Fig 1.
Map of the Grand Staircase-Escalante National Monument.
The Kaiparowits Formation outcrop is green. Solid yellow denotes newly reestablished monument boundaries. DMNH loc. 4150, the Lost Valley locality, is denoted by a star. Adapted from Crystal et al. [27].
Fig 2.
Representative stratigraphic column for the Kaiparowits Formation.
Redrawn from Roberts [29] showing major sedimentary modes. The stratigraphic position of DMNH loc. 4150, where the type and referred material for Catula gettyi was collected, was located by directly measuring section from the contact with the Wahweap Formation with the assistance of J. Hagadorn and M. Marshall in 2015.
Fig 3.
Photographs of Hell Creek taxa.
(A) Marmarthia pearsonii, (B) M. trivialis (C) “Artocarpus” lessigiana, and (D) “Ficus” planicostata. Black scale bars = 10 mm; white scale bar = 10 mm. Photographs by R. Wicker, DMNS.
Fig 4.
Overall and detail images of the holotype specimen (DMNH 54376, DMNH loc. 4150) of Catula gettyi Maccracken et al., gen. et sp. nov.
(A) Complete leaf. Arrow indicates mucronate apex. (B) Detail showing upper left section of the leaf with looping and simple brochidodromous secondary venation. The dark area on the leaf is a blotch mine. The primary vein of the leaf parallels the right side of the figure. (C) Detail showing a medial section on the right side of the leaf, with the primary vein on the left side of the figure. Note the epimedial tertiary veins with variable course (upper arrow), and the irregular to regular reticular quaternary venation (lower arrow). (D) Detail showing lower left section of the leaf with simple agrophic veins, looping ultimate marginal venation, and a fimbrial vein indicated by arrow. (E) Detail showing the base of the leaf with 2 pairs of acute basal secondary veins and laminar tissue extending down the petiole. Note the decrease in the primary vein width with the departure of the secondary veins. All scale bars = 0.5 cm.
Fig 5.
Paratype specimens of Catula gettyi.
(A) Attached leaves of C. gettyi on a stem showing opposite leaf attachment, probable distichous arrangement, and an odd-pinnate leaf terminus (DMNH 54378). Scale bar = 1 cm. (B) Enlarge section of dashed inset box in Fig 4 a showing leaf attachment (DMNH 54378). Arrows highlight axillary buds. Scale bar = 0.5 cm. (C) Attached leaves of C. gettyi on a stem showing opposite leaf attachment and an odd-pinnate leaf terminus (DMNH 54377). Scale bar = 1 cm.
Fig 6.
These specimens show the range of leaf architecture exhibited by this taxon. (a) DMNH 41570. (b) DMNH 54371. (c) DMNH 54379. (d) DMNH 54370. (e) DMNH 41584. (f) DMNH 41567. (g) DMNH 54380. Scale bars = 1.0 cm.
Fig 7.
Histogram of all damage types encountered on Catula gettyi by functional feeding group.
Red bars = generalist host specificity, gold bars = intermediate host specificity, and green bars = specialized host specificity.
Table 1.
Richness of damage types by functional feeding group and host plant specialization on Catula gettyi.
Fig 8.
Eleven hole-feeding damage types found on Catula gettyi produced by mandibulate insects.
(A) DT1; Circular holes under 1mm in diameter (DMNH 41564); (B) DT2; Circular holes between 1 mm and 5 mm in diameter (DMNH 41580); (C) DT4; Circular holes greater than 5 mm in diameter (DMNH 41596); (E) DT3; Polylobate holes between 1 mm and 5 mm in diameter (DMNH 41576); and (H) DT5; Large polylobate holes over 5 mm in diameter (DMNH 41583). Less common hole-feeding types consist of (G) DT7 & DT8 (DMNH 41590); Rectilinear holes and slot feeding, respectively (DMNH 41590); (F) DT9; Scattered, comma-shaped holes (DMNH 39733); (D) DT64; Holes located along the margin of the leaf (DMNH 41584); (I) DT51; Overlapping slot feeding holes (DMNH 41571); and (J) DT50; A series of holes associated with a primary vein (DMNH 41574). Black scale bars = 5 mm; white scale bars = 1 mm.
Fig 9.
Margin feeding damage types found on Catula gettyi at the Lost Valley Locality.
(A) DT14; Excision of the leaf to the primary vein (DMNH 41575); (B) DT13; The removal of the apex of leaf by the insect (DMNH 41578); (C) DT12; A common semi-circular excision of the leaf margin (DMNH 41579); (D) DT15; An excision that expands medially (DMNH 41585); (E) DT198; A deep, narrow excision with broad reaction tissue surrounding the herbivorized section (DMNH 41595); and (F) DT214; Multiple, connected excisions along the leaf margin (DMNH 41586). Black scale bars = 5 mm; white scale bars = 1 mm; arrows denote reaction tissues.
Fig 10.
Skeletonization damage types on Catula gettyi.
