Genetic diversity of two Southern African cichlids (Oreochromis andersonii and O. macrochir) in the Zambezi and Congo River basins

Oreochromis andersonii and O. macrochir are two important cichlid species native to Southern Africa. We describe in this paper their genetic population structure in the Upper Zambezi River, Kafue River, and Lake Bangweulu representing part of the Congo and Zambezi River Basins. Microsatellite genetic markers were employed to analyse the genetic population structure of the two species using 177 tissue samples. The average allele richness of O. andersonii was higher in the Zambezi River (10.500) than in the Kafue River (9.583) though not statistically different. For O. macrochir, it was highest in the Zambezi River (11.170) followed by the Kafue River (9.781) and least in Lake Bangweulu (7.067) and their differences were significant. The gene diversity indices; gene diversity (hs), observed heterozygosity (H_O), expected heterozygosity (H_E) were high and similar in O. andersonii populations. However, in O. macrochir H_E was significantly lower in Lake Bangweulu (0.678) compared to the Kafue River (0.799) and Zambezi River (0.802) populations. Population differentiation estimated by R_ST and D_EST revealed high differentiation in both species (R_ST = 0.598, D_EST = 0.777 for O. andersonii; R_ST = 0.379, D_EST = 0.710 for O. macrochir). The highest source of variation was among populations (84.71%) for O. andersonii and within populations (67.09%) for O. macrochir. Comparisons of population pairs revealed a close genetic similarity between the Zambezi River and Lake Bangweulu populations of O. macrochir. Bottlenecks were observed in both species using the Two-Phase Model (T.P.M.) indicative of a recent genetic loss or reduction in effective population size. Though our results indicate that the populations of both species still maintain sufficiently high levels of genetic diversity in the sampled areas, the bottlenecks observed are a source of concern. We recommend a more robust study of genetic diversity of these species in all sections of these river systems and that some key conservation sites should be identified to protect the gene pool of these native species.     

The growth ofOreochromis andersonii (Pisces: Cichlidae) from the Okavango Delta,Botswana, and a comparison of the scale and otolith methods of ageing

Synopsis Otoliths and scales were used for age and growth determination ofOreochromis andersonii from the Okavango Delta, Botswana. Marginal increment analysis showed that an annulus was formed in both the scales and otoliths during the dry summer period. Using scales, the growth ofO. andersonii was described by L_t = 285.27(1-e^{-0.26(t+2.02)}) mm SL and using otoliths by the equation L_t = 267.48(1-e^{-0.25(t+2.18)}) mm SL. Maximum age estimates of 10 years using scales and 13 years using otoliths were obtained and the growth curves were significantly different (p < 0.01). Age estimation using scales tended to over-emphasise growth inO. andersonii resulting in larger predicted lengths-at-age. For this reason, otoliths are considered to be more reliable and suitable than scales in determining the age and growth of this species.     

Postprandial changes in pH and enzyme activity from the stomach and intestines of Tilapia rendalli (Boulenger, 1897), Oreochromis mossambicus (Peters, 1852) and Clarias gariepinus (Burchell, 1822)

The ability to utilise dietary components differs among fish species. Digestive enzymes may be used to determine the efficiency of the digestive process. In this study, the activities of the digestive enzymes in Tilapia rendalli, Oreochromis mossambicus and Clarias gariepinus were explored. Protease, amylase, lipase and cellulase activities were measured in different parts of the digestive tract of the three fish species. The pH dynamics along the digestive tract were monitored. In all fish species, the presence of food led to a reduction in stomach pH, whereby pH values of 1.54, 1.58 and 2.01 were recorded 12 h after feeding in O. mossambicus, T. rendalli and C. gariepinus, respectively. Protease and amylase activities were significantly higher (P < 0.05, anova ) in the tilapias than in C. gariepinus. The tilapias may be pre‐adapted to produce more protease and amylase to digest plant material, which is more difficult to digest than animal matter. In all species amylase activity was significantly higher in the proximal intestine than in the other parts of the digestive tract (P < 0.05, anova ). The highest protease activity was recorded in the distal intestines. This is because of the alkaline pH recorded in the proximal and distal intestines, which favours amylase and protease activity, respectively. Lipase activities were significantly higher (P > 0.05) in C. gariepinus than in both tilapias. Marginal cellulase activities were recorded in all species. It is inferred here that phylogeny and not diet may be the main factor influencing enzyme activities, as all fish were fed a similar diet.     

