Reproductive biology of Hemiramphus brasiliensis and H. balao (hemiramphidae): maturation,spawning frequency,and fecundity

Analyses of life-history data show that both the size-specific batch fecundities and the age-specific spawning frequencies differ for two halfbeak species, Hemiramphus brasiliensis, the ballyhoo, and H. balao, the balao. Halfbeak ages were determined from sectioned otoliths; histological data was used to describe oocyte development and estimate spawning frequency; and batch fecundity was measured from counts of whole oocytes in final maturation. Hemiramphus brasiliensis lived longer (4 versus 2 years) and had a higher survival rate (14.9% versus 7.5% annually) than H. balao did. Of the two species the larger and longer-lived congener, H. brasiliensis, reached sexual maturity at a larger size (fork length 198 versus 160 mm). The spawning period of age-0 females was strongly related to season, whereas spawning by older females occurred throughout the year. Reproduction by both species peaked during late spring or early summer, and all mature females were spawning daily during April (H. brasiliensis) or June (H. balao). This is the first demonstration of iteroparity for the family Hemiramphidae. H. brasiliensis had a lower batch fecundity (about 1164 versus 3743 hydrated oocytes for a 100-g female) than H. balao did. Such low batch fecundities are typical of the order Beloniformes, but quite different from those of other fishes that live in association with coral reef habitats. H. balao's higher batch fecundity is consistent with the life-history theory that predicts higher numbers of eggs for shorter-lived species; this is possible because H. balao produces smaller hydrated oocytes than H. brasiliensis (modal diameter about 1.6 versus 2.4 mm). The high spawning frequency of Hemiramphus species compensates for their low batch fecundity. The annual fecundity of both species is similar to that of other reef fish species, after adjusting for body size and spawning frequency. The lifetime fecundity of H. balao was very similar to that of H. brasiliensis, after accounting for the differences in survival for each species. This suggests a fine tuning of different reproductive traits over the entire life cycle that results in roughly equivalent lifetime fecundity for both species.     

A hierarchical response to differences in ration size in the reproductive performance of female three-spined sticklebacks

The response of female three-spined sticklebacks ( Gasterosteus aculeatus ) to differences in food ration during the breeding season was quantified for several variables related to reproductive performance. The measured protein and lipid contents (mg g⁻¹) of eggs were unaffected by ration. Egg size increased with an increase in both ration and female size, but the proportional increase in egg size was much smaller than the proportional increase in ration. The best predictor of mean batch (clutch) fecundity and weight was female size. There was a small but significant increase in batch fecundity with ration, but the increase was not directly proportional to the increase in ration. The rate of spawning, total breeding season fecundity and total weight of eggs spawned over the breeding season were sensitive to ration with breeding season fecundity and weight increasing in direct proportion to ration. Thus, in female sticklebacks, there is a hierarchy of sensitivity to ration in the response of variables related to reproduction.     

Reproductive biology of the armoured catfish Loricariichthys castaneus (Castelnau, 1855) in Lajes reservoir,southeastern Brazil

Sex ratio, gonadal development, breeding season and fecundity of the armoured catfish Loricariichthys castaneus were described to assess its reproductive strategy in a Brazilian tropical reservoir. In total, 226 specimens (199 females and 27 males) were captured from September 2005 to August 2006 and examined in the laboratory. Females outnumbered males and achieved sizes larger than 330 mm TL. Oocyte development, determined by histological analysis, was asynchronous with oocyte size, ranging from pre‐spawning (27–270 μm) to spawning (243–3460 μm), followed by a sharp decrease in the mean oocyte diameter postspawning (590–730 μm) as the spawning proceeded. Spawning occurred throughout most of the year, peaking in August–September and reaching a low in April–May, according to variations in GSI and frequencies of stages of gonadal development. Batch fecundity ranged from 242 to 833 vitellogenic oocytes (relative fecundity = 2.27 oocytes g^?1), averaging 483 oocytes, and was positively related to gonad weight (P = 0.00003). Oocyte diameters ranged from 0.027 to 5.59 mm, with vitellogenic diameters ranging from 2.08 to 5.59 mm. Continuous development of oocytes throughout the year suggests that L. castaneus presents indeterminate fecundity and is a batch‐spawner. These attributes, associated with parental care and a wide reproductive period, correspond to an equilibrium strategy that has proved to be effective in the Lajes reservoir.     

