Hendricks—Miocene Conidae from the Gatun Formation of Panama follows.


Second, the asymmetry of the SSF (ASSF) was calculated as ASSF =SSFW1 = SSFW2


(5) Species with ASSF values <1 have asymmetrical SSFs,


while those with ASSF values of ~1 have symmetrical SSFs. Note that Harzhauser and Landau (2016) very recently pioneered an alternative approach to quantifying the shape of the SSF, wherein measurements were collected from images taken at an oblique angle relative to the SSF. Subsutural flexure data for individual specimens are presented in Supplementary data set 3. The terminology used here to characterize and describe


preserved shell coloration patterns revealed by UV light follows the recently developed approach of Hendricks (2015), which in turn was built upon earlier descriptive terminology developed by Röckel et al. (1995), Hendricks (2009), and Kohn (2014). Briefly, some cone snail shells show elements of coloration patterning that seemto occupy a single layer (e.g., rows of spiral blotches covering ashell’s last whorl). Others seemingly show two layers of patterning, with a basal (or, primary) layer that appears to be overprinted by a secondary layer (e.g., axial streaks overprinted by spiral bands). In some cases, these two layers of patterning show no evidence of interactions (i.e., a noninteracting pattern, where one pattern simply appears to cover another). In other cases, however, the two layers may interact with one another where they intersect (i.e., an interacting pattern); these interactions are described on a case-by-case basis. See Hendricks (2015) for additional discussion. Some of the Conidae species from YN020 have also been


reported from other tropical American localities and strata. The new morphological descriptions below pertain only to material from YN020 and studied type material. Future work will more broadly consider intraspecific differences across the Neogene of tropical America.


Repositories and institutional abbreviations.—All newly collected specimens from YN020 are reposited in the Florida Museum of Natural History Division of Invertebrate Paleontology collections at the University of Florida (UF). Previously published fossils, including type and figured specimens, are from the following museum collections: the Academy of Natural Sciences of Drexel University, Philadelphia, Pennsylvania (ANSP); the California Academy of Sciences Department of Invertebrate Zoology and Geology, San Francisco (CASG); the Colección Nacional de Paleontología, Instituto de Geología, Universidad Nacional Autónoma de México, Mexico City (IGM); the Natural History Museum, London (NHMUK); the Naturhistorisches Museum Wien, Austria (NHMW); the Paleontological Research Institution, Ithaca, New York (PRI); the University of California Museum of Paleontology,Berkeley (UCMP); and the Smithsonian Institution National Museum of Natural History, Washington D.C. (USNM).


Systematic paleontology Family Conidae Fleming, 1822


Remarks.—While repeated tests have confirmed the monophyly of the Conidae (e.g., most recently by Uribe et al., 2017), the


809


internal classification of this hyperdiverse clade has been in a state of continuous upheaval in recent years, with significant disagreement about how to subdivide the clade into manageable Linnaean categories consistent with modern hypotheses of phylogeny (e.g., see varied views in Röckel et al., 1995; Tucker and Tenorio, 2009; Hendricks et al., 2014; Hendricks, 2015; Petuch et al., 2015; Puillandre et al., 2015; Harzhauser and Landau, 2016; Landau et al., 2016). While a suitable classifi- cation for extant cone snails is now available (Puillandre et al., 2014, 2015; Uribe et al., 2017), morphological features of their shells have yet to be coded and mapped upon the existing molecular trees. Because of this, detailed morphological diag- noses of individual clades supported by molecular sequence data remain largely lacking. For example, while molecular data suggest a great genetic divide between the two largest genera, Conasprella and Conus, morphological features of the shell present no obvious features that separate the two clades. Thus, detailed analyses of the phylogenetic distribution of cone snail shell characters in light of molecular sequence data are badly needed, most especially to diagnose the shell features of the different genera and subgenera of Conidae. The assignment below of fossil species of Conidae to individual clades, there- fore, is based largely on comparison with the shells—and especially the shell coloration patterns—of similar modern taxa of known phylogenetic position. Species with no obvious rela- tionship to modern taxa are assigned for now to Conus, reflecting the traditional classification of cone snails (see Röckel et al., 1995; Hendricks, 2009; Kohn, 2014).


Genus Conasprella Thiele, 1929


Type species.—Conasprella pagoda (Kiener, 1847) by sub- sequent designation (Tucker and Tenorio, 2009). Species is extant and occurs in the Indo-Pacific.


Remarks.—Based on molecular phylogenetic results, Puillandre et al. (2014, 2015) divided the extant clade Conasprella into seven subgenera, three of which (Kohniconus Tucker and Tenorio, 2009; Dalliconus Tucker and Tenorio, 2009; Ximeni- conus Emerson and Old, 1962) include tropical American members. All extant species of Conasprella are vermivorous (Puillandre et al., 2014), so this feeding ecology can also be assumed for fossil taxa assigned to this clade.


Conasprella imitator (Brown and Pilsbry, 1911) Figure 3.1–3.14


1911 Conus dalli Toula, p. 509, pl. 31, fig. 23a–d (not Conus dalli Stearns, 1873, an extant eastern Pacific species).


1911 Conus imitator Brown and Pilsbry, p. 342, pl. 23, fig. 4. 1917 ?Conus dalli; Maury, p. 212, pl. 7, fig. 15. 1921 Conus imitator; Pilsbry, p. 327. 1928 Conus imitator lius Woodring, p. 209, pl. 10, figs. 5, 6.


1970 Conus imitator imitator; Woodring, p. 354, pl. 55, figs. 1, 2.


2009 Gradiconus imitator (Brown and Pilsbry); Tucker and Tenorio, p. 97.


2010 Conus imitator; Landau and da Silva, p. 101, pl. 20, fig. 9a–c, pl. 21, fig. 1a, b.


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