Paternal mouthbrooding bagrid catfishes in Lake Tanganyika

 This article gives the first information about paternal mouthbrooding in bagrid catfishes, based on analysis of specimens collected while SCUBA diving in Lake Tanganyika, Africa. Lophiobagrus aquilus and L. cyclurus, endemic to the lake, were nocturnally active, and in the daytime were found concealed singly or in pairs beneath rocks. Among 39 L. aquilus and 52 L. cyclurus collected, 8 and 1 solitary males, respectively, were incubating eggs or young in the buccal cavity. Paired males and females and solitary females never took care of the offspring. In both species, females of pairs showed much higher gonadosomatic index values and had larger oocytes in the ovaries than did solitary females. This fact suggests that pair formation takes place at a time near the gonadal maturation of the female and that the pair separate after spawning. We tentatively propose an evolutionary transition route of care patterns in bagrid catfishes. Received: January 25, 2002 / Revised: April 13, 2002 / Accepted: April 17, 2002     

A taxonomic review ofXenotilapia sima andX. boulengeri (Cichlidae; Perciformes) from Lake Tanganyika

Two cichlid fishes from Lake Tanganyika,Xenotilapia sima Boulenger, 1899 andX. boulengeri (Poll, 1942) are often confused owing to insufficient information on the diagnostic morphological characters for each species. Morphological and morphometric analyses showed thatX. sima differs fromX. boulengeri in having a greater number of teeth and anal and pectoral fin soft rays, thicker caudal peduncle, larger eyes, and a shorter snout and anal fin base.     

Systematics of Tanganyikan cichlid fishes (Teleostei: Perciformes)

The relationships among 53 genera of Tanganyikan cichlid fishes were analyzed based on internal and external morphological features. Comparison of the morphological cladistic tree with a previously proposed classification showed 5 of 12 tribes to be nonmonophyletic. Sixteen tribes were recognized, the changes in classification being that Trematocarini was treated as a junior synonym of Bathybatini; 5 new tribes were established for each of the following genera, Benthochromis, Boulengerochromis, Ctenochromis benthicola, Cyphotilapia, and Greenwoodochromis; Ctenochromis horei was transferred from Haplochromini to Tropheini; and Gnathochromis pfefferi was transferred from Limnochromini to Tropheini. The revised classification was supported by previously proposed molecular trees.     

Taxonomic status of the six-band morph of Cyphotilapia frontosa (Perciformes: Cichlidae) from Lake Tanganyika, Africa

Six- and seven-band morphs have been identified in a cichlid, Cyphotilapia frontosa, that is endemic to Lake Tanganyika. These color morphs have allopatric distributions; the six-band morph is widespread in the northern half of the lake while the seven-band morph is restricted to Kigoma on the east coast of the lake. Because no specimens of the seven-band morph have been available for taxonomic study except for the holotype of C. frontosa, the taxonomic status of these morphs has not been discussed. In a recent survey at the lake, 21 specimens of the seven-band morph were collected. A comparison of these with existing collection specimens of the six-band morph showed significant differences in morphometric and meristic characters; however, because all characters largely overlapped between these morphs, they are regarded as conspecific.     

Evolution of the tribe Tropheini from Lake Tanganyika: synchronized explosive speciation producing multiple evolutionary parallelism

One of the most surprising outcomes of recent molecular studies on cichlid fishes of the three Great East African Lakes Victoria, Malawi and Tanganyika, was the stunning rapidity of speciation and cladogenesis at early stages of adaptive radiation. Despite their rapid pace, speciation events were so far intuitively assumed to proceed in a bifurcating and tree-like fashion, even if they could not be resolved by gene phylogenies due to a lack of resolution. On the basis of phylogenetic analyses of the Tropheini, a lineage of endemic rock-dwelling cichlid fishes from Lake Tanganyika, we suggest a pathway of explosive speciation that accounts for a non-bifurcating manner of cladogenesis. This pattern is likely to be the result of the contemporaneous origin of a multitude of founder populations in geographically isolated rock habitats among which gene flow was interrupted simultaneously by a major change of the lake habitat in the form of a rapid rise of the lake level. As a consequence, all new species arising from that vicariance event must exhibit almost equal genetic distances to each other, within the scope of genetic diversity of the founder population(s), even if the actual processes of subsequent speciation and eco-morphological diversification followed independent routes. Our phylogeny also suggests a high frequency of parallel evolution of equivalent trophic specialization in the Tropheini. This phenomenon seems to be an inherent feature of this pathway of speciation, due to the action of similar selective forces on the same set of species colonizing isolated habitats of the same type. Explosive speciation via synchronization of genetic divergence triggered by rapid environmental changes seems to be particularly likely to occur at advanced stages of adaptive radiation, when species are already adapted to particular habitats and have a reduced ability for dispersal.     