Effect of spawning number and ration on reproductive performance of the batch-spawning three-spined stickleback Gasterosteus aculeatus

An experiment quantified the effect of food ration and spawning number on the breeding season reproductive performance of batch-spawning, female three-spined sticklebacks Gasterosteus aculeatus . Individually housed females were maintained on fixed rations of 2, 4, 8 or 16% of their initial postspawning mass of live enchytraeid worms from immediately after their first spawning until they ceased spawning. Number of spawnings correlated positively with ration. Total breeding season reproductive investment (total wet mass of eggs produced as a percentage of initial postspawning female mass) ranged from 38% at the 2% ration to 147% at the 16% ration. At the lower rations, postspawning mass, batch fecundity and wet and dry masses of the batch declined over successive spawnings, with the rate of decline inversely related to ration. At the highest ration, there was no decline in batch fecundity and postspawning mass increased over successive spawnings. Mean reproductive investment per inter-spawning interval was higher at the highest ration, but at all rations declined over successive spawnings. Mean reproductive effort per inter-spawning interval (wet mass of eggs spawned as a percentage of the wet mass of food consumed over the inter-spawning interval) was inversely related to ration. At the higher rations, reproductive effort showed no trend over successive spawnings. Neither spawning number nor ration had a systematic effect on egg diameter, wet mass per egg, dry mass per egg or total lipid content of the eggs. If the rate of food intake was insufficient, although batch fecundity declined, the main adjustment was a reduction in the number of spawnings in the breeding season.     

Reproductive ecology of Brazilian freshwater fishes

We used metadata on nine reproductive traits of 67 species of Brazilian iteroparous, oviparous, teleost freshwater fishes to test phenotypes associations to discriminate species that only spawn in large rivers (lotic fishes) from those capable to spawn in lentic habitats (lentic fishes). We tested the hypothesis that lotic fishes present spawning migration, shorter spawning season, single spawning, no parental care, free eggs, higher relative fecundity, faster embryogenesis, and larger size, while lentic fishes present no spawning migration, longer spawning season, multiple spawning, parental care, adhesive eggs, lower relative fecundity, slower embryogenesis, and smaller size. Our analyses supported the hypothesis but not all phenotypic associations satisfied it, specifically with regard to lentic fishes or to pairs of phenotypes typical of lentic fishes. We also concluded that spawning in large rivers is a better predictor of bionomic characters than spawning in lentic habitats, and lotic fishes are specialists compared to lentic fishes.     

Effect of seawater temperature on reproductive seasonality and fecundity of Pseudoplexaura porosa (Cnidaria: Octocorallia): latitudinal variation in Caribbean gorgonian reproduction

Abstract. The majority of tagged colonies of Pseudoplexaura porosa in Bermuda were reproductive over 2 months in the summer. They spawned 5–8 d after the full moon, with a peak on the sixth day, similar to colonies in Panama. The months of spawning were August and September in 1998, but July and August in 1999 and 2000. This temporal difference between the months of spawning corresponded to inter-annual variations in seawater temperature profiles. Initial gamete development each year occurred only when the daily mean seawater temperature during the month before spawning exceeded 27°C. There was a significant positive relationship between reproductive effort (gamete volume) of colonies and rising seawater temperature in the month preceding spawning; this was true for both the initial and the second spawning months. The end of the reproductive season each year was triggered by the decline in seawater temperature past the summer maximum. The duration of the reproductive season of conspecifics at the central Caribbean reef of Panama is 2 months longer than in Bermuda. This can be explained by the smaller annual temperature range at the lower latitude and the earlier onset of temperatures favorable for gamete development. Fecundity estimates for members of P. porosa (mean oocyte and spermary densities) in Bermuda were lower than for conspecifics in Panama. The shorter reproductive season in Bermuda, in addition to the lower fecundity of colonies, indicates that reproduction in P. porosa is compromised at this high latitude reef.     

