Swimming out of Africa: mitochondrial DNA evidence for late Pliocene dispersal of a cichlid from Central Africa to the Levant

The rich Levantine fauna and flora were shaped by millions of years of migration across the region, from Africa to Eurasia and vice versa. Most large-scale processes that led to this diversity have been relatively well studied. However, small-scale processes, and details such as the area of origin of particular groups, and the route and time of dispersal are often not as clear. This is the case with the endemic Levantine representatives of the fish family Cichlidae. In this work we combine genetic, palaeontological and geological data in an attempt to understand the dispersal of the cichlid fish Astatotilapia flaviijosephi (Lortet, 1883) from sub-Saharan Africa to the Levant. A. flaviijosephi is unique among the Levantine cichlids in being the only non-tilapiine. It is also the only haplochromine cichlid to be found out of Africa. A partial sequence of the control region of the mitochondrial DNA was used to determine A. flaviijosephi 's phylogenetic relationships with other African haplochromines, and to estimate its time of divergence from this group. Combining our findings with palaeontological and geological data, we suggest that A. flaviijosephi separated from the other haplochromines during the middle to late Pliocene (2.5–3.3 Mya) and probably dispersed from Africa to the Levant via the Nile.  © 2004 The Linnean Society of London, Biological Journal of the Linnean Society , 2004, 82 , 103–109.     

Food and feeding habits of Mirogrex fingerlings in Lake Kinneret (Israel)

The food composition of the fingerlings (13–41 mm TL) of Mirogrex terraesanctae terraesanctae was studied on specimen collected at four stations in Lake Kinneret. The fingerling feed mostly on pelagic zooplankters and zoobenthic forms whereas algae were found in the intestines in low biomass. The number of zooplankton organisms in the guts of the fingerling that were sampled during June–August was higher by 234% than in fingerlings that were sampled during March–April. The role of sardines fry in the Kinneret ecosystem is discussed.     

Effect of water level fluctuation on shore spawning of Mirogrex terraesanctae (Steinitz), (Cyprinidae) in Lake Kinneret, Israel

Mirogrex terraesanctae (Steinitz), is an endemic, open water, zooplanktivorous fish. It spawns in the shallow littoral (0–50 cm) of Lake Kinneret, from November to May with a peak in mid winter (January-February). Spawning begins shortly after nightfall, when schools of the fish move along the shoreline in rocky regions, and release their milt and eggs. The adhesive eggs are attached to the surface of recently inundated, algae-free stones. A negative relationship exists between epilithic growth and egg density and survival. Algal growth may play a key role in the determination of the spawning depth. Breeding success of M. terraesanctae depends on the rate and extent of rise in lake level which, in turn, determines the availability of a suitable spawning substratum.     

Food composition of Clarias gariepinus (= C. lazera) (Cypriniformes,Clariidae) in Lake Kinneret (Israel)

The feeding behaviour of the freshwater piscivore, Clarias gariepinus (C. lazera) (C & V 1840) was studied over two periods: 1973–1975 and 1981–1982, in Lake Kinneret (Israel). The total number of fish analysed was 264 and their sizes (SL) and weights varied between 238 and 830 mm (146 to 5728 g). More than fifty species of plants and animals from the plankton, benthos and nekton of Lake Kinneret were identified in the intestines of C. gariepinus. Preyed fish were the most abundant food component (81%) and constituted the highest biomass, with Mirogrex terraesanctae representing the majority (although other species were also found). The potential impact of piscivory in the Kinneret ecosystem is considered.     

Feeding habits of larval Mirogrex terraesanctae (Steinitz, 1952) in Lake Kinneret (Israel) II. Experimental study

Interactions between the larvae of Mirogrex terraesanctae (Steinitz, 1952) in Lake Kinneret, Israel, and their zooplankton prey were studied experimentally. Prey species preference and size selectivities were measured. Larvae were hatched in the lab from eggs collected in the field, and fed different food items in various concentrations. The food items included lake zooplankton, algae, and commercial pellets. It was shown that small, first feeding larvae (7–8.5 mm SL) prefer small bodied zooplankters (< 180 µ). The effect of these food sources on larval growth was measured. It was found that larval Mirogrex grew at a higher rate when fed zooplankton prey sized from 63 µ–250 µ. Food items smaller than 63 µ, larger than 250 µ and Scenedesmus sp., produced less than optimal growth rates. The importance of Mirogrex feeding habits and their potential influence on the Kinneret ecosystem is considered.     

