Finally, a single basal therapsid: Cutleria

Updated May 10, 2016 with a new cladogram. 

Earlier we considered Cutleria (Fig. 1, Lewis and Vaughn 1965, USNM 22099, early Permian, 299-290mya) a basal therapsid based on its morphology, but not tested in phylogenetic analysis. Laurin (1994) redescribed Cutleria wilmarthi as a sister to the Sphenacodontidae + Therapsida at a time when therapsids were thought to descend from sphenacodontids. Wiki considers Cutleria a basal sphenacodontid.

Before today
The large reptile tree recovered a diphyletic Therapsida, with Anomodontia on one branch and the remainder of the therapsids on another.

Figure 3. Basal therapsid tree.

Figure 3. Basal therapsid tree.

Today
a phylogenetic analysis nests Cutleria as THE basal therapsid (among the 50 included taxa, Fig. 2) and it still descends from Ophiacodon. So Cutleria is still just another ophiacodontid. It had simply taken the next evolutionary step toward mammals, dicynodonts, etc. etc. And it probably was not too far from sphenacoodontids even so.  Evidently there was an early Permian radiation that included tailback pelycosaurs and basal therapsids, largely at the same time, both derived from ophiacodontids.

Figure 1. Ophiacodon, Cutleria and Stenocybus, the lineage of basal therapsids.

Figure 1. Ophiacodon, Cutleria and Stenocybus, the lineage of basal therapsids.

Earlier
we looked at the diphyletic nature of the therapsid family tree. One branch led to the Anomodontia (dromasaurs + dicynodonts and kin). The other branch (let’s call them Biarmosuchiformes) led to dinocephalians, theriodontids and by way of extension mammals and us humans.

Today we have a common ancestor for these two therapsid branches, Cutleria, in the therapsid tree at right. Cuteria does not have a single large canine, but two small canines, as in Raranimus and Stereophallodon.

Liu et al. (2009) reported, “While sphenacodontid synapsids are considered the sister-group of therapsids, the place of origin of therapsids is an enigma, largely because of a long standing morphological and temporal gap (Olson’s Gap) in their fossil record.”

Note: Tetraceratops, once thought to be a basal therapsid, is not part of this clade, as it nests with the limnoscelid, Tseajaia.

Benson (2012) recovered several basal synapsid trees, almost all with Cutleria close to the base of the Therapsida, but far from Ophiacodon. Benson erred when he included caseasaurids, which nest with millerettids in the large reptile tree. His outgroups (save Protorothyris) were largely new lepidosauromorphs like Captorhinus and Limnoscelis, which are not recovered by the large reptile tree. Hylonomus and Coelostegus would have been more appropriate and phylogenetically closer. Benson’s tree did recover Haptodus as a basal sphenacodontid, not far removed from Ophiacodon, if we get rid of caseasaurs.

References
Benson RBJ 2012. Interrelationships of basal synapsids: cranial and postcranial morphological partitions suggest different topologies. Journal of Systematic Palaeontology 10(4):601-604.
Brinkman D and Eberth DA 1986. The anatomy and relationships of Stereophallodonand Baldwinonous (Reptilia, Pelycosauria). Breviora 485: 1-34.
Cheng Z and Li J 1997. A new genus of primitive dinocephalian – the third report on Late Permian Dashankou lower tetrapod fauna. Vertebrata PalAsiatica 35 (1): 35-43. [in Chinese with English summary]
Laurin, M 1994. Re-evaluation of Cutleria wilmarthi, an Early Permian synapsid from Colorado. Journal of Vertebrate Paleontology 14(1): 134-138.
Lewis GE and Vaughn PP 1965. Early Permian Vertebrates from the Cutler Formation of the Placerville Area Colorado. United States Geological Survey Professional Papers 503-C:1-50.
Liu J, Rubidge B and Li J-L 2009. New basal synapsid supports Laurasian origin for therapsids. Acta Palaeontologica Polonica 54(3): 393-400.