Reproductive guilds (maturation,spawning frequency and fecundity) in the black pomfret,Parastromateus niger (Carangidae) in the Kuwaiti waters of the Arabian Gulf

Oogenesis, oocyte maturation pattern, spawning rhythm, spawning frequency, batch fecundity and oocyte diameter–frequency distribution of the black pomfret, Parastromateus niger (Bloch, 1795) in Kuwaiti waters were investigated from October 2003 to September 2005, using histological and morphological methods. The process of development is divided into four major phases: (i) primary growth phase; (ii) secondary growth phase; (iii) maturation phase; and (iv) spawning phase, followed by the regressed phase. Development of the yolky oocyte is an asynchronous process resulting, by the time of oocyte maturation, in a clear differentiation between a ready batch of oocytes (ready for spawning) and a reserve pool. Consequently, P. niger is capable of spawning multiple times throughout the reproductive season. Spawning frequency estimates, based on final oocyte maturation (FOM) method indicated that the species spawns once every 2.8 days during an 8‐month spawning season lasting from February to September, with a potential annual number of 22.4 spawns. Batch fecundity (BF) (2132–2001 648, mean 406 010 eggs), was significantly positively related to both standard length (SL) (P < 0.05) and ovary‐free body weight (OFBW) (P < 0.05), both parameters being good predictors of BF (r² = 30.8% for SL, from 22 cm onwards, and r² = 29.6% for OFBW, from 129.5 g onwards). No significant differences in monthly BF were found throughout the spawning season. Relative batch fecundity was 336 eggs/g OFBW; thus, estimate for potential annual relative batch fecundity was 7526 eggs g^?1 OFBW. The oocyte diameter–frequency distribution analysis revealed a multimodal distribution (at 100–200, 300–400 and 500–700 μm), confirming the evidence of multiple spawning.     

Maturity and spawning of painted lizardfish,Trachinocephalus myops (Bloch and Schneider, 1801) in the southeastern China Sea

Ovaries of 1159 Trachinocephalus myops were examined, and six oocyte stages were histologically identified whereby each stage of ovary development was observed almost every month. The sex ratio was not different from 1 : 1 (P > 0.01). The weight‐length relationship was estimated as BW = 0.0077FL^3.1207 (r² = 0.976, n = 2256). Monthly composition of the oocyte stages gonadosomatic index (GSI), and condition factor (CF) were examined. The spawning season had two peaks: February to April and August to October. Sizes at 50% maturity were estimated at 18.0 and 18.7 cm FL for females and males, respectively. Fecundity estimates ranged from 11 100 to 426 200 (mean: 178 600), based on an oocyte diameter >0.2 mm. Batch fecundity was estimated to range from 1724 to 120 951 (mean: 60 200), germinal vesicle migration and mature oocytes >0.8 mm in diameter were counted, and the relative fecundity was estimated to range from 346 to 3115 (mean: 1087). The results suggest that reproductive activity occurred throughout the year, suitable for the highly variable environments utilized by this species.     

Reproductive biology ofAwaous guamensis, an amphidromous Hawaiian goby

Synopsis Spawning season, size at first reproduction, oocyte maturation, and fecundity ofAwaous guamensis, an amphidromous Hawaiian goby, were studied from June 1989 through May 1991 in the Wainiha River, Kau'ai, Hawai'i. Female fish larger than 73 mm standard length (SL) had mature gonads from August through December in 1989 and 1990. Gonadosomatic index (GSI) values for mature females ranged from 0.2 to 14.5 during the spawning season. Male fish larger than 64 mm SL had elevated GSI values from June 1989 through December 1989 and from August 1990 through December 1990. Mature sperm were found in two male fish collected in January and February. GSI values for mature males ranged from less than 0.01 to 4.0 in the spawning season. Size-frequency distributions of measurements of vitellogenic oocyte diameters and microscopic observations of oocytes indicated this species has group-synchronous oocyte development. Ovarian maturation stages examined over a 29-month period suggest that members of the stock spawned at different times within the spawning season, although mass spawning events have been documented for this species. Estimates of clutch sizes from nests measured in situ were comparable to estimates of potential fecundity from in vitro examination of ovaries, and indicated that female fish deposited an entire clutch during a spawning event. No evidence for multiple spawning by an individual fish in a single season was found. However, microscopic observations of brown bodies in some ovaries suggested that individual fish probably spawn more than once in a lifetime.     