(A) DT22; Skeletonization which follows a primary vein (DMNH 41565); (B) DT19; Broad swaths of skeletonized tissue in a rectilinear shape (DMNH 41568); (C) DT24; Circular areas of skeletonization adjacent to the primary vein (DMNH 41578); (D) DT61; Elongate areas of skeletonization adjoining primary and secondary venation (DMNH 41591); and (E) DT16; Unadorned and common areas of tissue removal between veins (DMNH 41594). Black scale bars = 5 mm; white scale bars = 1 mm.
Fig 11.
Surface feeding on Catula gettyi at the Lost Valley Locality.
These damage types consist of three previously known damage types and one new damage type: (A,B) DT333 (DMNH 39725); This new damage type (DT) entails large areas of herbivory in which one surface of the leaf is removed and third-fourth order venation is left intact. Part A exhibits the rank of undamaged venation and does not have surface tissue removed. Counterpart B illustrates the tissue removal and the presence of intact third and fourth order veins. The other three surface feeding damage types are: (C) DT31; Removal of surface tissue with a distinct circular to ellipsoidal reactions rim (DMNH 41589); (D) DT30; Surface feeding with a polylobate reaction rim (DMNH 41566); and (E) DT29; Surface feeding with a weak reaction rim (DMNH 41593). Black scale bars = 5 mm; white scale bars = 1 mm.
Fig 12.
Insect damage caused by piercing-and-sucking insects.
(A) DT219; This damage type (DT) consists of two mirrored lines of puncture marks, which are potentially made by mandibles (DMNH 39724). Additional evidence of piercing-and-sucking insects comes from circular, concave puncture marks DT46 (B) (DMNH 41590); and circular, convex puncture marks DT47 (C) (DMNH 41588). A second new piercing and sucking damage type (D) involves many puncture marks across large portions or the entirety of the leaf lamina (DT330) (DMNH 41569). Scale bars = 1 mm; arrows indicate a single puncture for clarity.
Fig 13.
Galling and oviposition damage on Catula gettyi.
(A) DT33 & DT34; Galls located on primary veins and secondary veins, respectively (DMNH 41577). (B) DT32; Galls located on the laminar surface, but avoiding primary and secondary venation (DMNH 41573). (C) DT85; Galls located on the petioles of leaves or petiolules of leaves (DMNH 41592). (D) DT101; Oviposition consists of multiple, scattered ovate-shaped scars produced by an insect bearing a robust ovipositor (DMNH 41570). Black scale bars = 5 mm; white scale bars = 1 mm.
Fig 14.
Mining insect damage at the Lost Valley Locality, Kaiparowits Formation, Utah.
(A–E) DT45; Leaf mines attributed to the lepidopteran family Gracillariidae. These five mines illustrate the variation in overall shape, projection, and length, potentially due to the number of instars completed during the mining life history stage (A: DMNH 39732; B: DMNH 41582; C: DMNH 39735; D: DMNH 47124; E: DMNH 39733). The three types of blotch mines include: (F) DT36; A blotch mine lacking internal frass and a central chamber (DMNH 41572); (G) DT35; A blotch mine with frass present in circular central chamber (DMNH 41581); and (H) DT37; A blotch mine with an internal serpentine phase (DMNH 39734). Black scale bars = 5 mm; white scale bars = 1 mm.
Fig 15.
New leaf mine damage type on Catula gettyi.
DT332 includes a range of leaf mine shapes, including (A) roughly circular mines with serpentine trajectories visible within the mine (DMNH 39726), close up of mines outlined (B, D); (E) leaf mines with a circular chamber and no evident serpentine trails (DMNH 41587); (C) a linear phase followed by a solid, circular chamber (DMNH 47126), (F) (DMNH 39737), close up of mine outlined (G,I) (DMNH 41597); and (H) (DMNH 41597), (J) (DMNH 41597) mines with linear trajectories. Differences in mine form are likely attributable to the number of instars completed by each individual. These mines are typically found along the margin of C. gettyi leaves. Black scale bars = 5 mm; white scale bars = 1 mm.
Fig 16.
Rarefaction of damage types and total sampled surface area for Catula gettyi and the four Hell Creek taxa.
The Hell Creek taxa are: Marmarthia pearsonii, M. trivialis, “Artocarpus” lessigiana, and “Ficus” planicostata.
Table 2.
Comparisons of herbivory between Late Cretaceous taxa in the family Lauraceae.
Fig 17.
Non-metric multidimensional scaling (NMDS) ordination subsampled at 85,000 mm2.
This ordination includes elipses, which denote 84% of the datapoints closest to the centroid, for Catula gettyi and the four Hell Creek taxa (Marmarthia pearsonii, M. trivialis, “Artocarpus” lessigiana, and “Ficus” planicostata).
Fig 18.
Non-metric multidimensional scaling (NMDS) ordination subsampled at 140,000 mm2.
This ordination includes elipses, which denote 84% of the datapoints closest to the centroid, for Catula gettyi and three Hell Creek taxa (Marmarthia pearsonii, M. trivialis, and “Ficus” planicostata).
Table 3.
Late Cretaceous floras analyzed for herbivory.