Double canines at the ophiacodont/therapsid transition

Rarely will I cover scrappy specimens, known from just a few parts (Fig. 1). With so few traits, it’s difficult to nest them on the large reptile tree while maintaining complete resolution. So, generally, they are to be avoided.

However,
I recently came across the Early Permian Stereophallodon (Brinkman and Eberth 1986), described as a basal ophiacodontid with double canines. That made me think about Raranimus, described as a basal therapsid with double canines. Here (Fig. 1) they are together for the first time.

Figure 1. Stereophallodon and Raranimus, two synapsids and the ophiacodontid-therapsid transition with double canines.

Figure 1. Stereophallodon and Raranimus, two distinct synapsids with double canines at the ophiacodontid-therapsid transition. Both had tiny teeth posterior to the canines, like Biarmosuchus.

Stereophallodon (Earliest Permian) was described by Brinkman and Eberth (1986) as an ophiacodontid more primitive than Ophiacodon (Fig. 2), a taxon nesting at the base of the Therapsida in the large reptile tree. Raranimus (Early Middle Permian) was described by Liu, Rubidge and Li (2009) as a basal therapsid. Funny, they both had double canines, a trait not present in other basal therapsids, or ophiacodontids.

That kind of messes things up, doesn’t it? We may have to dismiss our linear thinking and remember that evolution produces bushes, not ladders.

Liu et al. (2009) reported, “While sphenacodontid synapsids are considered the sister-group of therapsids, the place of origin of therapsids is an enigma, largely because of a long standing morphological and temporal gap (Olson’s Gap) in their fossil record.” Unfortunately, Liu et al. (2009) did not recognize the basal nesting of Stenocybus, but did nest Tetraceratops in their tree, not realizing it nests more parsimoniously with the diadectomorph, Tseajaia. Liu et al. (2009) did not include Ophiacodon (Fig. 2) or Stereophallodon in their taxon list. Taxon exclusion is the problem once again.

Ophiacodon skull with canines highlighted in green.

Figure 2. Ophiacodon skull with double canines highlighted in green.

Not sure what else can be said of these two or their phylogenetic nesting at present, until more is known, but I thought it was worth noting.

References
Brinkman D and Eberth DA 1986. The anatomy and relationships of Stereophallodon and Baldwinonous (Reptilia, Pelycosauria). Breviora 485: 1-34.
Liu J, Rubidge B and Li J-L 2009. New basal synapsid supports Laurasian origin for therapsids. Acta Palaeontologica Polonica 54(3): 393-400.

More on Ophiacodon + therapsids

A new nesting for the two new Varanosaurus specimens according to the large reptile tree. These two nest at the base of the main group of synapsids and close to the protodiapsids (synapsid taxa leading toward the diapsid, Petrolacosaurus). Note, Ophiacodon nests three nodes away.

Figure 1. New nestings for Varanodon here, but therapsids still evolve from a sister to Ophiacodon.

Yesterday I updated my earlier Varanosaurus post because new data (taxa) had come in and certain old data reexamined with a more critical eye. Mickey Mortimer’s instructive note (see below) was important to this next step. As a result, Varanosaurus moved closer to Ophiacodon in the large reptile tree (still an outgroup, Fig. 1) and Archaeothyris nested further away, more primitively. If Archaeothyris is still considered an ophiacodontid, then Apsisaurus and Varanosaurus are too. So are all the sphenacodontids.

Moreover, as you’ll see below, several basal therapsids also share traits with Varanosaurus + Ophiacodon. This makes therapsids and mammals ophiacodontids. This is a heretical relationship previously and still overlooked that we examined and demonstrated earlier here. Now, courtesy of Mortimer’s list of Benson’s (2012) traits describing the Ophiacondontidae, the evidence is more substantial for a therapsid relationship.