Spawning frequency, fecundity, egg weight and spawning type of silver pomfret, Pampus argenteus (Euphrasen) (Stromateidae), in Kuwait waters

Silver pomfret, Pampus argenteus, were collected by fishing with drift gillnets on one spawning ground in Kuwait waters during 1998–2000. Fish size frequency, sex ratio, maturation cycle, spawning frequency, fecundity and egg weight were assessed. The length–weight relationship differed between sexes whereby females were significantly bigger than males. Spawning started in mid‐May and continued until early October. During this time the water temperature ranged from 26.0 to 32.8°C, salinity was ? 39.0‰ and water depth ranged between 5 and 12 m. Large females spawned earlier than young spawners and the overall percentage of males during the spawning period was 70.3%. Spawning occurred after 13.00 h, with peak spawning between 15.00 and 18.00 hours during outgoing tide. Mean daily spawning frequency amounted to 63.2%. Spawning activity was found to be associated with the lunar cycle and spawnings were concentrated during the first and third quarters of the moon period, indicating a semilunar reproduction cycle. It was concluded that a female would spawn at least six times during the season. No change was observed in relative fecundity during the peak spawning season (June–August). Average relative batch fecundity was 176.3 eggs g^?1 somatic weight (SW), corresponding to a relative total fecundity of 1058 eggs g^?1 SW, which is 1.5 times higher than estimates obtained from counting the standing stock of oocytes. Bigger fish produced heavier eggs and the egg weight decreased as the spawning season progressed. Based on gonadal cycles, oocyte size frequency distribution and total fecundity, we concluded that silver pomfret is a multiple batch spawner with indeterminate fecundity.     

Comparative oocyte morphology and fecundity of five characid species from São Francisco River basin,Brazil

The morphology of vitellogenic oocytes and the batch fecundity of five tropical forage species belonging to the family Characidae, were studied in 104 late‐maturing ovaries. Significant morphological differences between vitellogenic oocytes and ovarian follicles were found. The lowest batch fecundity values were recorded in Hemigrammus marginatus (480 ± 163) and Orthospinus franciscensis (1701 ± 562), which were the smaller species in terms of total length, body weight and oocyte diameter. The highest batch fecundity value was observed in Tetragonopterus chalceus (8384 ± 3944) having the highest GSI and oocyte diameter. Batch fecundity and gonad weight was highly correlated followed by body weight and total length. Relative fecundity was estimated per unit total length, body weight and gonad weight. The wide variation in fecundity observed between the specimens and the species analysed is possibly related to the multiple spawning reproductive strategy of these fishes. Since T. chalceus have higher values of batch and relative fecundity, it is concluded that this species has a higher reproductive potential than the other forage species studied.     

Reproduction and growth of Craterocephalus marjoriae and C. stercusmuscarum (Pisces: Atherinidae) in south-eastern Queensland, Australia

SUMMARY. 1. The reproductive cycle and growth of Craterocephalus marjoriae and Craterocephalus stercusmuscarum (Atherinidae) were studied in Brisbane, south-eastern Queensland, over a 17-month period (1981–82).
2. Both species had a 5-month breeding season in spring and summer, with a breeding peak in spring (September-October). Multiple spawning occurred but life time fecundity was not determined.
3. Gonosomatic index values and mean fecundity per female were higher in Craterocephalus marjoriae than in C. stercusmuscarum. Fecundity increased with body length in both species according to the relationship, F=aL^b. Growth rates were similar and growth was adequately described by the von Bertalanffy equation. Both species grew rapidly during the first year and reached maturity the following season. Reproductive strategies are considered in relation to seasonal flooding in the streams inhabited.     

The multiple spawning pattern of weakfish in the Chesapeake Bay and Middle Atlantic Bight

Weakfish Cynoscion regalis were collected from commercial fisheries in the Chesapeake Bay and the Middle Atlantic Bight (n=4380) during 1989–1992 and their reproductive biology assessed using the gonadosomatic index, macroscopic gonad stages, oocyte diameter distributions, microscopic whole oocyte analysis and histology. Sex ratios were approximately 3:1, females to males, in 1990–1992. Most fish (90%) attained sexual maturity by age 1 and at a small size. Estimated mean length at first maturity was: 164mm total length (TL) for males, and 170 mm TL for females. Weakfish spawn within the Chesapeake Bay, as far north as the Virginia/ Maryland border. Although spawning occurred during May–August and gonad development and initiation of spawning was synchronous, cessation of spawning was asynchronous. There was no indication that older fish exhibited a more extended spawning season than younger fish. Weakfish are multiple spawners with indeterminate fecundity. Oocyte development is asynchronous with oocytes of all stages being present in developed ovaries. Because of the complex and dynamic weakfish ovarian cycle, typical methods of assessing reproduction, such as the GSI and macroscopic gonad stages, are inadequate for this species if not used in conjunction with more detailed methods such as histology.     

