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Alternative Reproductive Tactics (ARTs) (Taborsky et al. 2008: 1; Fleming and Huntingford 2012: 290), also known as “alternative reproductive phenotypes” (Sefc et al. 2008: 2531), or Alternative Mating Tactics (Taborsky 1994: 9), have been studied extensively on zoological species in general (see summaries in Oliveira et al. 2012 and Ota et al. 2012c), and in fish species in particular (e.g.

Taborsky 1994: 11, Table 1; Taborsky 2008). In Lake Tanganyika there are several species practising some kind of ART, with the best known among Lamprologini perhaps being Lamprologus callipterus (Sato 1994), Neolamprologus brevis (Ota et al. 2012c), Telmatochromis temporalis (Katoh et al. 2005), and T. Vittatus (Ota and Kohda 2006; Ota et al.

More species in the lake are likely to be added to this group as more details become known on their possibly ART-like behaviour, e.g. Telmatochromis dhonti, Altolamprologus compressiceps; on possibly ART-like behaviour in Lepidiolamprologus, see Karlsson and Karlsson (2012: 16). Recently, an alternative mating tactic was discovered regarding Variabilichromis moorii (Sefc et al. As stated above, V. Moorii is a biparental species with rather fierce defence behaviour, especially when defending the brood. Moorii would seem a very solid monogamous species. Yet, a lake-based genetic study of 10 broods of V.

Moorii (Sefc et al. 2008) revealed multiple paternity (several different fathers), suggesting that solitary wandering males of V. Moorii are likely to perform reproductive parasitism as sneakers. The sneaker is an alternative category of reproductive males (of which there are among fishes not only two types but at least four, normally genetically correlated, i.e. Phenotypes: sneaker males, satellite males, territorial males and piracy males), which are typically smaller in size, that may dart into pair-spawning territories and ejaculate sperm in order to fertilise the spawn (Ota and Kohda 2006); a sneaker male may in mouthbrooding fish taxa also be a mimicry of a female, e.g.

Mchenga eucinostomus (McKaye 1983) and Ophthalmotilapia ventralis (Haesler et al. The study (Sefc et al. 2008) suggests that there are at least two different reproductive types of V. Moorii males: small solitary sneakers and large monogamous territory holders. The Lake Tanganyika cichlids are much older than those of, for example, Lake Malawi, which is reflected not only by the greater morphological and genetic differentiation between Lake Tanganyikan species, but also by the greater genetic structures within populations and species of Lake Tanganyika cichlids, e.g. As seen in the study of V. Moorii (Duftner et al.

The Lamprologini exhibit perhaps the greatest ecological diversity in their trophic morphology and habitat of all tribes of cichlid fishes. However, all cichlid species in Lake Tanganyika are concluded to have diversified very slowly, and the assemblage seems to be derived from a prolonged accumulation of species, rather than rapid, recent radiation (Day et al. But evidence on rapid phenotypic diversification in fish, due to alternative environments during biological invasions, is commonly reported, e.g. By Lucek et al. Not unlikely, the diversification and radiation events in Lake Tanganyika took place at an early stage, and then slowed down to the present morphological stasis. Of course, exceptions exist. Isolated species with a limited range of geographical distribution may represent a relatively recent diversification and speciation event, such as the newly discovered Lepidiolamprologus kamambae (Kullander et al.

2012; Karlsson and Karlsson 2013). The morphological stasis in V. Moorii is concluded to correlate to an ecologically adaptive top, and prevails in the effect of stabilising selection favouring a certain ideal of morphology. The population structure and pattern of differentiation in V. Moorii are similar to those of the sympatric Tropheus and Eretmodus, and the three taxa also share the absence of eco-morphological diversification among populations, with colouration excluded (Duftner et al. 2006: 2388; Sefc et al.

The apparent absence in morphological diversification is further seen in another dark-coloured, multi-populational, highly stenotopic, rock-dwelling lamprologin species, Chalinochromis cyanophleps (Kullander et al. 2014b; Karlsson and Karlsson 2012); see also Staeck (2014). Regarding morphological stasis in other types of fishes, the lungfish clade, the Dipnoi, traces back in fossil records to about 400 million years, with little morphological diversification; and the West African lungfish, Protopterus annectens, a living fossil (a species being morphologically similar to its fossil ancestor), is thought to be the same species today as it was 100 million years ago, with no significant morphological alteration. Another extant lobe-finned fish and living fossil is the amazing coelacanth species Latimeria chalumnae, which has been morphologically unchanged for 70 million years; there is no doubt that morphological stasis really exists (Skelton 2001; King et al. 2011; Amemiya et al.

2013; Russell et al. 2014: 721, Fig. So80s Presents Culture Club By Blank here.

The results of the studies of V. Moorii (Duftner et al. 2006) and Eretmodus and Tropheus partially (Sefc et al.

2007), that isolated allopatric intraspecific populations with distinctly different genotypes may have more or less identical phenotypes, show not only that morphological diversification does not necessarily accompany genetic diversification, but also that morphological stasis may prevail in the presence of genetic diversification, which in turn may suggest that evolution does not necessarily always proceed at a constant and gradual pace, but rather in an erratic and stepwise manner. It is often assumed that Charles Darwin believed evolution to be smooth and gradual. Of course, for Darwin it was important to dissociate himself from the prevailing ideas of catastrophism and progressive creationism (species being supernaturally created at intervals), and possibly as a necessary consequence thereof, a perfectly gradual pace of evolution became associated with his work.

However, Darwin did not believe in a perfectly smooth and gradual type of evolution, but rather in prolonged stasis and stepwise diversification, as seen in the following passage from his The Origin of Species, 5th edition: “Many species when once formed never undergo any further change but become extinct without leaving modified descendants; and the periods, during which species have undergone modification, though long as measured by years, have probably been short in comparison with the periods during which they have retained the same form” (Darwin 1869: 551). Species may exist for millions of years without altering important specific characteristics; widespread, less specialised species may give rise to more advanced forms without becoming extinct themselves; and diversification and speciation may proceed in stepwise manners. These and many more ideas were parts of the Punctuated Equilibria (Eldredge and Gould 1972), the widely criticised macro-evolutionary theory that received increasing recognition in the nineties and onwards (Mayr 1992).

Being a theory about the change in the natural world, it generally also agrees with Darwin’s theory of evolution by natural selection (Eldredge 2006). The phenomena of morphological stasis and interrupted stability, as seen in the history of African lake faunas, are further studied in Fryer et al. On morphological stasis in Lepidiolamprologus, see Karlsson and Karlsson (2013: 16f); on stepwise diversification in Ophthalmotilapia, see Konings (1992).

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