Phylogenetic analysis of Cyprichromini (Perciformes: Cichlidae) endemic to Lake Tanganyika and validation of the genus <Emphasis Type="Italic">Paracyprichromis</Emphasis>

The phylogenetic relationships among two Paracyprichromis and five Cyprichromis species, included in the Tanganyikan cichlid tribe Cyprichromini, were investigated using morphological features. The previously proposed diagnostic characters of Paracyprichromis are not synapomorphies, because the nonelongated swim bladder is plesiomorphic, the numbers of dorsal and anal fin rays and scales on longitudinal line and around the caudal peduncle overlap with those of Cyprichromis, and these counts and number of vertebrae are all included within the ranges of other Tanganyikan cichlids. The monophyly of Paracyprichromis is supported by a unique condition of infraorbitals to this genus. Additionally, the monophyly of Cyprichromis was reconfirmed by one of the previously proposed diagnostic characters, the presence of an elongated swim bladder.     

Description of a new Lake Tanganyikan cichlid fish of the genus Cyprichromis(Perciformes: Cichlidae) with a note on sexual dimorphism

Cyprichromis coloratus , a new cichlid species, is described based on 41 type specimens from the Zambian coast of Lake Tanganyika. It differs from Cyprichromis microlepidotus by having fewer scales in the longitudinal line (41–44 v . 59–70 in C. microlepidotus ) and upper lateral line (31–36 v . 47–59), from Cyprichromis leptosoma by more dorsal‐fin spines (14–15 v . 11–13 in C. leptosoma ) and longitudinal line scales (41–44 v . 38–40), from Cyprichromis pavo by fewer dorsal‐fin spines and soft rays (29–31, usually 30, in total v . 30–32, usually 31, in C. pavo ), and from Cyprichromis zonatus by shallower body depth [24·0–28·5 (26·4 ± 1·1)% of standard length v . 24·5–30·5 (28·4 ± 1·3)% in C. zonatus ] and smaller eye [eye length 25·2–30·9 (27·2 ± 1·1)% of head length v . 27·9–33·3 (30·2 ± 1·2)%]. The new species is also separable from C. microlepidotus , C. pavo and C. zonatus by absence of distinct small spots on the body ( v . pearl grey and yellowish small spots along scale rows on body at least in males of C. microlepidotus and C. pavo ), its distinct dichromatism (particularly on the caudal fin) between males of the same population ( v . dichromatism absent in C. zonatus ) and absence of vertical bands ( v . three or four distinct vertical bands beneath dorsal fin base in live males of C. zonatus ). Cyprichromis coloratus n. sp. also exhibits sexual dimorphism, females having a larger head and males having longer pelvic fin, differences which are discussed in relation to reproductive behaviour. A key to the five species of Cyprichromis is included. A lectotype is designated for Cyprichromis leptosoma .     

Diatoms from surface sediments of the northern part of Lake Tanganyika

227 Diatom taxa were observed in the surface sediments of the northern part of Lake Tanganyika, including 1 new to science: Amphora tanganyikae. The diatom community of these sediments is mainly composed of benthic organisms while planktonic diatoms are rather rare. Many brackish-water and a few marine organisms were observed. Cosmopolitan organisms (77.1%) dominate the diatom flora but tropical, tropical African and African taxa are also well represented (22.9%)Deceased.Deceased.     

Validation of the periodicity of increment formation in the otoliths of a cichlid fish from Lake Tanganyika, East Africa

Tetracycline was used as a chemical tag in a mark‐recapture study to examine the pattern of increment formation in the otoliths of Tropheus moorii , a rock‐dwelling cichlid from Lake Tanganyika. A total of 256 fish were captured by divers and injected with tetracycline. Of these, nine were recaptured after either 1 or 2 years at liberty and eight retained tags within their otoliths. Comparison of the number of growth increments formed after the tag and the time at liberty demonstrated that increments were deposited on an annual basis in the otoliths of this species. Furthermore, there was a strong relationship between otolith mass and age suggesting that otoliths grew at a predictable rate throughout the life of the fish. Validation of an annual pattern of increment deposition allowed age and growth information to be derived from otoliths. This showed that T. moorii grew rapidly to attain adult size by 3 years of age. Males grew faster than females and also attained a larger size than females (8·74 v . 7·91 cm L _S respectively). The longevity of some of these small freshwater fish was surprising; the oldest individual had an age of 10 years, while the average age of adults was 4 years.     