Validation of annulus formation in otoliths of largemouth bass Micropterus salmoides outside their native range

The periodicity of growth zone formation was validated for largemouth bass Micropterus salmoides using edge analysis (EA) and mark recapture of chemically‐tagged wild fish (MRCT) to test the hypothesis that one opaque and hyaline zone was deposited annually in sagittal otoliths sampled from temperate South African M. salmoides populations. For 35 fish recaptured in the MRCT experiment, the relationship between the number of growth zones posterior to the chemical mark and the time at liberty (0.04–1.38 years) did not differ significantly from a 1 : 1 relationship (t‐test, t = 0.76, d.f. = 2,33, P = 0.45). This result was supported by EA, where periodic logistic regression and a binomial model linked with a von Mises distribution for circular data demonstrated that the frequency of otoliths with opaque margins followed a unimodal distribution (maximum October–January). Both the timing of growth zone deposition (spring) and the annual rate were consistent with results from validation studies conducted globally in localities ranging from 45°N to 33°S, and indicate that the growth zone deposition rate is annual throughout the native and introduced range of this species.     

Breeding pattern of Oreochromis niloticus (Linnaeus, 1758) versus native congeneric species,Oreochromis macrochir (Boulinger, 1912), in the upper Kabompo River,northwest of Zambia

Investigating the determinants of the reproductive biology of fishes is an essential component of fisheries research. Tilapia breeding patterns were investigated to determine the impact of non‐native Oreochromis niloticus on the native congeneric Oreochromis macrochir in the upper Kabompo River in the Northwest of Zambia using the gonadosomatic index and the sex ratios. Oreochromis niloticus was the most abundant fish caught (221, 63.5%) than O. macrochir (127, 36.5%). Results showed that the overall gonadosomatic index means of O. macrochir in both sections were similar. Oreochromis macrochir bred in December and February–March, with no reproduction in June. However, O. niloticus in the invaded section indicated all year reproduction through reduced spawning in May–June, with increased spawning activity in February–March. The sex ratio (females: males) was 1:1.3 and 1:1.7 for O. niloticus and O. macrochir, respectively, and both significantly deviated from the sex ratio of 1:1 (ꭓ² = 8.42 and 9.37, p < .05). Our study has revealed that O. niloticus was able to spawn across all sampled months with a 23% higher breeding population than O. macrochir, which might explain the suppression in the abundance of native O. macrochir. Due to the superior breeding patterns of O. niloticus, fisheries, wildlife, and aquaculture practitioners need to make contingency plans to alleviate its impacts further downstream of the Kabompo River.     

Validation of annual periodicity in otoliths of red snapper, <Emphasis Type="Italic">Lutjanus campechanus</Emphasis>

The periodicity of otolith growth increments (opaque and translucent zones) from adult red snapper (Lutjanus campechanus) was examined through a mark and recapture study (2005–2010), and laboratory holding of hatchery reared red snapper over a 2 year period (2002–2004). Wild red snapper (n = 295) were caught hook-and-line, marked with anchor tags, injected with oxytetracycline dihydrate (OTC), and released in the Gulf of Mexico 15–40 km south of Dauphin Island, Alabama. Marked fish were recaptured up to 2.8 years after release (n = 35) and sagittal otoliths were dissected, sectioned and examined under white and blue-violet light. The number of opaque growth zones past the OTC mark was compared to time at liberty for each fish and supported an annual periodicity of growth increment formation. Also, most (87%) of the hatchery reared fish showed two opaque zones that supported an annual increment formation rate. However, an unusual timing of opaque zone formation was shown for mark-recaptured fish. Based on known timing of OTC marking, otoliths from mark-recapture fish showed opaque zone formation from late summer (August) to early winter (December). This fall formation of opaque zones is in contrast to previous studies and its timing may relate to the end of spawning for this species.     

