Sunday 20 September 2020

EVOLUTION OF FISH

The evolution of fish began about 530 million years ago with the first fish lineages belonged to the Agnatha, a superclass of jawless fish.

We still see them in our waters as cyclostomes but have lost the conodonts and ostracoderms to the annals of time. 

Like all vertebrates, fish have bilateral symmetry; when divided down the middle or central axis, each half is the same. Organisms with bilateral symmetry are generally more agile, making finding a mate, hunting or avoiding being hunted a whole lot easier. While we still find them on our menus, the ability to move quickly means they avoid being the snack of choice, an honour that falls more to the invertebrates with whom they share the sea.

When we envision fish, we generally picture large eyes, gills, a well-developed mouth. The earliest animals that we classify as fish appeared as soft-bodied chordates who lacked a true spine. While they were spineless, they did have notochords, a cartilaginous skeletal rod that gave them more dexterity than the cold-blooded invertebrates who shared those ancient seas and evolved without a backbone.

Fish would continue to evolve throughout the Paleozoic, diversifying into a wide range of forms. Several forms of Paleozoic fish developed external armour that protected them from predators. The first fish with jaws appeared in the Silurian period, after which many species, including sharks, became formidable marine predators rather than just the prey of arthropods.

Fishes in general respire using gills, are most often covered with bony scales and propel themselves using fins. There are two main types of fins, median fins and paired fins. The median fins include the caudal fin or tail fin, the dorsal fin, and the anal fin. Now there may be more than one dorsal, and one anal fin in some fishes.

The paired fins include the pectoral fins and the pelvic fins. And these paired fins are connected to, and supported by, pectoral and pelvic girdles, at the shoulder and hip; in the same way, our arms and legs are connected to and supported by, pectoral and pelvic girdles. This arrangement is something we inherited from the ancestors we share with fishes. They are homologous structures.

When we speak of early vertebrates, we're often talking about fishes. Fish is a term we use a lot in our everyday lives but taxonomically it is not all that useful. When we say, 'fish' we generally mean an ectothermic, aquatic vertebrate with gills and fins.

Rhacolepis Buccalis, an extinct genus of ray-finned fossil fish
Fortunately, many of our fishy friends have ended up in the fossil record. We may see some of the soft bits from time to time, as in the lovely Rhacolepis Buccalis, an extinct genus of ray-finned fossil fish in carbonate concretion from the Lower Cretaceous, Santana Formation, Brazil.

Not surprisingly, vertebrates with hard skeletons have a much better chance of being preserved than those with just soft parts and no teeth or bone to speak of.

In British Columbia, we have lovely two-dimensional Eohiodon rosei, a common freshwater fossil fish well-represented in Eocene deposits from the Allenby of Princeton and McAbee Fossil Beds near Cache Creek. We also have the Tiktaalik roseae, a large freshwater fish, from 375 million-year-old Devonian deposits on Ellesmere Island in Canada's Arctic. Tiktaalik is a wonderfully bizarre creature with a flat, almost reptilian head but also fins, scales and gills. We have other wonders from this time. There are also spectacular antiarch placoderms, Bothriolepsis, found in the Upper Devonian shales of Miguasha in Quebec.

There are fragments of bone-like tissues from as early as the Late Cambrian with the oldest fossils that are truly recognizable as fishes come from the Middle Ordovician from North America, South America and Australia. At the time, South America and Australia were part of a supercontinent called Gondwana. North America was part of another supercontinent called Laurentia and the two were separated by deep oceans.

Eohiodon rosei, McAbee Fossil Beds
These two supercontinents and others that were also present were partially covered by shallow equatorial seas and the continents themselves were barren and rocky. Land plants didn't evolve until later in the Silurian Period.

In these shallow equatorial seas, a large diverse and widespread group of armoured, jawless fishes evolved: the Pteraspidomorphi. The first of our three groups of ostracoderms. The Pteraspidomorphi are divided into three major groups: the Astraspida, Arandaspida and the Heterostraci.

The oldest and most primitive pteraspidomorphs were the Astraspida and the Arandaspida. You'll notice that all three of these taxon names contain 'aspid', which means shield. This is because these early fishes and many of the Pteraspidomorphi possessed large plates of dermal bone at the anterior end of their bodies. This dermal armour was very common in early vertebrates, but it was lost in their descendants.

Arandaspida is represented by two well-known genera: Sacabampaspis, from South America and Arandaspis from Australia. Arandaspis have large, simple, dorsal and ventral head shields. Their bodies were fusiform, which means they were shaped sort of like a spindle, fat in the middle and tapering at both ends. Picture a sausage that is a bit wider near the centre with a crisp outer shell.