Palæos:		Unit 530: Altungulata
The Vertebrates		100: Altungulata

Altungulata

UNGULATOMORPHA 
|--CETARTIODACTYLA
`--Altungulata
   |--Hilalia  
   `--+--+--Perissodactyla 
      |  |  |--Hippomorpha
      |  |  `--Tapiromorpha
      |  `--Tubulidentata 
      `--Paenungulata 
         |--Hyracoidea 
         `--+--Sirenia 
            `--Proboscidea 

Taxa on This Page

1. Altungulata
2. Hilalia X
3. Hippomorpha  
4. Hyracoidea
5. Paenungulata
6. Perissodactyla
7. Proboscidea
8. Sirenia
9. Tubulidentata

Descriptions

Altungulata: horses > cows.

Phylogeny: Ungulata: Eparctocyona + * : Hilalia + (Paenungulata + (Perissodactyla + Tubulidentata))

Characters: $ petrosal does not contact basioccipital [SM98]; $ clavicle completely absent [SM98]; $ astragalar head with short neck or neck absent with flat head [SM98]; $ penial glandular fossa present [SM98].

Links: Phylo.gen; Altungulata after McKenna & Bell 1997

References: Shoshani & McKenna (1998) [SM98]. APW 020617.

Hilalia:

Range: mEc of Turkey

Phylogeny: Altungulata : (Paenungulata + (Perissodactyla + Tubulidentata)) + *.

Characters: Small to medium-sized; markedly selenodont ("selenolophodont") molars; upper canines relatively large; P2 double-rooted; P4 metacone reduced or absent; P4 metacone, if present, close to paracone; P4 without posterolingual cusp; lower premolars with talonid narrower than trigonid; p3 lacks trigonid and talonid basins (differs from perissodactyls); premolars more molariform than artiodactyls, but less than hyracoids or perissodactyls; upper molars with large mesostyle, parastyle, and paraconule; metaconule reduced or absent; crista obliqua from hypoconid terminating in large metastylid equal in size to metaconid; postcristid complete and unnotched from entoconid to hypoconid; hypoconulid absent or indistinct on m1-m3; supraorbital foramen absent; lateral pterygoid process present; tibia with shallow distal articular surface for astragalus; calcaneum with prominent, laterally-directed sustentaculum; calcaneum with large, transversely-oriented cuboid facet and no navicular facet.

Links: N/A APW 020617

References: Maas et al. (2001) [M+01].

Perissodactyla: Hyracotherium, horses, rhinos & tapirs.  Horses > elephants?

Range: from the Late Paleocene

Phylogeny: Altungulata ::: Tubulidentata + * : Hippomorpha + Tapiromorpha. 

Characters: elongate skulls; complete set of incisors; premolars molariform, often hypsodont & lophodont; horns, if present are unpaired on frontals or nasals & lack bone core; cervical vertebrae opisthocoelous; tall neural spines on thoracic vertebrae; clavicle absent; scapula is long & narrow with small coracoid process; broad, tall ilium with large gluteal (thigh) muscles; forelegs bear most weight, with hind legs providing propulsion; digits 3-4/3 (equids 1/1); "odd-toed" or mesaxonic, i.e. plane of symmetry passes through a large central digit; astragalus is primary weight-bearing tarsal; intestinal fermentation, normally requiring constant foraging, but superior use (compared to artiodactyl ruminants) of proteins and high-quality food materials. 

Links: Introduction to the Perissodactyla; Fossil Horses in Cyberspace; The Tapir Gallery - Welcome!; Perissodactyla; Order PERISSODACTYLA; Order Perissodactyla: Rhinoceroses, Horses; ISIS - taxon perissodactyla; Journal of Molecular Evolution 50: 11-21 (2000); Order Perissodactyla; Rhinoceros Skulls and Rhino Horns from Skulls Unlimited; Perissodactyla (Hoofed (Odd-toed)); Bibliography of Ungulates: Perissodactyla and Artiodactyla, etc.; Breeds of the World; Vertebrates : Mammals : Perissodactyla; World's oldest horse runs free in Mongolia.  APW020617.

