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18 Freshwater Fishes Freshwater Fishes Freshwater fishes: high biodiversity and density I. Evolution II. Types of FW fishes Habitat % of total # individuals/species % of III. Ecology species water volume Marine 58 1010 97 Freshwater 41 108 0.01 Diadromous 1 ? • 7500x more species per unit volume in FW than SW! (1 sp. per 15 km3 vs. 1 per 113,000 km3) • 75x higher density (fish/area) in FW than SW Why are there so many FW fish species? Cause of high rate of speciation in FW: rapid evolution • productivity: freshwater habitats are generally more Evolution: a change in gene frequencies between generations productive than marine environments - shallower more sunlight more photosynthesis Causes of evolution - more terrestrial input of nutrients 1. Natural selection: best adapted individuals pass on more genes to the next generation than do less adapted individuals • isolation: FW habitats tend to be 2. Genetic drift: random processes cause certain genes to disappear isolated by: from or become fixed in a population drainages, drought, landslides, waterfalls, plate tectonics, etc. 3. Gene flow: genes enter a population from outside sources impedes gene flow 4. Sexual selection: selected genes become more common in the next can lead to explosive speciation when generation when one or both sexes (1) prefer to mate with individuals new habitats are invaded with certain phenotypes that (2) have a genetic basis BUT… • speciation will only occur if evolving populations become • in contrast, marine habitats are more connected, separated mainly by reproductively isolated continents) Case study: Causes of reproductive isolation Evolution of African Cichlids 1. Physical (geographic) isolation: populations cannot come into (family Cichlidae) contact due to physical barriers 2. Environmental isolation: populations live in different habitats 3. Behavioral isolation: mating behaviors of individuals from different populations are too different for successful reproduction 4. Mechanical isolation: sex organs are too different for mating to occur 5. Physiological isolation: hybrid offspring are not formed or have lower fitness than pure offspring 1 Evolution of African Cichlids Causes of massive speciation in African Cichlids: • most rapid and extreme case of speciation in any vertebrate group • extremely adaptable feeding structures – highly specialized & variable teeth on premaxilla • Great Rift Valley of Africa – highly specialized & variable pharyngeal jaws • ~1500 species from a handful of ancestral species • has allowed very specialized feeding: – Lake Victoria: 450 species in < 1 my (possibly 15,000 yr) – algal turfs from 1 ancestor – invertebrates – Lake Malawi: 850 species in 4-9 my from a few ancestors – fishes – scales – Lake Tanganyika: 215 species in 9-12 my from a few – eyeballs ancestors – broods of young from mothers mouth How can so many species have evolved so quickly?! numbers of species estimated by Kaufman 2007 variable tooth shape eye biter Dimidiochromis compressiceps algae scraper variable pharyngeal jaws Melanochromis auratus arthropod eater Aulonocara stuartgranti Mbuna: rock dwelling cichlids Utaka- open water zooplanktivores, Lake Malawi aufwuchs (turf aglae & inverts) 2 Surf Zone – Tanganyikan Goby Cichlids Death feigner, Lake Malawi Nimbochromis livingstoni TILEFISH Lake Tanganyika Lake Malawi Convergent GOBY SUNFISH Convergent evolution - African evolution in two cichlids that look like of the African Rift other fishes Lakes RED SNAPPER SERRANID JAWFISH TOADFISH WRASSE PARROTFISH SNOOK Causes of massive speciation in African Cichlids: Causes of massive speciation in African Cichlids: • extremely adaptable feeding structures • extremely adaptable feeding structures • specialized life history • specialized life history - mouthbrooders • extremely territorial - very limited dispersal - also limits dispersal 3 Causes of massive speciation in African Cichlids: Causes of massive speciation in African Cichlids: • extremely adaptable feeding structures • extremely adaptable feeding structures • specialized life history • specialized life history • extremely territorial • extremely territorial • geographic isolation • geographic isolation - lakes have been subdivided into isolated pools • very small populations during droughts - increases genetic drift & founder effects - lakes are huge & habitats are patchy Causes of massive speciation in African Cichlids: • extremely adaptable feeding structures • specialized life history • extremely territorial • geographic isolation • very small populations • complex mating systems - complex behaviors - elaborate color patterns (strong sexual selection) - (but not physiologically isolated — successful hybrids) Extinction! II. Types of Fishes in Freshwater • about 250 of 450 species extinct in Lake Victoria • Primary FW fishes: families strictly confined to FW, cannot tolerate SW; have a long evolutionary history in FW • causes: - e.g., characins, minnows, catfishes, sunfishes – introduced predator: Nile perch • Secondary FW fishes: families generally restricted to FW but