The impact of zooplankton status on the management of Lake Kinneret (Israel)

Monthly averages of standing stock wet biomass of zooplankton in Lake Kinneret (Israel) varied between 11 and 76 g m^–2 during 1969–1981, with the exception of two months. Averaged contributions of different groups were: Cladocera 58%, Copepoda 35% and Rotifera 7%. Total standing crop wet biomass is highest during January–June, averages varied between 35 and 50 g m^–2, and decreases during summer–fall (23–36 g m^–2). The winter biomass of Cladocera fluctuated between 22 and 35 g m^–2 and dropped to a range of 9–23 g m^–2 in summer, whereas copepod biomass varied very little around an average of 18 g (ww) m^–2 with the exception of low values from April to June. The stock biomass of Rotifera is relatively high during winter floods season (December-March) whilst in summer it is very low.Young stages of fish in Lake Kinneret feed mostly on zooplankton and zoobenthic forms. The most abundant fish in the Kinneret ecosystem, Mirogrex terraesanctae terraesanctae, also feed on zooplankton at the adult stage throughout the year, and herbivorous fish consume zooplankton during the summer when lake plankton resources are limited.The summer ecosystem of Lake Kinneret is characterised as a steady state type, in which the impact of the zooplankton-chain is of great importance. Increase of predation pressure on zooplankton by fish can disequilibrate the balanced trophic relations existing between nannoplankton production and zooplankton grazing capacity. Such a situation can lead to organics accumulation as nannoplankton blooms, resulting in water quality deterioration. Management options aimed at preventing collapse of zooplankton populations are discussed.     

Food composition of the barbelTor canis (Cyprinidae) and its role in the Lake Kinneret ecosystem

Synopsis Gut contents of 294 individuals ofTor canis from Lake Kinneret (Israel), sampled throughout a full year indicated a wide trophic spectrum including 52 identified animal and plant species. Nevertheless a major part of ingested biomass (5.7 g per fish) was due to the Kinneret sardine (Mirogrex sp.). The biomass of other ingested organisms was relatively low (0.01–0.8 g per fish) consisting of planktonic organisms, zoobenthos, macrophytes and a few others. The role ofT. canis in the Kinneret ecosystem and management design is discussed.     

The effect of environmental factors on the nesting and courtship behaviour ofTilapia zillii in Lake Kinneret (Israel)

Tilapia zillii is a polyphilic, guarding nest spawner which is widespread in west and north Africa and in the Middle East. A study of the nesting and courtship behaviour of this species in Lake Kinneret (Sea of Galilee), revealed that nesting and brood care differ according to the exposure of the nesting site to wave action and the nature of the substrate. The nests are simple and parental care is abbreviated in exposed, sandy sites whereas elaborate nests with brood chambers are excavated in sheltered areas with clayey substrates, and parental care is extensive. The implications of these findings to studies on cichlid breeding and the conservation ofT. zillii in Lake Kinneret are discussed.     

Aphanizomenon ovalisporum (Forti) in Lake Kinneret, Israel

The filamentous cyanobacterium Aphanizomenon ovalisporum wasobserved for the first time in Lake Kinneret in August 1994and formed a prominent bloom from September through October.Aphanizomenon ovalisporum reappeared in diminished amounts inthe summer and fall of 1995. These events are the first recordof significant quantities of a potentially toxic nitrogen-fixingcyanobacterium in this lake. No definite provenance of inoculumhas been identified, although A.ovalisporum was also observedin a newly reflooded area (Lake Agmon) in the catchment. Unusuallyhigh water temperatures and low wind inputs were observed priorto and during the A.ovalisporum bloom period. These, togetherwith possibly enhanced availability of phosphorus or other growthfactors, may have contributed to the cyanobacterium growth in1994. Phosphorus limi tation, as indicated by high cellularalkaline phosphatase activity, the onset of stormy conditionsand a fall in water temperatures led to the demise of the 1994bloom. Although the A. ovalisporum bloom in 1994 had no seriousdirect impact on water quality, the continued presence of apotentially toxic cyanobacterium in Lake Kinneret, a major nationalwater supply source, is a cause for serious concern.     

Sex ratio in Mesocyclops leuckarti (Claus) populations in Lake Kinneret (Israel)

Sex ratio (F/M) values of adult Mesocyclops leuckarti (Claus) populations in Lake Kinneret during 1970 and 1974 showed a periodic pattern of winter (January–March) and summer (August–October) female peaks. It is suggested that water temperature governs the appearance of both peaks but through different pathways.Contribution no. 163, Israel Oceanographic & Limnological Research Ltd.     

Two species of eriophyoid mites (Acari, Prostigmata) in wheat cultivation (Triticum aestivum L.) and associated grass community in Wielkopolska, Poland

Abstract: Quantitative characteristics of the occurrence of eriophyoid mites on wheat and associated grasses were studied. Three groups of hosts: cultivated wheat ( Triticum aestivum ), quackgrass ( Agropyron repens ) in cultivation, and quackgrass in balks were analysed. Two species of eriophyoid mites were recorded: Aceria tosichella Keifer and Abacarus hystrix (Nalepa). A. hystrix appeared more frequent [present in 65.0% of all collected samples; (95% confidence interval: 50–80)] and abundant [mean number of individuals per sample: 207.3 (94.4–351.5)] than A. tosichella (17.5%; (7.5–3.0), 13.8 (0.9–32.8), respectively). Significant differences in numbers among studied mite species populations on different host plants have been found. It is concluded that wheat is the marginal host for these two eriophyoids when compared to quackgrass.     