Hippomorpha: Equus + Pachynolophus 

Range: Ypresian (Early Eocene) [A+98] [NA00] (both based on Hyracotherium).  Dominant ungulates in the Eocene.  

Phylogeny: Perissodactyla : Tapiromorpha + * 

References: Arnason et al. (1998) [A+98], Norman & Ashley (2000) [NA00].  

Tubulidentata: aardvarks Archaeorycteropus, Leptorycteropus, Myorycteropus, Orycteropus, Palaeorycteropus, Plesiorycteropus. 

Range: From the Early Miocene of Europe and Africa. 

Phylogeny: Altungulata ::: Perissodactyla + *. 

Characters: Skull elongate & conical; thin, protrusible tongue; dentary elongate and slender, bladelike anteriorly & wider at molars; elaborate turbinals and highly-developed olfactoray centers of brain; nares can be sealed with hair; small tentacular (chemosensory?) structures on nares; zygomatic arch complete; poor vision; palate ends at the posterior end of the palatines & is not extended posteriorly by the pterygoid (see tubulidentata.jpg from the ADW); auditory pinnae long and pointed; dental formula (adult) 0/0, 0/0, 2-3/2, 3/3; teeth rootless and ever-growing, without enamel, columnar, and made up of ~100µ hexagonal prisms of dentine and pulp core; teeth surrounded by cementum; zygapophyses well-developed and interlocked; moderately long, muscular tail; very powerful forelimbs; digits 4/5, with long, spatulate claws; exceptionally fast digging ability; digitigrade, but slow locomotion; thick skin, almost hairless; insectivores, largely by digging for social insects; solitary; low basal metabolism and slightly lower than expected temperature (34.5°); nocturnal. 

Links: Order Tubulidentata; Aardvarks!; Tubulidentata; Order Tubulidentata: Aardvark; Comparative Mammalian Brain Collections: TUBULIDENTATA; Tubulidentata; Tubulidentata; tubulidentata page; Order Tubulidentata; Aardvark - Wikipedia; TUBULIDENTATA; ORYCTEROPODIDAE: Aardvark, or Ant Bear; Aardvark; Google Directory - Science > Biology > Flora and Fauna > Animalia ...; Aardvarks (Tubulidentata) - MavicaNET.  ATW031107.

Paenungulata: elephants, sea cows, & hyraxes. 

Range: Fr upPc. 

Phylogeny: Altungulata :: (Perissodactyla + Tubulidentata) + * : Hyracoidea + (Sirenia + Proboscidea). 

Characters: Bifurcated styloglossus muscle (see also Fig. 1019); nails, rather than hooves; digits usually 4/3; 

Links: Phylo.gen; Paenungulata; SIRENIA: Manatees, Dugong, and Sea Cow APW 020617.

Hyracoidea: Hyraxes. 

Range: Fr lwEc of Afr & Eur; fl lwOc. 

Phylogeny: Paenungulata : (Sirenia + Proboscidea) + *. 

Characters: Small rodent or rabbit-like forms with some large extinct relatives. Short skull with short rostrum; dental formula 1/2, 0,0, 4/4, 3/3; upper incisors, widely separated, triangular, tusk-like & ever-growing, with enamel only on lingual face; lower incisors wedge-shaped & tricuspid; small diastema present; upper molars with ectoloph & 2 cross-lophs; lowers lilambdodont (?); deep lower jaw; orbital bar present; zygomatic arch complete; jugal large & contributes to glenoid; small auditory bullae; compact body; tail highly reduced; clavicle absent; digits 4(5?)/3, with flat nails ("hoof-like"?) except for claw on pes 2; mesaxonic (III); palm/sole covers metapodials with thick, elastic, glandular skin which can be cupped for traction; stomach simple, with 3 ceca; mainly herbivorous, with some omnivory; metabolism low & behavioral thermoregulation common; able climbing & jumping; fairly wide variety of guilds and ecological settings. 

Links: Introduction to the Hyracoidea; Order Hyracoidea/Family Procaviidae; hyrax page (but the line drawing has 2/3 incisors, no postorbital bar & jugal does not contribute to glenoid. Hyracoid?); Hyracoidea; Other Videos; Order HYRACOIDEA; Order Hyracoidea: Hyraxes; Schliefer; Hyracoidea; Subungulates. 