Parasites of fishes in the recently inundated ephemeral Lake Liambezi,Namibia

Lake Liambezi forms the periodic connection between the upper Zambezi, Kwando and Okavango rivers. A full parasitological assessment was conducted on 86 fish, representing 14 species in six families sampled in August 2011. Parasite diversity was low and dominated by species with complex life cycles involving intermediate hosts. Most prevalent were larval nematodes (Contracaecum sp.) infecting 12 and Trypanasoma sp. infecting nine of the 14 host species. The intra-erythrocytic parasite Babesiosoma mariae was found in the blood of Coptodon rendalli and Oreochromis andersonii with prevalence of 50% and 60%, respectively. The host-specific monogenean Annulotrema hepseti was recorded only from H. cuvieri with a prevalence of 100%. Notable absences were the copepod and branchiuran parasites that have direct lifecycles and usually occur in high prevalence and abundance in the region. Because parasites with direct life cycles can only be transported into the lake on the host fish, their absence suggests limited immigration of infected fishes into the lake. This suggests that internal recruitment dominates over immigration in the fish population dynamics in Lake Liambezi.     

Age and growth of <Emphasis Type="Italic">Schizothorax waltoni</Emphasis> in the Yarlung Tsangpo River in Tibet,China

Age and growth were studied for Schizothorax waltoni in the middle reaches of the Yarlung Tsangpo River in Tibet, southwest of China, from April 2004 to September 2006. A total of 201 specimens were collected ranging from 110 to 580 mm in standard length (SL). In contrast to other otoliths, sectioned lapillus showed a clear pattern of alternating opaque and hyaline zones. Marginal increment analyses showed that the increments, each composed of one opaque and one hyaline zone, are deposited annually. Opaque edges were prevalent from May to August. The von Bertalanffy growth parameters based on sectioned otolith data were L _t  = 689.8{1 − exp[0.051 (t + 3.275)]} for males, and L _t  = 691.1{1 − exp[0.056 (t + 2.466)]} for females. The slow growth and long life indicate that S. waltoni is vulnerable to overfishing and that harvesting strategies for the species should be conservative.     

Age and growth of longnose trevally (Carangoides chrysophrys) in the Arabian Sea

The ages and growth of longnose trevally (Carangoides chrysophrys), caught in the northwest Arabian Sea between April 2005 and September 2006, were investigated. Age and growth of 336 fish specimens were determined using sectioned sagittae otoliths. Annual opaque growth rings were formed between December and March, with the majority being laid down in February and March, coinciding with the spawning period and high water temperatures. Marginal zone or edge analysis was used to validate the annual deposition of the opaque zone in the otoliths. This species showed large variations in length‐at‐age, suggesting large growth variations among individuals of the same cohort. The estimated von Bertalanffy growth model differed significantly between the sexes, with males having larger mean lengths at age and reaching a larger asymptotic size. The von Bertalanffy growth models were TL (cm) = 73.34[1‐exp (?0.25 (t + 1.21))] and TL(cm) = 73.26[1‐exp (?0.24 (t + 1.20))] for males and females, respectively.     

Aspects of the Life History of the Salema, Sarpa salpa (Pisces, Sparidae), off the Canarian Archipelago (Central-East Atlantic)

Life history of the salema, Sarpa salpa, off the Canarian archipelago was studied from samples collected between January 1998 and December 1999. Fish ranged from 119 to 452 mm in total length and from 24.9 to 1491.7 g in weight. The species was characterized by a protandric hermaphroditism. The overall sex ratio was unbalanced in favour of males (1 : 0.41). The reproductive season extends from September to March, with a peak in spawning activity in December–January. Males reached maturity at a smaller length (226 mm, 2 years old) than females (294 mm, 3 years old). Males, females and all fish showed an isometric growth. The morphometric relationship between length and mass for the whole population was described by the parameters: a = 0.0000134 and b = 3.01. A concentric pattern of opaque and translucent zones was readily distinguishable on the otoliths. Two rings, one opaque and one translucent, were laid down each year on the otoliths; the opaque zone was formed during the summer months, and the translucent zone during the winter months. Individuals aged 0 to 11 years were found. The parameters of the von Bertalanffy growth equation for all individuals were: L = 479 mm, k = 0.212 year^–1, and t₀ = –1.08 year.     