M. Mortimer wrote in response to my earlier Varanosaurus post here (answers follow bullets — this starts slow, but gains momentum):

MM: Benson (2012) found the following [traits] to support Ophiacodontidae-


7. Temporal fenestra morphology: narrow dorsoventrally (<0.5x temporal height) with deep temporal bar [posterior process of the jugal] (0). (not in your matrix)

  • good trait. In evolution, fenestrae tend to start small (in this case narrow), coming from nothing (solid cheek). But note the small temporal fenestra has the same shape in therapsids only larger.

37. Maxilla, supracanine buttress on medial surface: present, may be expanded into lateral margin of internal naris [choana] (1). (not in your matrix)

  • I have few to no traits visible on inner bone surfaces. For my work I need more commonly found external traits for 340 taxa.

94. Stapes, shaft: rod-like, quadrate process small or indistinct (0); blade-like with prominent quadrate process, substantially longer than dorsal process (1). (not in your matrix)

  • I have few to no traits so tiny as this.

192. Humerus, ridge connecting deltopectoral crest to head: single, fossa absent (1). (not in your matrix)

  • I have no traits based on limb fossa.

219. Pubis, pubic tubercle anteroventral to acetabulum: present, projects dorsally (2). (not in your matrix)

  • This is character 189 on the large reptile tree. I did not notice the dorsal projection on ophiacodontids.

Further, he [Benson 2012] found Varanosaurus and Ophiacodon (but not Archaeothyris, or unknown in the latter) to share-


Here’s where it gets interesting… Yesterday I found Varanosaurus and Apsisaurus to be the outgroup to the Ophiacodontidae (including Therapsida).

2. Snout proportions, antorbital length in relation to temporal length: snout very long, antorbital length at least twice the temporal length (3). (you have a few snout length characters, but I’m not sure if any are equivalent)

14. Premaxilla, orientation of ascending (supranarial) process: slopes posterodorsally at an angle < 75° (2). (your character 13 is similar, but seemingly not equivalent)

  • Also in Biarmosuchus, Nikkasaurus many other therapsids

17. Premaxilla, marginal tooth count: five–six (1). (character 108 in yours)

  • Also in Biarmosuchus, Stenocybus, Nikkasaurus.

27. Maxilla, ventral surface: pronounced convexity (1). (character 28 in yours)

  • Also in Biarmosuchus, Stenocybus, many other therapsids

32. Maxilla, precanniniform tooth count: six or more (2). (not in your matrix)

  • Yes, perhaps by convergence or homology, not in Archaeothyris and unknown in Apsisaurus.

47. Nasal, length: longer than frontal (2). (character 25 in yours, but I’d say it’s correlated with character 2 in Benson’s, so that’s a case of his matrix being equally bad)

  • Perhaps. Also in BiarmosuchusStenocybus, many other therapsids

54. Prefrontal, lateral surface: concave, forming antorbital recess [prefrontal pocket] (1). (not in your matrix)

70. Postorbital and jugal, medial orbital process (deep, dorsoventrally tall medial flange): present (1). (not in your matrix)

  • Also in Stenocybus, unknown in Archaeothyris.

75. Jugal, posterior ramus length: long, extending to posterior end of temporal fenestra (or past temporal midlength in taxa that lack a temporal fenestra) (1). (character 63 in yours)

  • Also in Stenocybus.

76. Jugal–squamosal ventral contact, perforated by small, elongate fenestra: present, upper margin enclosed by anteroventral exension of the squamosal (1). (not in your matrix)

  • Also in Stenocybus, Nikkasaurus, many other therapsids

79. Squamosal, anteroposterior breadth of posttemporal portion on lateral surface of skull: broad, just over half the breadth of the temporal region, restricting the posterior extent of the temporal fenestra (1). (not in your matrix)

  • Also in Stenocybus, Nikkasaurus, many other therapsids

86. Tabular morphology: subrectangular sheet located dorsal to posttemporal fenestra (0). (not in your matrix)

  • Also in Stenocybus, Nikkasaurus, many other therapsids

96. Opisthotic, paraoccipital process: short and knob-like (2). (not in your matrix)

  • Also in Biarmosuchus (hard to tell, impossible on the other basal therapsids listed above)

102. Basal articulation, orientation of basipterygoid process: posteroventrally directed (2). (not in your matrix)

  • Difficult to assess in available crushed data

137. Lateral mandibular fenestra between dentary and angular: present (1). (character 124 in your matrix)

  • Also present in Stenocybus, Anomodonts.