A new species of the deep-sea fish genus Paraliparis Collett (Liparididae) from the eastern North Atlantic, with notes on its ecology

A new species of liparidid, Paraliparis hystrix , is described from the slope waters (255–1140 m) to the west of Britain, in the area 49–59°N and 07–18°W. This pallid species is not uncommon, and is represented by a strong imbalance of females. Specimens close to ovulation have been collected in all seasons and have an estimated mean fecundity of 83 eggs.     

Seasonal Rotifer Dynamics in the Long-term (1969–2002) Record from Lake Kinneret (Israel)

Long-term (1969–2002) data record of biomass distribution of rotifers in Lake Kinneret is combined with previously published information on their metabolic activity and newly calculated population dynamics parameters to synthesize a model of their seasonal dynamics in Lake Kinneret. Nineteen rotifer species were recorded in routine samples collected in Lake Kinneret (Israel) in 7 offshore (deeper than 5 m), stations, at 12 discrete depths during 1969–2002. Organisms were sorted and counted (including external egg carrying females), biomass was measured and calculated for the entire lake stock (g_w.w m⁻²; mg l⁻¹). Rates of grazing, respiration and production were measured experimentally at three different temperature ranges. Results were extrapolated to the lake community for months with similar temperatures. Rotifera comprised 7% of total zooplankton biomass in Lake Kinneret whilst Cladocera and Copepoda 58 and 35% respectively. Rotifers were found to be more abundant during December–June and decline in summer months. Monthly (1969–2001) means indicated total grazing capacity of rotifers as 11%, respiration as 9% and production as 3.7% of the total zooplankton metabolic activity. Positive relations were indicated between rotifer and small bodied cladoceran numerical concentrations. Population growth models suggest that rotifers are not food limited in Lake Kinneret but that fish predation plays an important role in regulating abundance in spring-summer and fall.     

Sphaerospora epinepheli N. Sp. (Myxosporea: Sphaerosporidae) Observed in Grouper (Epinephelus malabaricus)

Sphaerospora epinepheli n. sp. is described from grouper, Epinephelus malabaricus , in cage-cultured and wild fish collected from both coastal lines of southern Thailand. Subspherical to spherical spores and mono- or disporous pseudoplasmodia were observed in the lumen of kidney tubules. Pseudoplasmodia were round to elongate, size range 15.6–22.9 μm (length) × 8.4–21.6 μm (width). Spores were 7.8–10.0 μm (length) × 12.3–14.5 μm (thickness), and 7.0–9.5 μm (width) with two spherical polar capsules of equal size measuring 2.9–4.4 μm in diameter and containing polar filaments with six or seven windings. Two uninucleate sporoplasms showed iodine vacuoles. Blood stages, similar to C-blood protozoans observed from freshwater fish in Europe, were found from peripheral blood smears of grouper. Ultrastructural studies of blood stages showed a similar structure to unidentified mobile protozoans from the blood of carp. Electron dense bodies were observed in the cytoplasm of the primary cell blood stages. Infected proximal-tubular epithelial cells showed highly vacuolated cytoplasm and pycnotic nuclei.     

Morphological Comparison of Diophrys scutum (Dujardin, 1841) and Diophrys peloetes n. sp. (Hypotrichida, Ciliophora)

SYNOPSIS. Diophrys scutum , collected from four locations on the New Hampshire coast, ranged from 89–195 7mu; in length, 50–105 μ in width, and 68–88 μ in buccal cavity length. The end of the adoral zone of membranelles (AZM) extends 37–59 μ (average = 45.6 μ) posteriorly in a groove on the right side of the body. Dorsally are five rows of stiff cilia. The silverline system (Chatton-Llvoff technique) appears as a fine meshwork, entirely different from that found in Euplotes or Uronychia . There are two elongate macronuclei (Feulgen reaction) and several micronuclei. Diophrys peloetes n. sp., collected from one location in Alligator Harbor. Florida, ranged from 95–134 μ in length, 62–84 μ in width. and 60–80 μ in buccal cavity length. The terminal portion of the AZM extends posteriorly in a groove 30–44 7mu; (average = 36.8 μ) on the right side of the body. Dorsally are eight rows of stiff cilia. The details of the silverline system are similar to those of D. scutum .
There is insufficient difference in ranges of body length, width. and buccal cavity length to use these characters in separating the two species. However, a statistical analysis shows that the length of the portion of the AZM on the right side of the body in D. scutum is significantly different (longer) from that of D. peloetes. Furthermore, these two species differ not only in number of dorsal ciliary rows, but also in the number of cilia per row. The degree of difference in these two species is similar to that between closely related species in other hypotrich genera, and also to that between some varieties of Paramecium aurelia.     