Comparing the fish assemblages and food‐web structures of large floodplain rivers

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FIRST RECORD OF OREOCHROMIS MACROCHIR (BOULENGER, 1912) (PISCES:CICHLIDAE) FROM THE LIMPOPO RIVER IN SOUTHERN AFRICA.

Summary

During 1973/74, Barbus poechii, Oreochromis macrochir, O. andersonii, Serranochromis angusticeps, S. codringtonii, S. robustus and S. thumbergi from the Okavango System were introduced into the Shashe Dam (Limpopo System, Botswana). These species are alien to the Limpopo System. A fish survey during September 1991 disclosed the presence of two specimens of the greenhead tilapia O. macrochir in a large natural pool in the Limpopo River 7 tan downstream from the Shashe/Limpopo confluence. In August 1992 another specimen was caught in this pool. It is surmised that the likely origin of these specimens is the Shashe Dam. There is a possibility that the greenhead tilapia may interbreed with the indigenous Mocambique tilapia O. mossambicus. Consequently the genetic integrity of O. mossambicus in the Limpopo River may become suspect. If the other alien species introduced into the Shashe Dam reach the Limpopo River, O. andersonii may also interbreed with O. mossambicus while the Serranochromis spp. could be detrimental for feeding relationships in the Limpopo River and its tributaries. The impact that B. poechii could have is uncertain but it does not appear to co-exist with the closely related B. trimaculatus of the Limpopo System. Eradication of O. macrochir in the Limpopo River is not practical. Further stocking of the Shashe Dam as well as other impoundments in the Limpopo System with alien fish species should be discouraged.     

Trophic interrelationships between the exotic Nile tilapia, <Emphasis Type="Italic">Oreochromis niloticus</Emphasis> and indigenous tilapiine cichlids in a subtropical African river system (Limpopo River,South Africa)

The stable isotope ratio and seasonal changes in diet of two indigenous (Oreochromis mossambicus, Tilapia rendalli) and one exotic (Oreochromis niloticus) tilapiine cichlids in the subtropical Limpopo River, South Africa were investigated to determine patterns of resource partitioning. Stomach contents of O. niloticus and O. mossambicus indicated high dietary overlap across size class, habitat and season, with both species primarily feeding on vegetative detritus. However, stable isotope analysis revealed that the two Oreochromis species had different stable isotope ratios derived from different food sources. The relatively δ¹³C-depleted O. niloticus indicates a phytoplankton-based diet, while the δ¹³C-enriched O. mossambicus indicates a macrophagous diet dominated by vegetative detritus and periphyton. The high similarity in stomach contents and the interspecific differences in isotopic composition reveal fine-scale patterns of food resource partitioning that could be achieved through selective feeding. Tilapia rendalli was largely macrophagous and fed mainly on aquatic macrophytes and had a low dietary overlap with both O. niloticus and O. mossambicus. In the Limpopo River, detritus and algae are probably the most abundant food resources and the causal factors responsible for the observed patterns of resource partitioning among the tilapiines are usually difficult to ascertain. Fish may be able to perceive food resources in terms of the dynamics that determine their availability. Detailed studies of variation in food resource availability and fish habitat use within the system are needed to evaluate this hypothesis.     

Validation of annulus formation in otoliths of a temperate population of adult African sharptooth catfish Clarias gariepinus using fluorochrome marking of wild fish

The deposition rate of growth zones in sagittal otoliths of wild Clarias gariepinus that were fluorochrome marked with oxytetracycline and recaptured after a period of between 0·44 and 2·13 years was consistent with the deposition of one growth increment annually.     

