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The fossil record of the Solanaceae revisited and revised--the fossil record of Rhamnaceae enhanced.

Introduction

It is ironic that despite the size of the family (Hunziker, 2001; Martins and Barkman, 2005; Olmstead et al., 1999; Olmstead et al., 2008; Teteny, 1987), the fossil record of Solanaceae is very sparse perhaps due to undersampling of the fossil record in the Neotropics where the greatest diversity is found today, and to the inherent preservational bias against herbaceous, non-deciduous plants. The paucity of fossils assignable to the Solanaceae has prevented the establishment of a reliably calibrated timeline for its diversification, but the importance of the family imposes great significance on the fossils that have been reported and the validity of their identifications becomes critical.

Selection of Fossil Taxa

Based on the revision of the fossil record of the Asteridae by Martinez-Millan (2010), eleven fossil taxa plausibly assigned to Solanaceae have been identified (Table 1). Of these eleven taxa, five are represented by flowers of the same fossil genus, Solanites, and four of these are also potentially the earliest (Early Eocene) fossil members of the family. Due to available numbers of characters and their diagnostic values and relative ages, the fossil taxa Solanites saportanus, S. sarachaformis, S. crassus, S. pusillus were selected for further investigation along with a fruit fragment with seeds from the London Clay (Cantisolanum daturoides). Cantisolanum was included because the original description of the latter fossil was by reliable investigators (Reid & Chandler, 1933), and because its remains were reproductive in nature and likely to include diagnostic characters.

The protologues of these fossil taxa were revised and the type specimens located in their housing institutions. In the case of the four [presumably] Early Eocene species of Solanites, it was necessary to not only locate their type specimens, but also to locate the type specimen of type species for the genus, Solanites brongniartii even though it is from the Oligocene, in order to evaluate their identifications. On the other hand, Cantisolanum is a monotypic genus known from only one locality, therefore, only one type specimen needed to be located.

Direct Examination of Fossil Taxa

The type specimens of the five species of Solanites were observed in their housing institutions: The Museum National d'Histoire Naturelle (MNHN) in Paris, France for S. brongniartii and the Smithsonian Institution National Museum of Natural History (USNM) in Washington, DC for the remaining species. Some additional specimens from North American localities were obtained on loan from the Florida Museum of Natural History (FMNH) and the University of Connecticut (UCPC). The specimens were studied by direct observation using mainly stereoscopic microscopes at the corresponding institutions and at the Paleobotanical Laboratory, Department of Plant Biology, Cornell University. Anthers or portions of anthers of four specimens (UF15737-49566, UF15737-49567, UF15738-8214 and UCPC-P10) were removed from the fossil, mounted on an SEM stubs, sputter coated with gold/palladium and observed under a field emission Hitachi 4500 Scanning Electron Microscope at the Cornell Center for Materials Research. The stubs are kept at the Cornell University Paleobotanical Collection (CUPC-1554 to CUPC-1561). Digital photographs were taken of all specimens. Appropriate archived photos from the Paleobotanical Laboratory including those of pollen grains taken on a compound microscope were scanned in to include in this study.

The type specimen of Cantisolanum daturoides was observed in its housing institution, the Natural History Museum in London, UK.

Initial examination of these fossils based on evident morphological characters a priori (due to missing or conflicting characters), precluded definite affinities with Solanales for one subset (Cantisolanum and certain Solanites species--as discussed below in section "Discussion"), suggested the possibility of Solanaceous affinities pending further analysis for another (those associated with Solanites brongniartii--Sections Cladistic Analysis and Methods for S. brongniartii and S. pusillus, Results of Cladistic Analyses), and suggested that yet another appeared to have affinities outside of the Solanaceae (those associated with Solanites pusillus--Sections Cladistic Analysis and Methods for S. brongniartii and S. pusillus, Results of Cladistic Analyses).. For, two of the fossil species {S. brongniartii and S. pusillus), phylogenetic analyses were warranted based on an adequate number of preserved characters. Appropriate matrices were constructed for these that included morphological and molecular characters as discussed and enumerated below in section "Cladistic Analysis and Methods for S. brongniartii and S. pusillus".

