Scott et al.—New diminutive species of Catopsalis from Alberta
columnar in lateral view. Although the first and second labial cusps are well separated fromone another, with a deepU-shaped valley developed between them, subsequent cusps are more poorly separated, and the U-shaped valley becomes shallower between each subsequent pair of cusps. The anterior and posterior surfaces of the labial cusps are virtually flat, but weak infolding of the enamel can occur on the anterior, posterior, and lingual surfaces of the fourth and fifth cusps; subvertical grooves can occur on the labial side of the first three cusps. The labial cusps are connected to one another on their labial sides by weak crests that are developed near their bases. The ultimate labial cusp can be large and bulbous (e.g., TMP 2009.133.0114, Fig. 3.15), or narrower and crest-like (e.g., TMP 2012.024.0063, Fig. 3.18). The cusps of the lingual row are nearly vertical in lateral view; the cusps increase in size posteriorly, are subquadrate to subovate in horizontal section, and are separated from one another by deep but narrow U-shaped valleys. The lingual surfaces are strongly convex, whereas the remaining sides are flat to slightly convex, and weak infolding of the enamel occurs on the anterior, posterior, and labial sides of the third and fourth cusps. A sharp crest connects the ultimate cusps of the labial and lingual rows, closing off the posterior part of the slit-like intercusp valley; the crest can be relatively short (e.g., TMP 2009.133.0114, Fig. 3.15) or expanded poster- iorly (e.g., TMP 2012.024.0063, Fig. 3.15). The posterior side of the crown angles anteroventrally-posterodorsally, and a prominent interdental facet marks the area of contact with the m2. The crown is supported by two stout roots that are both anteroposteriorly com- pressed; a heavy interradicular crest is present in one specimen (UALVP 57542). m2.—The crown of m2 is both shorter and wider than that
of m1, with the labial cusp row slightly longer than the lingual row. The anterior side of the crown is long and slopes steeply posterodorsally-anteroventrally, and the presence of a wide, strip-like interdental facet indicates that the anterior part of the m2 crown abutted the m1 underneath its sloping posterior margin. As with m1, the two cusp rows on m2 are parallel, but the individual cusps are staggered with respect to those in the opposite row. The three labial cusps decrease in size, although not height, posteriorly; each of the cusps is anteroposteriorly compressed, but the first cusp becomes appreciably swollen toward its base. The enamel of the second and third labial cusps is infolded posteriorly and lingually, and weak crests connect each of the cusps on their labial sides toward the base. The two cusps in the lingual row are massive: the first cusp is semi- circular in horizontal section, with convex anterior, labial, and lingual sides, whereas the second cusp is more quadrate. A high crest extends labially and slightly posteriorly from the second lingual cusp, then curls anteriorly toward the ultimate labial cusp, closing off the posterior part of the deep, slit-like intercusp valley. Two stout roots, with the posterior one being the larger, support the crown.
903
Etymology.—Kakwa, Cree, porcupine; the name is in reference to the occurrence of Catopsalis kakwa n. sp. in the Porcupine Hills Formation.
Materials.—Specimens of Catopsalis kakwa n. sp. were discovered at four localities in southwestern Alberta, as follows: From Zagas Quarry.—TMP 2011.110.0073, RI2; TMP
2012.024.0225, incomplete RM1; TMP 2012.024.0207, LM2; TMP 2009.133.0129, Li1; TMP 2014.043.0027, incomplete Ri1; TMP 2009.131.0001, incomplete Ri1; TMP 2009.133. 0114, Rm1; TMP 2009.133.0041, Rm2. From Narcissus.—TMP 2010.097.0020, Rm1; TMP
2010.097.0015, Rm2; TMP 2010.097.0037, incomplete Rm2. From Nordic Ski Quarry.—UAVLP 57538, LM1; UALVP
57539, incomplete LM1; TMP 2009.131.0149, incomplete RM2; UALVP 57540, 57537, Ri1; TMP 2009.131.0001, incomplete Ri1; UALVP 57541, Rp4; UALVP 57542, Rm1; UALVP 57543, incomplete Lm1. From Jumpingpound Creek Site 1.—TMP 2015.071.
0141, Lp4.