Using otoliths to estimate age and growth of a large Australian endemic monocanthid, <Emphasis Type="Italic">Nelusetta ayraudi</Emphasis> (Quoy and Gaimard, 1824)

Nelusetta ayraudi (the ocean leatherjacket) is an endemic Australian monacanthid species distributed from North West Cape (Western Australia) south to southern Queensland. The commercial and recreational fisheries targeting Nelusetta ayraudi have expanded substantially along the coast of New South Wales (NSW) in recent years but there exists little biological information on which to base effective management of this growing fishery. World-wide, only a few studies have aged monacanthids. Of these, researchers have interpreted periodic increments in bony structures such as vertebrae and anterior dorsal spines in preference to those found in otoliths. In this study we estimated age of N. ayraudi by counting growth increments in sectioned otoliths. The periodicity of increment formation was validated using a vital stain, (oxy-tetracycline), injected into young-of-the-year fish. Growth was rapid especially as juveniles with N. ayraudi attaining approximately 220 mm after 1 year and 340 mm after 2 years. No differences in growth rates were detected between sexes or between fish captured at different latitudes (zones). The largest male (605 mm, Total Length—TL) and female (656 mm, TL) were both recorded from northern NSW, with both sexes attaining the maximum age of 6+ years from northern and southern NSW. The von Bertalanffy parameters describing growth for N. ayraudi were L_￥ {L_\infty }  = 591 mm (TL), k = 0.377 year⁻¹ and t _o = −0.247 years.     

Age determination,growth and mortality of Gerres subfasciatus Cuvier, 1830 in southeast Australia

Commercial beach‐seine (30–50 mm cod‐end mesh) catches of Gerres subfasciatus in Lake Macquarie southeast Australia were sampled monthly between December 2000 and April 2002 to investigate species age, growth and mortality characteristics using sectioned sagittal otoliths. Inclusion of fishery‐independent length data of young‐of‐the year collected from littoral seagrass beds with fine meshed beach seines augmented growth analyses. Fifty otoliths each month (total 448 female and 402 male) were examined with sections displaying alternating opaque and translucent zonation. Formation of otolith opaque zones occurred annually between August and December as determined by monthly marginal increment analyses. Spawning occurred between October and February, and a universal birth date of 1 December was assigned that allowed counts of opaque zones to be converted to age estimates in years and months (decimal ages). The von Bertalanffy growth function identified that both genders grew fast for their initial 2 years, but thereafter slowed. Females grew faster, had a greater mean fork length‐at‐age, estimated L_∞ and an observed greater maximum fork length in samples compared to males. Nevertheless, observed longevity was 10⁺ years for both genders. Commercial catches of both genders were dominated by the 2⁺ and 3⁺ year age classes. Catch curve analyses identified the instantaneous rate of total annual mortality to be ~0.8 and an exploitation rate of <0.5 for both genders. The data form a basis for developing assessment and management strategies of G. subfasciatus fisheries.     

Otolith age determination for adult tilapia, Oreochromis niloticus L. from Lake Awassa (Ethiopian Rift Valley) by interpreting biannuli and differentiating biannual recruitment

Age of mature Oreochromis niloticus in Lake Awassa, Ethiopia, was estimated by analysing optical macrozones (translucent and opaque) in sagittal otoliths from fish sampled over a 12-month period. Seasonal record on the type of macrozone at the edge of otoliths suggested that two translucent macrozones associated with biannuli were formed each year; one during January and February and another during June and July. Formation of translucent macrozones coincided with minimum water temperature, spawning associated loss in condition and presumably with reduction in the quantity and quality of the food consumed by the fish. Relative marginal increment analysis showed that biannulus formation may be completed in March and in August each year. A concurrent study has confirmed a biannual recruitment in O. niloticus in Lake Awassa, from which mid-February and mid-August were taken as median hatch-dates. A procedure to assign otolith age is described which considers median hatch-dates and uses the number of biannuli in otoliths without discriminating between fish from the two recruitment cohorts. We also show a procedure to discriminate between the two cohorts by using the number of biannuli, conditions on the edge of the otolith and time of capture to assign age for the two recruitment cohorts separately.     