Identification of acetate-utilizing Bacteria and Archaea in methanogenic profundal sediments of Lake Kinneret (Israel) by stable isotope probing of rRNA

Acetate is an important intermediate in the decomposition of organic matter in anoxic freshwater sediments. Here, we identified distinct microorganisms active in its oxidation and transformation to methane in the anoxic methanogenic layers of Lake Kinneret (Israel) profundal sediment by rRNA-based stable isotope probing (RNA-SIP). After 18 days of incubation with amended [U-(13)C]acetate we found that archaeal 16S rRNA was (13)C-labelled to a far greater extent than bacterial rRNA. We identified acetoclastic methanogens related to Methanosaeta concilii as being most active in the degradation and assimilation of acetate. Oxidation of the acetate-methyl group played only a minor role, but nevertheless 'heavy'(13)C-labelled bacterial rRNA templates were identified. 'Heavy' bacteria were mainly affiliated with the Betaproteobacteria (mostly Rhodocyclales and Nitrosomonadales), the Nitrospira phylum (related to 'Magnetobacterium bavaricum' and Thermodesulfovibrio yellowstonii), and also with the candidate phylum 'Endomicrobia'. However, the mode of energy gain that allowed for the assimilation of (13)C-acetate by these bacterial groups remains unknown. It may have involved syntrophic oxidation of acetate, reduction of chlorinated compounds, reduction of humic substances, fermentation of organic compounds, or even predation of (13)C-labelled Methanosaeta spp. In summary, this SIP experiment shows that acetate carbon was predominantly consumed by acetoclastic methanogens in profundal Lake Kinneret sediment, while it was also utilized by a small and heterogeneous community of bacteria.     

Non-siliceous algae in a five meter core from Lake Kinneret (Israel)

The composition and succession of non-siliceous algae, studied in a five meter core from Lake Kinneret (Israel), are described. Only Chlorophyta species were recorded, probably due to the standard palynological sample processing which was used. In the lower part of the core, from the bottom to 300 cm (interval 5500–2500 years B.P.), Botryococcus braunii was the only common alga. Relevant changes in algal diversity and abundance occur at 300 cm. Many species of green algae were recorded for the first time (Pediastrum, Scenedesmus, Coelastrum, etc.). These changes may be related to an increase in nutrient concentration as a consequence of cultural disturbance. In the interval 300-0 cm, a succession of Pediastrum species is followed. The recovered green algae are extant in the present plankton of Lake Kinneret. They also constitute an important part of the algae found in the profundal sediments today.     

Decreased salinity effects in Lake Kinneret (Israel)

Kinneret is the only freshwater lake in Israel. It currently supplies about 30% of national water demands. Most pumped water is for drinking, and water quality is of major concern. During 1970–1987 temporal changes were observed in the lake ecosystem: decrease of salinity, decrease of total N (TN) and increase of total P (TP) mass contents, decline of TN/TP ratio, increase of phytoplankton biomass and increase of algal photosynthetic specific activity. It is suggested that because of decrease in salinity carbonic anhydrase activities in algal cells and nitrifying bacteria were enhanced. The increase of NO₃ flux through nitrification consequently enhanced denitrification and nitrogen losses in lake waters. These increased N losses together with P increase, as reflected by the decline of TN/TP ratio might be a slight shift from the present both P and N deficiencies to a higher level of N limitation in the Kinneret ecosystem. This This may cause changes in phytoplankton community structure possibly without changing primary production levels but deteriorate water quality.     

Morphological evidence for the occurrence of two electric catfish (Malapterurus) species in the White Nile and Omo–Turkana systems (East Africa)

Morphological analysis of 28 Malapterurus specimens collected from the Nile and Omo–Turkana basins revealed two groups. The first was characterized by low vertebral counts (41–44) and numerous gill rakers (14–22) situated along the entire first ceratobranchial, the second group was characterized by high vertebral counts (47–51) and a low number of gill rakers (6–13) situated only on the upper portion of the first ceratobranchial. Through comparison with the type specimens of three currently recognized Malapterurus species and comparison with museum collections from the Nile, Omo–Turkana, and Chad basins, these two groups were identified as M. electricus and M. minjiriya . A principal component analysis of 27 measurements allowed partial discrimination among species. Biogeographic aspects of finding M. minjiriya in the tributaries of the White Nile and in the middle reaches of the Omo River are discussed.