Remarks.—Taeniolabidoids are most obviously distinct among Multituberculata in their large size, but also in their possession of a suite of unusual dental features, including gigantoprismatic enamel, an anteroinferiorly restricted band of enamel on the hypsodont lower incisors, upper and lower fourth premolars that are greatly reduced in size, and enlargedmolars (Granger and Simpson, 1929; Kielan-Jaworowska and Hurum, 2001; Weil and Krause, 2008). These and several characters of the cranium were at one time thought to be synapomorphies of a larger group of multi- tuberculates, the Taeniolabidoidea, which included taeniolabidids and, variously, eucosmodontids, microcosmodontids, cimolomyids, djadochtatheriids, and several other poorly known taxa (Sloan and Van Valen, 1965; Clemens and Kielan-Jaworowska, 1979; Kielan-Jaworowska, 1980; Hahn and Hahn, 1983; Simmons, 1993). The monophyly of Taeniolabidoidea, however, was challenged by Fox (1999, 2005), who demonstrated that many of the purported taeniolabidoid synapomorphies, particularly the restricted enamel on i1, likely originated several times in multituberculates and other mammals. As such, ‘Taeniolabidoidea,’ in any of its previous incarnations, were then considered nonmonophyletic, and the phylogenetic relationships of the various ‘taeniolabidoid’ taxa (e.g., Eucosmodontidae, Microcosmodontidae, Djadochtatherioidea), both to one another and to other multi- tuberculates, remained uncertain. The evolutionary relationships of the various taeniolabidid species have been explored more recently, resulting in Taeniolabidoidea once again being resurrected, although the constituent taeniolabidoid taxa differ among the studies (com- pare, e.g., Xu et al., 2015, Williamson et al., 2015, and Mao et al., 2016). We follow Williamson et al. (2015) in recognizing a monophyletic Taeniolabidoidea that minimally includes the Taeniolabididae, aNorthAmerican family that contains Taeniolabis and Kimbetopsalis,and the genera Valenopsalis and Catopsalis,
Figure 3. Catopsalis kakwa n. sp. from localities in the Porcupine Hills Formation, early Paleocene, southwestern Alberta: (1–4) UALVP 57537, Ri1 (NSQ) in (1) lateral, (2) medial, (3) anterior, and (4) occlusal views; (5–6) TMP 2009.133.0129, Li1 (ZQ) in (5) lateral and (6) medial views; (7–9) TMP 2015.071.0141, Lp4 (JPC) in (7) labial, (8) lingual, and (9) occlusal views; (10–12) UALVP 57541, Rp4 (NSQ) in (10) labial, (11) lingual, and (12) occlusal views; (13–15) TMP 2009.133.0114, Rm1 (ZQ) in (13) labial, (14) lingual, and (15) occlusal views; (16–18) TMP 2012.024.0063, holotype, Lm1 (ZQ) in (16) labial, (17) lingual, and (18) occlusal views; (19–21) TMP 2009.133.0041, Rm2 (ZQ) in (19) labial, (20) lingual, and (21) occlusal views. JPC=Jumpingpound Creek Site 1; NSQ=Nordic Ski Quarry; ZQ=Zagas Quarry. Scale=2mm.
Page 1 |
Page 2 |
Page 3 |
Page 4 |
Page 5 |
Page 6 |
Page 7 |
Page 8 |
Page 9 |
Page 10 |
Page 11 |
Page 12 |
Page 13 |
Page 14 |
Page 15 |
Page 16 |
Page 17 |
Page 18 |
Page 19 |
Page 20 |
Page 21 |
Page 22 |
Page 23 |
Page 24 |
Page 25 |
Page 26 |
Page 27 |
Page 28 |
Page 29 |
Page 30 |
Page 31 |
Page 32 |
Page 33 |
Page 34 |
Page 35 |
Page 36 |
Page 37 |
Page 38 |
Page 39 |
Page 40 |
Page 41 |
Page 42 |
Page 43 |
Page 44 |
Page 45 |
Page 46 |
Page 47 |
Page 48 |
Page 49 |
Page 50 |
Page 51 |
Page 52 |
Page 53 |
Page 54 |
Page 55 |
Page 56 |
Page 57 |
Page 58 |
Page 59 |
Page 60 |
Page 61 |
Page 62 |
Page 63 |
Page 64 |
Page 65 |
Page 66 |
Page 67 |
Page 68 |
Page 69 |
Page 70 |
Page 71 |
Page 72 |
Page 73 |
Page 74 |
Page 75 |
Page 76 |
Page 77 |
Page 78 |
Page 79 |
Page 80 |
Page 81 |
Page 82 |
Page 83 |
Page 84 |
Page 85 |
Page 86 |
Page 87 |
Page 88 |
Page 89 |
Page 90 |
Page 91 |
Page 92 |
Page 93 |
Page 94 |
Page 95 |
Page 96 |
Page 97 |
Page 98 |
Page 99 |
Page 100 |
Page 101 |
Page 102 |
Page 103 |
Page 104 |
Page 105 |
Page 106 |
Page 107 |
Page 108 |
Page 109 |
Page 110 |
Page 111 |
Page 112 |
Page 113 |
Page 114 |
Page 115 |
Page 116 |
Page 117 |
Page 118 |
Page 119 |
Page 120 |
Page 121 |
Page 122 |
Page 123 |
Page 124 |
Page 125 |
Page 126 |
Page 127 |
Page 128 |
Page 129 |
Page 130 |
Page 131 |
Page 132 |
Page 133 |
Page 134 |
Page 135 |
Page 136 |
Page 137 |
Page 138 |
Page 139 |
Page 140 |
Page 141 |
Page 142 |
Page 143 |
Page 144 |
Page 145 |
Page 146 |
Page 147 |
Page 148 |
Page 149 |
Page 150 |
Page 151 |
Page 152 |
Page 153 |
Page 154 |
Page 155 |
Page 156 |
Page 157 |
Page 158 |
Page 159 |
Page 160 |
Page 161 |
Page 162 |
Page 163 |
Page 164 |
Page 165 |
Page 166 |
Page 167 |
Page 168 |
Page 169 |
Page 170 |
Page 171 |
Page 172 |
Page 173 |
Page 174 |
Page 175 |
Page 176 |
Page 177 |
Page 178 |
Page 179 |
Page 180 |
Page 181 |
Page 182 |
Page 183 |
Page 184 |
Page 185 |
Page 186 |
Page 187 |
Page 188 |
Page 189 |
Page 190 |
Page 191 |
Page 192 |
Page 193 |
Page 194 |
Page 195 |
Page 196 |
Page 197 |
Page 198 |
Page 199 |
Page 200 |
Page 201 |
Page 202 |
Page 203 |
Page 204 |
Page 205 |
Page 206 |
Page 207
