The Vertebrates Ferungulata

Laurasiatheria: Ferungulata

Abbreviated Dendrogram
Eutheria ├─Euarchontaglires ├?─Chiroptera └─Laurasiatheria ├?─Insectivora └─Ferungulata ├─┬─Ferae │ └─Pholidota └─┬─┬─Phenacodontidae │ └─Perissodactyla └─Eparctocyona ├─┬─Arctostylopida │ └─┬─Mesonychia │ └─Cetartiodactyla └─Bulbulodentata ├─Hyopsodontidae └─Meridiungulata


Taxa on This Page

  1. Hyopsodontidae X
  2. Phenacodontidae X
  3. Ferungulata


Ferungulata Simpson 1945

Synonyms: Cetferungulata, Fereuungulata

Range: from the Latest Cretaceous? (assuming Protungulatum - controversial palcement) or Earliest Paleocene

Laurasiatheria : ?Insectivora + ?Chiroptera (both controversial placements because of molecules - morphology incongruency) + * : (Ferae + Pholidota) + ((Phenacodontidae + Perissodactyla) + Eparctocyona)

Comment: Originally (in the old Palaeos) this place was held by Ungulata defined as "Mammals with hooves. Used here as horses + cows." This morphology-based clade is incompatable with molecular phylogeny, which divides ungulates among the Afrotheria and Laurasiatheria. The morphology-based cladistic analysis of Wible et al 2007 and 2009 rejects a monophyletic Laurasiatheria but fails to recover the Ungulata. As no current workers , in this field accept that the ungulates are a monophylertic taxon, we have likewise rejected it in the revised Palaeos., although we have included the original entries (should the phylogenetic consnesus swing the other way) . Until then, Ungulata is replaced by Ferungfulata

Ferungulata is traditionally a cohort (a non-standard taxon rank, here between infraclass and superorder) within the placental mammals. Established by rnowned mamalian zoologist and plaeontologist George Gaylord Simpson on morphological grounds, and uniting both ungulate herbivores and carnivorous mamamls, it includes the extant Carnivora, Perissodactyla, Artiodactyla (but not whales (Cetacea) which were at the time still considered distinct), Tubulidentata, and Paenungulata, as well as a number of fossil taxa.. With the paradigm shift resulting from rise to dominance of molecular phylogeny in the 1990s and 2000s, morphology-based criteria are now rejected whenever they contradict genome sequencing. Nevertheless, a modified form of Simpson's Ferungulata was recovered, with the Pholidota added but the Tubulidentata and Paenungulata are relocated to the Afrotheria . To distinguish this new taxon a variant (and somewhat unpronounceable) spelling has been used: "Fereuungulata". Since this is still essentially the same taxon (uniting hooved and carnivorous mammls) we see no reason to change the original spelling. - MAK120717

Currently, the Ferungulata includes the living orders Carnivora, Pholidota, Cetartiodactyla and Perissodactyla. Though supported by molecular analyses (Arnason et al. 2002; Asher et al. 2003), the clade has not yet gained morphological support, with the possible exception of a compatible clade being identified by Wible et al. (2007) (who did not reconstruct supporting character states for any specific nodes). As well as the surviving orders, there are a number of extinct 'ungulate' groups that may be referrable to this clade. In the tree below, the extinct groups are related to the modern orders on the basis of a number of studies of smaller subgroups, but a broader analysis of the extinct placentals is sorely needed. - CKT110403

The original Ungulata entry reads as follows: Characters: $ Stylomastoid foramen opens dorsal to level of cheek teeth [SM98]; Canines reduced [d&C00]; bunodont molars; m3 hypoconulid larger than m2 hypoconulid [d&C00]; astragalus with short, robust head; $? patella rounded or approximately ball-shaped [SM98]; $? distal phalanges spatulate [SM98].

Links: UNGULATA; The Ultimate Ungulate Page; Mammalia/UNGULATA/ungulata.html">Mammalia, Ungulata - Huftiere; ordo Ungulata; Ungulates; Ungulata Catalog (mostly catalog -- a few, very high quality, technical photos); C. Savona-Ventura: Maltese Unglulates.; Salles; Wikipedia (with a very handsome llama); Ungulata Baskin's course notes, with some good links); Paleocene mammals of the world Best on the Web).

References: de Muizon & Cifelli (2000) [d&C00]; Shoshani & McKenna (1998) [SM98]. ATW0.

