Comparative phylogeography of the Yellow River schizothoracine fishes (Cyprinidae): Vicariance, expansion, and recent coalescence in response to the Quaternary environmental upheaval in the Tibetan Plateau

National Natural Science Foundation of China (NSFC) [30225008, 30300036, 30530120]; Key Innovation Plan [KSCX2-SW-106]; National Basic Research Project in China [2005cb422005]; National Natural Science Foundation of China [30600062]

The youngest split in sympatric schizothoracine fish (Cyprinidae) is shaped by ecological adaptations in a Tibetan Plateau glacier lake

Although new empirical evidence shows that sympatric speciation has occurred in some species, there are few indisputable model organisms for this process of speciation. The two subspecies (Gymnocypris eckloni eckloni and G. e. scoliostomus) of the schizothoracine Gymnocypris fish species complex from a small glacier lake in the Tibetan Plateau, Lake Sunmcuo, fit several of the key characteristics of the sympatric speciation model. We used combined mitochondrial control region sequences and the cytochrome b gene (1894 bp) to address the phylogenetics and population genetics of 232 specimens of G. e. eckloni and G. e. scoliostomus, as well as all of its closely related sister species. We found that: (i) a total of four old lineages were uncovered in the widespread G. e. eckloni, of which only one was shown to be shared with all G. e. scoliostomus individuals and (ii) the new subspecies (G. e. scoliostomus) evolved in Lake Sunmcuo from the ancestral G. e. eckloni population within approximately 0.057 Ma. These two taxa of the species complex are morphologically distinct, and reproductive isolation is further suggested. Ecological disruptive selection based on morphological traits (e.g. mouth cleft characters) and food utilization may be a mechanism of incipient speciation of two sympatric populations within Lake Sunmcuo. This study provides the first genetic evidence for sympatric speciation in the schizothoracine fish.

Age and Growth of an Endemic Tibetan Fish, Schizothorax o'connori, in the Yarlung Tsangpo River

We investigated the age and growth of Schizothorax o'connori in the Yarlung Tsangpo River by examination of annuli from otoliths. The von Bertalanffy model was the most acceptable statistical growth model. Its parameters were as follows: SL infinity = 492.4 mm, K = 0.1133, t(0) = -0.5432 year and W-infinity = 1748.9 g for females; SL infinity = 449.0 mm, K = 0.1260, t(0) = -0.4746 year and W-infinity = 1287.0 g for males. Theoretical longevity was 25.9 years for the female and 23.3 years for the male. Moreover, females had larger asymptotic length and weight compared with males.

Extraordinarily thick-boned fish linked to the aridification of the Qaidam Basin (northern Tibetan Plateau)

Scattered with numerous salt lakes and approximate to 2,700-3,200 m above sea level, the giant Qaidam inland basin on the northern Tibetan Plateau has experienced continuing aridification since the beginning of the Late Cenozoic as a result of the India-Asia plate collision and associated uplift of the Tibetan Plateau. Previous evidence of aridification comes mainly from evaporite deposits and salinity-tolerant invertebrate fossils. Vertebrate fossils were rare until recent discoveries of abundant fish. Here, we report an unusual cyprinid fish, Hsianwenia wui, gen. et sp. nov., from Pliocene lake deposits of the Qaidam Basin, characterized by an extraordinarily thick skeleton that occupied almost the entire body. Such enormous skeletal thickening, apparently leaving little room for muscles, is unknown among extant fish. However, an almost identical condition occurs in the much smaller cyprinodontid Aphanius crassicaudus (Cyprinodonyiformes), collected from evaporites exposed along the northern margins of the Mediterranean Sea during the Messinian desiccation period. H. wui and A. crassicaudus both occur in similar deposits rich in carbonates (CaCO3) and sulfates (CaSO4), indicating that both were adapted to the extreme conditions resulting from the ariclification in the two areas. The overall skeletal thickening was most likely formed through deposition of the oversaturated calcium and was apparently a normal feature of the biology and growth of these fish.