湖南沅水下游?繁殖期内繁殖力和卵径的变化研究

为分析鱼类繁殖期内繁殖力和卵径变化趋势, 研究于4—10月在沅水下游逐月采集180尾?(Hemiculter leucisculus), 分析81尾雌性成熟个体的繁殖力。研究结果表明: ?的性体指数在5、6月最高, 其次在7、8和9月, 在4和10月最低。绝对繁殖力平均值为(30116±19390)粒, 以10000—30000粒为主, 相对繁殖力的平均值为(650±324)粒/g, 以400—800粒/g为主。绝对繁殖力与体长、空壳重呈幂函数关系(P<0.05)。绝对繁殖力和相对繁殖力在5和6月最大, 7、8和9月次之, 在4和10月最小。不同月份成熟卵子的卵径无显著差异, 补充卵子的卵径在5和6月最大, 暗示5和6月分批繁殖次数较多。不同繁殖时期的分批繁殖力、繁殖次数不一样, 对评估种群的繁殖潜能具有重要的指导意义。比较不同水系繁殖力发现, 湖泊种群的个体繁殖力普遍大于河流种群, 沅水种群的个体繁殖力跟湖泊种群的相似, 可能与其具有类似湖泊的缓流水或静水生境有关。研究揭示了?的基本繁殖特征, 为渔业资源管理提供了基础数据资料。     

Reproduction mode of European Bitterling (Rhodeus amarus,Bloch, 1782) determined through rapid oocyte counts and size determination using digital imaging

This study focuses on assessing the reproduction mode of an important model species for evolutionary and behavioural ecology by using digital image analysis: the European bitterling Rhodeus amarus (Bloch, 1782). Specifications of mode of reproduction were determined using oocyte size distribution, seasonal dynamics in mean oocyte diameter and total number of oocytes in ovary samples gained from April to July 2007. The rapid oocyte count was enabled by using lucia image analysis software, which also provided measurement and colour estimations of oocytes. Bitterling ovaries showed features typical for indeterminate spawners, i.e. a continuous distribution of oocyte size over the reproductive season and recruitment of new pre‐vitelogenic oocytes in the second half of the reproductive season. These results are consistent with the view that the European bitterling is a batch spawning fish with indeterminate fecundity.     

Reproductive strategy of the European perch,Perca fluviatilis Linnaeus, 1758 (Osteichthyes: Percidae) in the Anzali wetland,southwest Caspian Sea

Temporal variation in reproductive traits of geographically distributed fish is supposed to take place in response to the spatial and environmental variations. With regard to the wide distribution of the European perch in the northern hemisphere, important reproductive traits such as the initiation and duration of the spawning activity are likely to vary in different latitudinal gradients. In this study, reproductive biology of the European perch, Perca fluviatilis, is described, based on 324 specimens caught in the Anzali wetland (southwest Caspian Sea) between June 2008 and May 2009. The gonadosomatic index, oocyte frequency distribution and histological examination suggested a long vitellogenic process (October to February) and a short spawning season (January and February). The size‐frequency distribution of the oocytes showed that this perch is a species with group‐synchronous ovarian development. Ovarian development occurred only in one clutch of oocytes (700–900 μm oocyte diameter) with no indication of maturation of any subsequent clutch in the spawning season. The average of (realized) fecundity (±SD) was estimated to be 16177 ± 5846 eggs in late vitellogenic stage, which was lower than the potential fecundity (17188 ± 6917 eggs). Histological examination of the gonads revealed the existence of atretic oocytes in early vitellogenic stages (October and November). This investigation highlights the temporal variation in the initiation and duration of the reproductive activity of the European perch in this region compared to other geographical regions. The results emphasize the necessity of specific temporal management in fishing of European perch based on spatial differences in reproductive biology.     