140. Splenial, contact with posterior coronoid: present (1). (not in your matrix)

  • Don’t have many any traits visible medially.

43. Angular, cross-section shape of ventral border of angular: prominent, sheet-like keel with strongly convex posterior edge (1). (not in your matrix)

  • Don’t have any traits visible in cross-section.

155. Dorsal centra, anteroposterior length: short, subequal to height (0). (not in your matrix)

  • I have cervical ratios, but not dorsals, true.

178. Scapula, posterolateral surface of blade immediately dorsal to glenoid: deep, triangular concavity bounded anteriorly by prominent supraglenoid buttress (1). (not in your matrix)

  • I am not able to see this in that data available for Ophiacodon or Varanosaurus. Something like this appears in the later therapsid Cynognathus. Good posterior views of posterior scapulae hard to find. That’s why it’s not in my matrix.

227. Femur, mound-like eminence on dorsal surface of proximal end: small (1). (not in your matrix)

Also in Biarmosuchus. Other basal therapsids known only from skulls.

So in summary
Benson’s traits describing Ophiacodon + Varanosaurus are good, but they also apply to basal therapsids more often than not. A larger study including several missing generic taxa might have separated the Apsisaurus + Varanosaurus clade from the Ophiacodon + Therapsid clade in the Benson (2012) tree. It’s touch’n’go. They’re all very close.

And
I do move taxa when the results indicate to do so. Still these often incomplete taxa are all within just a few steps of each other.

Soon we’ll look at two double-canine ophiacodontids known from scraps that may help cross the therapsid transition.

Cutleria, Another Basal Therapsid

Laurin (1994) redescribed a small synapsid, Cutleria wilmarthi, (Lewis and Vaughn 1965) as a sister to the Sphenacodontidae + Therapsida at a time when therapsids were thought to descend from sphenacodontids. Here a close relationship to the derived ophiacodontid and basal therapsid, Stenocybus is noted (fig. 1).

Cutleria compared to Stenocybus.

Figure 1. Cutleria compared to Stenocybus. Click to enlarge. Smaller canines in Cutleria nest it closer to the early herbivore clan of Dicynodontia + Dromosauridae. Color added to restore the skull

Laurin (1994) got it right. This specimen is a close taxon to the Therapsida and not far from the Sphenacodontia. Unfortunately, Laurin (1994) published three years before Stenocybus was described by Cheng and Li (1997).

The smaller canines and lower snout mark this taxon as close to Nikkasaurus and the herbivorous clade of Dromosauridae + Dicynondontia.

As always, I encourage readers to see specimens, make observations and come to your own conclusions. Test. Test. And test again.

Evidence and support in the form of nexus, pdf and jpeg files will be sent to all who request additional data.

References
Laurin, M 1994. Re-evaluation of Cutleria wilmarthi, an Early Permian synapsid from Colorado. Journal of Vertebrate Paleontology 14(1): 134-138.
Lewis GE and Vaughn PP 1965. Early Permian Vertebrates from the Cutler Formation of the Placerville Area Colorado. United States Geological Survey Professional Papers 503-C:1-50.
Cheng Z and Li J 1997. A new genus of primitive dinocephalian – the third report on Late Permian Dashankou lower tetrapod fauna. Vertebrata PalAsiatica 35 (1): 35-43. [in Chinese with English summary]