Lunar synchronization of spawning in cichlid fishes of the tribe Lamprologini in Lake Tanganyika

Lunar synchronization of spawning was investigated for eight substrate brooding cichlid fishes belonging to the tribe Lamprologini. Their spawning activities all peaked during the second quarter of the lunar cycle. Comparison between their breeding styles and the degrees of spawning synchronization suggested that the nocturnal guarding-efficiency of eggs, especially exposed ones, is improved by the maximal amount of moonlight during full moon and, in species whose young leave the breeding sites about 2 weeks after spawning, the survival of dispersing young is enhanced by the darkness of moonless nights.     

Seasonal patterns of sediment loading and benthic invertebrate community dynamics in Lake Tanganyika, Africa

1. This study investigated the spatiotemporal dynamics of benthic invertebrate communities located close to the mouths of the Kalambo and Lunzua Rivers, at the southern end of Lake Tanganyika, and subject to varying degrees of sediment load. 2. Metrics of exposure to riverine sediments were associated inversely with abundance and diversity of benthic organisms at sampling locations adjacent to both river mouths, indicating a major effect of riverine sediments on the structure of near‐shore benthic communities. 3. A lower abundance of benthos was found at the mouth of the Lunzua River, which exports significantly higher sediment loads than the Kalambo. 4. Seasonal cycles of abundance and diversity in the benthos varied with distance from river mouths. This has important implications for monitoring programmes investigating benthic communities close to the mouths of rivers. 5. Our findings suggest that recent increases in the sediment loading of Lake Tanganyika, owing to anthropogenic catchment disturbance, impact significantly on biological diversity throughout the lake littoral, which provides habitat for the majority of species in the lake.     

Ecology and conservation status of endemic freshwater crabs in Lake Tanganyika, Africa

Sedimentation resulting from riparian deforestation has a wide range of detrimental effects on aquatic biodiversity, but predicting the full consequences of such disturbances requires an understanding of the ecosystem’s key functional components. We investigated the ecology and response to sedimentation of the diverse, endemic freshwater crabs of Lake Tanganyika, which may occupy important positions in littoral foodwebs. Our surveys revealed crab distribution patterns to be patchy, and that crabs can be locally abundant (0–28 individuals m⁻²). Crab densities decreased with depth and the dry mass of crab assemblages ranged from 0.0 to 117.7 g m⁻². Comparisons among sites revealed significant effects of sedimentation on crab assemblage evenness, but provided no evidence that sedimentation has altered densities, incidence or species richness. The resilience of crabs to sedimentation might be related to their intraspecific dietary breadth. Stable isotope data (δ¹³C and δ¹⁵N) from crabs and their potential food resources indicated differences in trophic roles among endemic crab species. Overall, crabs occupy higher trophic positions than most other invertebrates, and they draw upon both benthic and planktonic energy pathways. The high biomass and top-predator status of some crab species suggests the potential for cascading effects on organisms lower in the food web.     

Fate of heterotrophic bacteria in Lake Tanganyika (East Africa)

Bacterial mortality was studied using two complementary methods between 2002 and 2004 in the two main basins (north and south) of Lake Tanganyika. The disappearance of radioactivity from the DNA of natural assemblages of bacteria previously labeled with tritiated thymidine was used to estimate the mortality due to grazing by predators (72%) and due to the cell lysis (28%). Measurements of ingestion rate of bacteria by protozoa using fluorescent micro-particles yielded protozoan grazing rates similar to those provided by the thymidine method, and showed that heterotrophic nano-flagellates were responsible for most of the grazing pressure on the bacterial community of the pelagic zone (92-99%). Bacterial cell lysis was the second process involved in bacterial mortality, ranking before ciliate grazing. Overall, bacterial mortality was balanced with bacterial production. With regard to the assessment of the trophic role of bacteria, it was estimated that c. 5-8% of the organic carbon taken up by bacteria was converted into protozoan biomass and was thus available for metazoans.     