C-values of seven marine mammal species determined by flow cytometry

C-values, which estimate genome size, have puzzled geneticists for years because they bear no relationship to organismal complexity. Though C-values have been estimated for thousands of species, considerably more data are required in order to better understanding genome evolution. This is particularly true for mammals, in which C-values are known for less than 8% of the total number of mammalian species. Among marine mammals, a C-value has been estimated only for the bottlenose dolphin (Tursiops truncatus). Thus examination of additional species of marine mammals is necessary for comparative purposes. It will enable a better understanding of marine mammal genome evolution, and it is also relevant to conservation, because larger genome size has been linked to increased likelihood of extinction in some plant and animal groups. Our study presents C-values of seven marine mammal species, including five cetacean species that are endangered to varying degrees. Similarly to the results for other groups, our results suggest that larger genome size in cetaceans is related to an increased likelihood of extinction.

The reproductive strategies of an endemic Tibetan fish, Gymnocypris selincuoensis

The reproductive traits of Gymnocypris selincuoensis from Selincuo Lake and its tributaries were investigated in 1997 and 1998. The youngest mature male was age 7 with a standard length (SL) of 172.0 mm, and the youngest mature female was age 8 with a SL of 194.0 mm. The L(50)s Of SL and age at first maturity were respectively 250.32 mm and age 9 for males and 224.71 mm and age 8 for females. The gonadosomatic index (GSI) significantly changed with seasons for mature individuals but not for immature individuals. GSIs of mature females at stages IV and V of ovary development increased with SL and reached a maximum value at the SL range from 370 mm to 390 mm; the GSIs of mature males were negatively correlated with SL. The breeding season lasted from early April to early August. Egg size did not significantly change with SL but increased with the delay of spawning. The individual absolute fecundity varied from 1,341 to 28,002 eggs (mean 12,607+/-7,349), and the individual relative fecundity varied from 6.4 to 42.0 eggs.g(-1) (mean 25.5+/-9.7). The individual fecundity increased with total body weight; it also increased with SL for those of SL less than 370 mm. There was a rest of spawning for mature individuals.

Molecular phylogeny of the specialized schizothoracine fishes (Teleostei : Cyprinidae), with their implications for the uplift of the Qinghai-Tibetan Plateau

Molecular phylogeny of three genera containing nine species and subspecies of the specialized schizothoracine fishes are investigated based on the complete nucleotide sequence of mitochondrial cytochrome b gene. Meantime relationships between the main cladogenetic events of the specialized schizothoracine fishes and the stepwise uplift of the Qinghai-Tibetan Plateau are also conducted using the molecular clock, which is calibrated by geological isolated events between the upper reaches of the Yellow River and the Qinghai Lake. Results indicated that the specialized schizothoracine fishes are not a monophyly. Five species and subspecies of Ptychobarbus form a monophyly. But three species of Gymnodiptychus do not form a monophyly. Gd. integrigymnatus is a sister taxon of the highly specialized schizothoracine fishes while Gd. pachycheilus has a close relation with Gd. dybowskii, and both of them are as a sister group of Diptychus maculatus. The specialized schizothoracines fishes might have originated during the Miocene (about 10 MaBP), and then the divergence of three genera happened during late Miocene (about 8 MaBP). Their main specialization occurred during the late Pliocene and Pleistocene (3.54-0.42 MaBP). The main cladogenetic events of the specialized schizothoracine fishes are mostly correlated with the geological tectonic events and intensive climate shift happened at 8, 3.6, 2.5 and 1.7 MaBP of the late Cenozoic. Molecular clock data do not support the hypothesis that the Qinghai-Tibetan Plateau uplifted to near present or even higher elevations during the Oligocene or Miocene, and neither in agreement with the view that the plateau uplifting reached only to an altitude of 2000 in during the late Pliocene (about 2.6 MaBP).