may – increased turbidity (loss of planktivores & eutrophication) occasionally tolerate SW; originally of marine origin - e.g., cichlids, poecilids – pollution (pesticides & others) • Freshwater representatives of marine families (peripheral) - e.g., sculpins, puffers, gobies, stingrays • Diadromous fishes (peripheral): migrate from SW to FW or vice versa at different stages in their life cycles - e.g. salmon, smelt, eels • Euryhaline marine visitors – e.g., sharks, snappers 4 Primary FW fishes (~85 families) Secondary FW fishes (11 families) • Characidae • Cyprinidae • Siluriformes Characidae • Centrarchidae Centrarchidae Poecilidae Cichlidae Cyprinidae Siluriformes Freshwater representatives of marine families Diadromous fishes Salmonidae Cottidae Osmeridae Tetraodontidae Dasyatidae Anguillidae Gobiidae Euryhaline marine visitors III. Ecology of Freshwater Fishes Bull shark, Charcharhinus leucas Factors that affect the abundance and distribution of FW fishes: • zoogeography (e.g., continental drift) • physical factors (e.g., water flow) • chemical factors (e.g., pH) • biological factors (e.g., competition, predation) Lutjanidae • introductions (e.g., rainbow trout & largemouth bass) 5 Zoogeography: Continental drift Percidae (perches & darters) • Laurasian distribution Osteoglossidae (bonytongues) Cichlidae (3rd largest family of fishes) • Gondwana distribution (w/ invasion of Mesoamerica) • Gondwana distribution ~ 100 spp. Non-overlapping distributions in ecological equivalents: -- caused by biological interactions? Centrarchid Cichlidae (cichlids) Gondwana Centrarchidae (sunfishes) Nearctic continuous two nostrils lateral line Cichlid 6 South American freshwater species (2500+ spp.) African freshwater species (2000+ spp.) Asian freshwater species (28 families) Physical factors that set FW fish distributions: • temperature • light (turbidity) • gradient (steepness) • substrate • flow regime • size of water body Chemical factors that set FW fish distributions: • pH Biological factors that set FW fish distributions: • dissolved oxygen • predator-prey interactions • salinity • competitive interactions • dissolved ions • symbiotic interactions • anthropogenic pollutants 7 Temperate streams as an example Freshwater Habitats: • biological interactions are typically • streams flexible because of strong influences of physical & chemical factors • rivers • puddles results in many different • ponds combinations of species • lakes • hot (& cold) springs • caves Physical and chemical factors that cause… 1. Erosional zone Zonation of Temperate Streams • physical characteristics: high gradient, rocky bottom, swift Zones current, cold water; long riffles and small pools are main • erosional habitat • intermediate • depositional • typical fishes: streamlined, active swimmers (i.e., trout), small bottom-dwellers (sculpins and dace) fishes tend to occur in areas with particular physical and chemical characteristics Erosional Cyprinidae (longnose dace) Salmonidae (brook trout) Intermediate Depositional 2. Intermediate zone 3. Depositional zones • physical characteristics: moderate gradients, warm water, • physical characteristics: lower reaches of rivers, where intermediate current; main habitats are shallow riffles and waters are warm, turbid, and slow flowing and stream bottom deep pools with rocky bottoms or mud bottoms, and runs is generally muddy; aquatic plants can be common with undercut banks • typical fishes: deep-bodied forms that are bottom feeders • typical fishes: minnows, suckers, sunfishes, darters, (carp, suckers), planktivores (shads), invertivores (sunfish), catfishes or predators (centrarchid basses). (Same as those found in nearby lakes) Catostomidae (suckers) Percidae (darter) Cyprinidae (minnow) Catostomidae (longnose sucker) Ictaluridae (catfish) Centrarchidae (large mouth bass) Centrarchidae (sunfish) Cyprinidae (carp) 8 Tropical streams and lakes: Deserts & other seasonally dry habitats • biological factors are usually more important than physical/ chemical factors in determining fish distributions and Challenges: Adaptations: Protopteridae abundances. Why? • drying (desiccation, • diapause (dormant eggs) – climate is more stable/constant (temp. is always warm and food is concentration of solutes & • accessory respiratory structures always abundant) waste products, lack of aqueous O ) • estivation – geological stability - tropics are old 2 • altered physiology (e.g., fat – more different types of food (detritus, plant material, fruits, etc.) • heat metabolism) • eurythermal • Freshwater fishes show an amazing array of morphological • euryhaline adaptations for feeding, predator defense, habitat use, prey capture, etc. Devils hole pupfish (Cyprinodontidae) African rivuline (Nothobranchiidae) Caves Challenges: blind cave catfish (Ictaluridae) • lack of light • lack of food Adaptations: • loss/reduction of eyes • hypertrophy of other senses cavefish (Amblyopsidae) • anguilliform shape • loss of scales • low reproductive output but large offspring blind cavefish (Characidae) cave catfish (Clariidae) 9 .
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