Asterropteryx atripes, a new gobiid fish from the Western Pacific Ocean (Perciformes: Gobioidei)

 A new gobiid fish, Asterropteryx atripes, is described on the basis of eight specimens from Iriomote-jima Island, Ryukyu Islands, and El Nido, Philippines. It differs from its congeners by having the following combination of characters: 3rd spine of first dorsal fin long, filamentous, distal tip usually over end of 2nd dorsal fin base when appressed in both sexes; pelvic fins almost separated, innermost (=5th) segmented rays connected by rudimentary low membrane between bases, and no frenum; 4–7 short spines on posterior margin of preopercle (the uppermost spine usually just behind the cephalic sensory canal pore N); eye large, 32.3–35.8% of head length; enlarged haemal arches on 1st two caudal vertebrae; a distinct black band from posterior margin of eye to caudal fin base (indistinct in dark-phase individuals); black pelvic fin (vivid in dark-phase individuals); numerous minute bright blue spots on head and body in life; no distinct dark spots on head and body; iris entirely reddish-brown or dusky (bright white ventrally in pale-phase individuals) when alive or fresh, and entirely black in preservation, without white transverse bar on middorsal surface; hovering habit. The new species appears to be most closely related with the other only known hovering species, A. striata; the latter is readily distinguished from the former in having no long, filamentous dorsal spine; semitranslucent pelvic fin; and a series of small black spots along dorsal fin base and dorsal edge of caudal peduncle. Asterropteryx contains two distinct groups, and the monophyly of the genus is open to question. Received: March 19, 2000 / Revised: February 25, 2002 / Accepted: April 25, 2002     

Foraging behavior of the scale-eater Plecodus straeleni (Cichlidae,Teleostei) in Lake Tanganyika,Africa

Synopsis Sixty-nine individuals of Plecodus straeleni were followed for 1 h each in the field with the aid of SCUBA, and time budgets, hunting techniques and prey selection were investigated in relation to sex and body size. The time of cruising in midwater and on the substrate amounted to 3/4 of the total time. The rest of the time was mainly spent on five hunting techniques named pursuing, waiting, mingling, aiming and stealthy approaching. Pursuing (following a flying prey at high speed) was frequently used by adults, especially males, mainly to attack the spiny eel Afromastacembelus moorii. Waiting (keeping motionless on the substrate, waiting for a known prey) was used by some adult females when they tried to steal eggs of the mouthbrooder Cyathopharynx furcifer on the bower and by adult males when they targeted an eel having hidden under a rock. Mingling (mixing in a school of prey to attack school members) was a favorite tactic of subadults to attack plankton-feeders. Aiming (directing the head to a target fish for a moment) commonly occurred when both adults and subadults attacked solitary fishes. In stealthy approaching, the scale-eater approached an unwary prey from behind or sideway. Attacks by these hunting techniques amounted to 97% of the total attacks, which were made on 38 cichlid species and 7 non-cichlid species. Hunting techniques and prey preference varied not only with sex and size but even among consexuals of similar sizes. A number of individuals successively attacked only one or a few prey species in 1 h. Food specialization among individuals was attributed to their learning of the behavior and life style of preferred prey species.     

A new species of the cardinalfish genus Gymnapogon (Perciformes, Apogonidae) from the Red Sea

 A new cardinalfish species, Gymnapogon melanogaster, is described from two specimens collected at night in the Gulf of Aqaba, Eilat, Israel. This species is characterized by having 9 dorsal and 8 anal fin soft rays; 14–15 pectoral fin rays; 2 + 11 gill rakers; a flat, bifurcated preopercular spine; a naked body without a papillae network; black pelvic fins; and a black stomach. It is similar to Gymnapogon vanderbilti (Fowler, 1938) that is known only from the Line Islands of the Central Pacific Ocean. Received: December 26, 2001 / Revised: June 10, 2002 / Accepted: June 24, 2002 Acknowledgments We thank D. Didier and M. Sabaj of the Academy of Natural Sciences, Philadelphia, for loans of and for taking data from type specimens; T.H. Fraser of the Mote Marine Laboratory, Sarasota, kindly provided data on type specimens. We are grateful to E. Heemstra of the South African Institute for Aquatic Biodiversity, Grahamstown, South Africa, for the artwork presented in this article and to A. Lerner of the Hebrew University, Jerusalem, for his assistance in collecting the specimens. Correspondence to:Ofer Gon