实度无显著性差异（P>0.05）,而0-30 cm土层土壤水分含量存在显著性差异（P<0.05）。结果表明在一定种群密度下,高原鼢鼠繁殖特性的变化与地下根可溶性糖含量和土壤水分有关,而繁殖特性与草地植物学组成、草地生物量无显著关系。     

The population dynamics and sexual strategy of Anodonta woodiana (Bivalvia: Unionacea) in Plover Cove Reservoir, Hong Kong

A 12-month investigation of the population dynamics, growth, ageing, reproduction and sexuality of Anodonta woodiana (Lea, 1834) in Plover Cove Reservoir, Hong Kong, has been undertaken. Recruitment occurs in the summer, May through July being the period of peak incidence of glochidia on captive fish hosts. Juveniles grow rapidly to a shell length of 30 mm and are sexually mature by the following spring. A study of the population structure of A. woodiana indicated that most animals were less than 8 years old. Results of a Walford plot for this species indicated a maximum theoretical shell length (in Plover Cove) of 162·5 mm and a life span of approximately 12 years; the latter is in good agreement with age estimates for European unionids.
Anodonta woodiana is dioecious with females predominating in a ratio of 60:40. A very small number of individuals (0·3%) are hermaphrodite. Males possess mature gonads throughout the year whereas females come into reproductive condition during the spring. There was no evidence of shell form sexual dimorphism, as has been recorded for other unionaceans.
The reproductive strategy and time of gonad maturation of A. woodiana is compared with that of unionids from elsewhere and with other fresh water bivalves from southern China.     

Age and terminal reproductive attempt influence laying date in the thorn‐tailed rayadito

Age‐specific variation in reproductive effort can affect population dynamics, and is a key component of the evolution of reproductive tactics. Late‐life declines are a typical feature of variation in reproduction. However, the cause of these declines, and thus their implications for the evolution of life‐history tactics, may differ. Some prior studies have shown late‐life reproductive declines to be tied to chronological age, whereas other studies have found declines associated with terminal reproduction irrespective of chronological age. We investigated the extent to which declines in late life reproduction are related to chronological age, terminal reproductive attempt or a combination of both in the thorn‐tailed rayadito Aphrastura spinicauda, a small passerine bird that inhabits the temperate forest of South America. To this end we used long‐term data (10 years) obtained on reproductive success (laying date, clutch size and nestling weight) of females in a Chilean population. Neither chronological age nor terminal reproductive attempt explained variation in clutch size or nestling weight, however we observed that during the terminal reproductive attempt older females tended to lay later in the breeding season and younger females laid early in the breeding season, but this was not the case when the reproductive attempt was not the last. These results suggests that both age‐dependent and age‐independent effects influence reproductive output and therefore that the combined effects of age and physiological condition may be more relevant than previously thought.     

Reproductive seasonality of the detritivorous fish <Emphasis Type="Italic">Cyphocharax abramoides</Emphasis> (Kner, 1958) (Characiformes: Curimatidae) in flooded rivers of the eastern Amazon

The Lower Anapu River, Eastern Amazon, has a discrete flood pulse with narrow flooded areas due to the reduced influence of tides. In this environment, local precipitation might act as an important factor in the input of detritus, which can serve as a trigger for fish reproductive period. Therefore, this paper aims to determine the effects of local precipitation on the reproductive ecology of an abundant detritivorous fish in flooded rivers. Between May 2012 and April 2013, 886 specimens (471 females and 415 males) were captured monthly using gill nets. Individuals were measured, weighted, sexed, and the gonadal development stage was defined macroscopically and confirmed histologically. Sex ratio, length-weight relationship (LWR), condition factor (k), Gonadosomatic Index (GSI), relative frequency of maturity stages, and size at first gonadal maturity (L₅₀) were estimated and tested in relation to rainfall periods. There was no difference between the expected sex ratio (1:1), and both males and females showed polyphasic growth. The condition factor indicated higher values at the beginning of the rainy season and lower values during the dry season for both sexes. GSI and relative frequency of mature individuals indicated a long reproduction period with gonadal investment starting within the dry season until early in the rainy season for spawning within the rains. The L₅₀ value showed that males reach sexual maturity before females. Results indicated a reproductive pattern strongly influenced by rainfall, corroborating the importance of local precipitation for lacustrine environments.