The more inclusive Ungulatomorpha (which also includes Zhelestidae and Dinocerata has turned out to be polyphyletic, the zhelestids being stem eutherians convergent with ungulates. The position of the dinocerata is uncertain, we have tentatively placed them in the stem eutherians as well, but more derived than zhelestids. For what it's worth, here's the original entry: Characters: premolars simple [d&C00]; small metaconid on p4 [d&C00]; trigonid cusps sharply conical & well separated (metaconid & paraconid fuse in almost all derived groups) [d&C00]; trigonid open lingually [d&C00]; lower molars with at least weak postcingulid [d&C00]; m2 trigonid slightly wider than talonid (reversed in Zhelestids?) [d&C00]. Links: Basal Ungulatomorpha; g00n1a4.pdf; MESOZOIC MAMMALS; 'basal' Ungulatomorpha, an internet directory: (Best on the Web -- most of the other links are taken from this one); vol3-p18; Welcome to Adobe GoLive 4; Re: Condylarths (TINGAMARRA REVISITED); Ungulate-Like Mammals from the Late Cretaceous of Uzbekistan and ....; The earliest eutherian mammal. References: de Muizon & Cifelli (2000) [d&C00]. ATW021027.  links checked ATW040213, updated MAK120321 (links not checked)

Life restoration of Meniscotherium

Phenacodontidae: Eoconodon, Meniscotherium, Phenacodus.

Range: Paleocene to Eocene of Europe and North America.

Ferungulata : (Ferae + Pholidota) + (Eparctocyona + (Perissodactyla + * ))

(from Thewissen 1990): Teeth bunodont to bunolophodont. P4 with talonid basin and subequal protoconid and metaconid. Paraconid of lower molars absent or small. P3 with small protocone, P4 with paracone, metacone, and protocone. M1-2 with hypocone and often with mesostyle. Dental formula / Humerus with weak deltopectoral crest and supracondylar foramen. Ulna strong with anterior process distally. Five digits in hand and foot. Femur with third trochanter. Fibula complete. Astragalus with convex head. -


Comments: The Phenacodontidae are a family of herbivorous cursorial mammals from the Palaeocene and Eocene of North America and Europe. Size estimates for phenacodontids vary from about three or four kilograms for the smaller Ectocion and Copecion species, to well over fifty kilos for the larger Phenacodus species (Thewissen 1990), or about the size of a cat to that of a small sheep. - CKT110413

Image: Life restoration of Meniscotherium from W.B. Scott's A History of Land Mammals in the Western Hemisphere. New York: The Macmillan Company, 1913 (copyright exzpired - public domain. Art by Robert Bruce Horsfall. Via Wikipedia

Bulbulodentata: stem group including the paraphyletic "Mioclaenidae" as well as the Periptychidae.

Range: Early Paleocene to Late Eocene of Africa, Europe, South America, North America

Characters: premolars "swollen";

Phylogeny: Eparctocyona : (Arctostylopida + (Triisodontidae + Mesonychia + Cetartiodactyla))) + * : Hyopsodontidae + Meridiungulata.

Links: Bulbulodentata Mikko); Species Skull Ht Lt Wt Time Range Acama probably Acamana U ... time & place); de Muizon & Cifelli important article with full text pdf -- Best on the Web). ATW021224.

HyopsodonHyopsodontidae:  Hyopsodon, Litomylus, Louisina, Paschatherium.

Range: Late Paleocene to Late Eocene of Europe, Africa, North America & ?China. Probably not South America. Most common in North America.

Phylogeny: Bulbulodentata : Meridiungulata + *.

Characters: insectivore-like animals with arboreal capabilities; hypocone large; parastylar lobe present and larger on M3 than M2; primitively, preparacrista & postmetacrista oriented mesiodistally; trigonid relatively open lingually; paraconid not appressed to metaconid; entoconid large; hypoconid well developed.

Image: Hyopsodus from Paleocene Mammals of the World, which credits Savage, RJG & MR Long (1986), Mammal Evolution. An Illustrated Guide. British Museum (Natural History).

Notes: Recent work suggests that this may be a synonym for Mioclaenidae, a well-known condylarth group. In addition, some hyopsodonts the group may be polyphyletic) may be the sister group, or even include, the "Afrotherian" clade of (at least) elephant shrews (Macroscelidea) [Z+05]. Note that this has resonances in the some of the web citations below.

Links: Hyopsodontidae Mikko); Untitled Document suggesting a relationship with elephant shrews and elephants); Bootstrap method with heuristic search-; Conddylarths (Steve & Casey's); Untitled (Czech); Turtles (and Afrotheria) (long); The "condylarths" (archaic Ungulata, Mammalia) from the early ...; Main Page Ben Berger's page, which was long extinct, seems to have been resurrected); Paleocene mammals of the world suggesting a link with insectivores); Turtles (and Afrotheria) (long) same); Volume 31 No. 2, abstract of Robinson article showing odd evolutionary pattern); Институт проблем экологии и эволюции имени А. Н. Северцова.(Russian: abstract of Agadzhanian article arguing hyopsodonts are ancestral to tapirs and indricotheres).

References: Zack et al. (2005) [Z+05]. ATW030714.    

checked ATW040213, this p[age MAK120321, revised MAK120717