Trophic interrelationships amongst cichlid fishes in a tropical African reservoir (Lake Chivero,Zimbabwe)

The diet of seven cichlid species of Lake Chivero, Zimbabwe, was investigated from October 2004 to April 2005 to determine the degree of dietary overlap and interspecific competition. These fish could be separated into four groups. Two microphages, Oreochromis niloticus and O. macrochir, fed mostly on blue-green algae and their diets, in all size classes, overlapped almost completely. This suggested a high degree of competition between the two species and may account for the fact that O. niloticus, introduced into the lake in the 1980s is now the dominant species. There was also a significant dietary overlap between the two macrophages Tilapia sparrmanii and T. rendalli, which were macrophages with macrophytes as a significant component of their diet. Interspecific competition between these two species may account for the increase in the abundance of T. sparrmanii and a decrease in the numbers of T. rendalli over the last two decades. The diet of two small omnivores Pharyngochromis acuticeps and Pseudocrenilabrus philander consisted of algae, zooplankton, plant material, fish and detritus and also overlapped extensively, but these species probably avoid competition by living in different habitats; P. philander is usually associated with vegetation while P. acuticeps tends to live in more open waters. The last species, Serranochromis robustus, was largely piscivorous. There were no ontogenetic variations in diet, apart from P. acuticeps where small fish fed mainly on zooplankton becoming omnivorous in larger ones, and S. robustus where the diet shifted from zooplankton in specimens <50 mm SL to fish in larger size classes. Data available from the 1960s and 1970s indicate that the diets of some species have changed with invertebrates such as chironomid larvae being previously more important in all diets, while most species now take a higher proportion of algae. This reflects changes in the lake, which is eutrophic and has experienced severe algal blooms since 1960. Handling editor: C. Strumbauer     

Ultrastructure of otolith increments and checks in the teleost fish Oreochromis niloticus

An otolith increment is composed of an incremental zone and a discontinous zone. Observation of the otoliths of Oreochromis niloticus with transmission and scanning electron microscopy reveals that organic fibers are concentrated in the discontinuous zone and are relatively scarce in the incremental zone. On the other hand, calcium carbonate crystals are chiefly packed in the incremental zone and are less dense in the discontinuous zone. Both fibers and crystals are oriented perpendicular to the growth increments. Otolith checks, or discontinuities, contain even denser fibers and fewer fine crystals than does the discontinuous zone. A higher proportion of individual fibers is prominently stained in the discontinuous zone and check than in the incremental zone. Other features of individual fibers appear to be the same among the three zones. The crystals on either side of a check or a discontinuous zone resemble each other both in size and orientation. Zonation of protein and mineral components of otoliths can be understood in terms of both incremental growth and the daily cycle of deposition.     

First record of anomalous otoliths of Menticirrhus americanus in the South Atlantic

Sagitta otoliths are usually formed of calcium carbonate polymorphs as aragonite. The objective of this study was to verify which carbonate polymorph is predominant in the sagitta otolith of Menticirrhus americanus and check whether this pattern remains in otoliths with morphological alterations. Otoliths of M. americanus were obtained from five sites on the southeast‐south coast of Brazil (São Sebastião (SS) 23°45′S–45°24′O, n = 29; Cananéia‐Iguape Estuarine Complex (CI) 25°02′S–47°54′O, n = 30; Paranaguá Estuarine Complex (PEC) 25°28′S–48°20′O, n = 35; Itapoá (IT) 26°07′S–48°36′O, n = 31; Laguna (LA) 28°28′S–48°46′O, n = 13). The characterization of carbonate polymorphs of otoliths was performed through Raman spectroscopy, a photonic and non‐destructive technique that analyzes molecular vibrations induced by laser. We analyzed 138 pairs of M. americanus otoliths, of which eight otoliths from different pairs presented morphological alterations (SS n = 1, CEP n = 5, IT n = 1, LA n = 1). The Raman spectra show that normal otoliths, that is, without morphological alterations, presented only aragonite in their structure. Among the otoliths that presented morphological alterations, the Raman spectra allowed to identify in six otoliths the deposition of aragonite and in only two otoliths the deposition of vaterite (one specimen of the PEC and one of SS).