Reproductive biology of an endemic,amphidromous goby Lentipes concolor in Hawaiian streams

Synopsis Egg size, fecundity, nest site selection, and breeding structure of the amphidromous Hawaiian goby Lentipes concolor were studied to determine if there was any relationship between these aspects of the breeding biology of this fish and the very long larval period of Hawaiian stream gobies. To do this, these parameters were compared with those known for other gobioid fishes. While eggs were small and females were found to lay up to four nests in a season, reproductive biology and mating behavior of L. concolor were typical of many gobies in spite of its amphidromous life history pattern. Lentipes concolor nests were found from October to June with a spawning peak in February. Nests tended to be positioned closer to the stream bank, and under larger rocks than if they were situated randomly. Depth and mean water column velocity over nest sites did not differ from that at randomly selected points in the stream.     

Reproductive patterns of blacktail comber (Serranus atricauda,Serranidae) from south-west Portugal seamounts

The knowledge of the reproductive strategy is essential for the comprehensive understanding of the population dynamics of any fish species. This study intends to define the spawning season, verify the type of fecundity regulation and estimate relative annual fecundity for Serranus atricauda inhabiting the south-west of Portugal seamounts (Gorringe, Ampère and Lion). Monthly samples, between September 2010 and December 2011, were obtained from commercial vessels. This species is synchronously hermaphroditic, and mature individuals were found all year round, with gonadosomatic index peak from February to September. No clear relation between hepatosomatic index and Fulton’s condition factor and gonadal development was found, and therefore, it is probable that reproductive success depends more on environmental conditions and food availability during the spawning season than on body energy reserves. No gap exists between pre-vitellogenic and vitellogenic oocytes and the oocyte size-frequency distribution during the main spawning season show no dominant cohort progressing through time. The number of standing stock of advanced vitellogenic oocytes showed an increase trend along the spawning season decreasing only at its end and the relative intensity of α-atresia was always low throughout this period with an increase towards the end. All these criteria corroborate the indeterminate fecundity type assigned to S. atricauda in previous studies. Mean relative annual fecundity estimated was 5,655 ± 4,321 oocytes per gram of gutted weight.     

Reproduction of the spiny puffer, Diodon holocanthus (Pisces: Diodontidae) in the continental shelf of Mexican Central Pacific

Diodon holocanthus is an important economic and ecological species of the demersal fish community, caught as bycatch from local shrimp fishery. The reproductive biology of this long-spine porcupinefish has not yet been described, and reproductive season, the sex ratio, length distribution, length at first gonad maturity, and the gonad macro and microscopic features are described. A total of 400 organisms, ranging from 5.0 to 40.3 cm (average 18.4 cm) total length, were caught from the continental shelf of the Central Mexican Pacific, from December 1995 and December 1998. Sex ratio was 1:0.86 females to males (n = 253). The length at which 50% of the individuals showed maturing gonads was 19.7 cm for females and 20.1 cm for males. Length of the smallest organism with ripe gonads was 12.2 cm for females and 13 cm for males. Four gonadal maturation stages were found in both sexes, and five oocyte development phases were identified. The oocyte development pattern is of asynchronous type, which means the species can reproduce several times a year. Testicle development is lobular type, as in most teleost fishes. Monthly mean values of the gonad-somatic index suggest the reproduction activity peaks in June, and September-December.     

Reproductive strategies of a temperate Australian sciaenid (teraglin,Atractoscion atelodus)

Worldwide there have been declines in stocks of exploited sciaenids. Despite a long history of exploitation, there has been no study of the life history characteristics of the fishery-important sciaenid, teraglin, Atractoscion atelodus. This study describes the reproductive characteristics of A. atelodus within the major area of its distribution and fishery, New South Wales (NSW) Australia. Characteristics studied included the size and age at maturity, fecundity, spawning mode and season, and these are compared to congeneric species. Atractoscion atelodus displays year-round batch spawning behaviour with asynchronous oocyte development and indeterminate fecundity. This is unusual for sciaenids, which typically display spring-summer spawning and only over a few months. The length and age at which 50% of the population matures for both males and females is 36 cm fork length (FL) and 1 year, a smaller size compared to other closely related sciaenids. While the species displays resilient reproductive characteristics such as protracted year-round spawning, constant supply of vitellogenic oocytes and relatively small/young age at maturity, there are also characteristics that make it susceptible to over-exploitation. These include highly female skewed sex ratios, as well as their schooling and voracious feeding behaviour leading to the potential for high catch rates.