Image: Procavia capensis from Utah's Hogle Zoo (the object which looks like a cigarette hanging out of the Hyrax's mouth is one of its modified upper incisors).

Sirenia: Manatees (Trichechus)& dugongs (Dugong). 

Range: Fr lwEc. 

Phylogeny: Paenungulata :: Proboscidea + *. 

Characters: Large coastal marine herbivores. Skull short, with strongly developed premaxillary; dentition variable among species; primitively 5(!) premolars; essentially modern postcranium from earliest known remains of mEc age; nostrils valvular, extending posteriorly beyond anterior rim of orbit; nasals reduced or absent; dentary deep; rostrum and dentary deflected ventrally at anterior end; tympanic semi-circular; periotic not connected to skull (as in cetaceans); heavy pachostotic ribs; muscular tail with flukes; digits 5/0, with forelimb as flipper; strongly developed deltopectoral crest [T+01]; hindlimbs absent; pelvis vestigial; lungs unlobed, long, & oriented horizontally with a horizontal diaphragm (used for floatation); heavy, dense bone; large, mid-gut cecum; very slow metabolism and low-nutrient diet; metabolism & relatively poor insulation restrict range to warmer waters; complex oral signals; social behavior not well known. 

Links: Encyclopedia Smithsonian: References on the Sirenia; Introduction to the Sirenia; Order Sirenia; Dugong Page (English); Sirenia; Manatee Skull - Trichechus manatus; Dugongs: The UAE is one of the last refuges of these endangered sea-mammals APW 020617. 

References: Thewissen et al. (2001) [T+01].

Image: from Welcome to Dubai - the desert's most exciting city!.

Proboscidea: Elephants. 

Range: Fr lwEc (upPc?) of Afr Asia, Eur, & NAm. 

Phylogeny: Paenungulata :: Sirenia + *. 

Characters: Very large (up to 7500 kg) herbivores with trunks. Skull anteroposteriorly compressed & tall, pneumatized; large tusks formed from I2 (I1?) with enamel only on tip; in recent forms; usually only one very large, loxodont, cheek tooth is functional at any given time; cheek teeth with numerous ridges formed from enamel, dentine coat and cementum between ridges; narial proboscis with 1-2 finger-like structures at distal end; premaxilla specialized as sheath for tusks; marked lambdoid crest (see the ADW elephantidae.jpg); neck short; small tail; long axis of pelvis almost 90º to spine; acetabulum faces ventrally; graviportal, unspecialized, columnar limbs; proximal segments longest; heel pad of fibrous tissue supports most of weight; short digits 5/5; massive bones; limb bones without marrow & with spongy bone core; non-selective browsers(?); complex social structures and vocalizations; infrasound (14-24 Hz) long-distance communications. 

Links: Introduction to the Proboscidea; Elephant Information Repository: Elephant Resources (mostly a conservation site, but some science); Illinois State Museum; Order Proboscidea;  proboscidea page; Order PROBOSCIDEA (always a great site, but this specimen is particularly worth a look); Order Proboscidea: Elephants; Mammoths; Proboscidea; Phylum Chordata / Class Mammalia / Order Proboscidea; Proboscidea (Trunked Mammals); ADW- Proboscidea- Information; proboscidea page (presumably a student page, with several layout errors -- but lots of good information); Proboscidea (the usual aging ToL page, but many good older references); Proboscidea (Ultimate Ungulate page); Proboscidea (grade-school level site); Basal Proboscidea (Mikko's Phylogeny); PROBOSCIDEA; ELEPHANTIDAE (extract from Walker's); Proboscidean Mammals- EnchantedLearning.com; Elephant Evolution (not very useful); Comparative Mammalian Brain Collections- PROBOSCIDEA (fair amount of background on Elephas as well); Elephants - facts and information about elephants ( a 300-page site on elephants.  Easily Best on the Web). 

Image: Ganesh riding on his bearer and good friend, the rat. 

Note: the heavy bones, columnar limbs and pneumatized skull may have developed before the evolution of great size. ATW040703.

checked ATW050727