Phytosaur

Parasuchia Huxley, 1875 Phytosaurs (Φυτόσαυροι in Greek, meaning 'plant lizard') are an extinct group of large, mostly semiaquatic Late Triassic archosauriform or basal archosaurian reptiles.

The clade Phytosauria was defined by Paul Sereno in 2005 as Rutiodon carolinensis and all taxa more closely related to it than to Aetosaurus ferratus, Rauisuchus tiradentes, Prestosuchus chiniquensis, Ornithosuchus woodwardi, or Crocodylus niloticus (the Nile crocodile).

[3] Phytosaurs were long-snouted and heavily armoured, bearing a remarkable resemblance to modern crocodilians in size, appearance, and lifestyle, as an example of convergence or parallel evolution.

Some studies of the evolutionary relationships of early archosauriforms have suggested that phytosaurs evolved before the split between crocodile- and bird-line archosaurs and are a sister taxon of Archosauria.

Fossils attributed to phytosaurs have been found in Early Jurassic rocks, possibly extending their temporal range beyond the Triassic-Jurassic boundary.

[7] Despite their great similarities in appearance and lifestyle, there are still a number of minor differences that distinguish phytosaurs from true crocodiles.

Phytosaurs were even better armoured than crocodiles, protected by heavy bony scutes (often found as fossils), and the belly reinforced with a dense arrangement of gastralia (abdominal ribs).

[9] Abler examined another sort of prehistoric predator, Dimetrodon, and found that it also lacked adaptations for guarding against crack propagation.

[16] A skeleton of Mystriosuchus planirostris, found in a marine setting and with evidence of little post-mortem transportation – indicating that it died either at sea or in a freshwater environment nearby – shows that this animal had paddle-like limbs, less adapted for terrestrial locomotion than in most other phytosaurs.

[18] Scans on various phytosaur braincases suggest that these animals generally had long olfactory tracts, weakly demarcated cerebral regions, dorsoventrally short endosseous labyrinths and various sinuses, including large antorbital and dural venous ones; the general bauplan is vaguely similar to that of crocodilians, but differs significantly in the presence of multiple sinuses, smaller cerebral hemispheres and smaller endosseous labyrinths.

The similarities are considered to be plesiomorphic in relation to the ancestral archosauriform design, lacking many features seen in avemetatarsalians, though convergence in terms of lifestyle might also play a role.

There are no clear intermediate forms, as even the earliest known phytosaurs are highly specialized aquatic animals, unlike most contemporary archosauriforms that were terrestrial.

The Carnian-Norian extinction meant that these animals died off, and the Early Norian sees new genera like Nicrosaurus and Pseudopalatus, both of which belong to the most derived clade of phytosaurs, the Pseudopalatinae.

Finally the large Redondasaurus in southwest North America and the long-snouted (altirostral) Angistorhinopsis ruetimeyeri in Europe continued the group into the Rhaetian.

Phytosaur footprints (the ichnotaxon Apatopus) are also known from the latest Rhaetian of the East Coast of USA (the Newark Supergroup) (Olsen et al. 2002).

In 1951, a partial upper jaw was discovered in the Early Jurassic Lower Lufeng Series in China and described as a new genus of phytosaur, Pachysuchus, but a study in 2012 reinterpreted the fossil as a sauropodomorph dinosaur.

[24] A fragment of a lower jaw from a longirostrine archosaur has been described from early Hettangian strata in the town of Watchet in Somerset, England.

The presence of phytosaurs in the earliest Jurassic may have prevented thalattosuchians from occupying similar ecological niches at that time.

[25] However, more recent work suggests that the jaw fragment came from a pre-Hettangian rock unit, and is therefore Late Triassic in age.

Camp Springs Formation Phytosaurs are generally regarded as the most basal group of Crurotarsi, a clade of archosaurs that includes crocodilians and their extinct relatives.

[27][28][29] Phytosaurs are often excluded from a clade called Suchia, which usually encompasses all other crurotarsans, including aetosaurs, rauisuchians, and crocodylomorphs.

[32] Below is a cladogram modified from Benton et al. (2010) showing the widely accepted phylogenetic relationships of phytosaurs:[29] Avemetatarsalia Phytosauria Aetosauria Crocodylomorpha Ornithosuchidae Rauisuchia A phylogenetic analysis of early archosaurs by paleontologist Sterling Nesbitt (2011) found strong support for a sister taxon relationship between phytosaurs and Archosauria.

Another one of the traits (an antorbital fossa contacting the horizontal process of the maxilla) was found in the basal phytosaur Parasuchus.

One trait (short metacarpals compared to metatarsals) was difficult to analyze in any crurotarsan, and another (a medial tuber on the femur) was found in both proterochampsids and Parasuchus.

The following cladogram is a simplified version the fourth strict reduced consensus tree of Ezcurra's third phylogenetic analyses within his study.

Nundasuchus Ornithosuchidae Suchia In the Late Triassic coprolite which could belong to a phytosaur, eggs of nematodes and probably protozoan cysts were found.

Illustration of two Smilosuchus species, illustrating brachyrostral and dolichorostral snout types
Phytosaur skull