Historical Perspective, Nomenclature and Descriptions of Taxa

European Solanaceae

Solanites

Nomenclatural History. In 1855, Oswald Heer, then Director of the Zurich Botanical Garden published the first part of Flora Tertiaria Helvetiae, "The Tertiary Flora of Switzerland". The third part of the series, published in 1859, included an appendix titled Ueber das Klima und die Vegetationsverhaltnisse des Tertiarlandes, "On the climate and vegetation conditions of the Tertiary lands" (Heer, 1859a, b). This appendix compared the different Tertiary floras known at the time, especially those of Europe: Switzerland, Italy, Austria, Germany, Greece, Hungary, Romania, Russia, France, England and Iceland.

In 1861, Charles Th. Gaudin translated this appendix from German to French and published it as a separate book under the title Recherches sur le climat et la Vegetation du Pays Tertiaire, "Investigations on the climate and the vegetation of the Tertiary" (Heer, 1861). In this version, the original 3-page-long section on France was replaced with some excerpts of the original text followed by a 55-page-long treatise on the Tertiary floras of Provence, written by the Count Louis Charles Joseph Gaston de Saporta and titled Examen analytique des flores tertiaries de Provence, precede d'une notice geologique etpaleontologique sur les terrains tertiaires lacustres de cette region par M. Phil. Matheron, "Analytical exam of the Tertiary floras of Provence, preceded by a geological and paleontological note on the lacustrine terrains of that region". In this treatise, Saporta provided an overview of the fossil flora of Aix-en-Provence and, on page 146, he discussed some flowers pointing out in particular, "[One] specimen, in perfect state of conservation" and naming them Solanum brongniarti Sap [sic] after Brongniart suggested that "... [valvate aestivation] along with anther structure ... is an indication of the probable assignation of these flowers to the Solanaceae" (de Saporta, 1861).

The following year, 1862, Saporta started a series of papers in the Annales des Sciences Naturelles under the title Etudes sur la vegetation du sud-est de la France a Tepoque Tertiaire, "Studies on the vegetation of Southeast France in the Tertiary period" systematically describing the fossils of Aix-en-Provence and the surrounding areas. In the second installment of this treatment (Ann. Sci. Nat. 4th series, vol. 17, page 262), Saporta formally described the fossil giving it the name of Solanites brongniartii and clearly referencing the 1861 treatise. In that publication, he mentioned the existence of "at least" three specimens although only two were figured, with the second one cited as just "another corolla" (Fig. 1). The third specimen was, at a later date (de Saporta, 1873), recognized as the counterpart of the one upon which both descriptions were based.

In 1863, the first three installments were bound and published together as Volume 1 of a series of books bearing the same title. The fact that the description of S. brongniartii occupies page 109 in this compilation book has caused some confusion regarding the correct citation of the name. Therefore, in accordance to the ICBN (Vienna Code; McNeill et al., 2006), the correct name and citation for the genus should be Solanites Sap 1862 and type species Solanites brongniartii (Sap) Sap 1862 in Ann. Sci. Nat. Bot. ser. 4. 17: 262 with holotype MNHN-14215b.

Generic Diagnosis. As given by de Saporta (1862): Corolla gamopetala, pentamera, rotata, cestivatione valvata, caduca. Stamina 5 corollce fauci inserta, incumbentia, antheris 2-locularibus in processum apiculatum superne coalitis, longitudinaliter dehiscentibus.

Generic Description. Isolated flowers, presumably bisexual, actinomorphic. Calyx unknown. Corolla pentamerous, gamopetalous and caducous, with valvate aestivation, presumably rotate and reflexed. Androecium of five stamens alternate to the corolla lobes with short filaments and elongated anthers. Anthers tapering distally and ending in a connective projection.

Diagnosis of S. brongniartii. As given by de Saporta (1862): S. corolla quinquefida, lobis acuminatis, staminibus exsertis, incumbentibus, filamentis brevibus, antheris 2-locularibus fusiformibus, in processum tenuissime apiculatum desinentibus.

Description of S. brongniartii. S. brongniartii is known from two isolated pentamerous flowers. One of the specimens represent an isolated corolla (MNHN-14215a, b), while the second specimen appears to be a complete flower (MNHN-14223). The calyx is unknown. The corolla is membranaceous with basally fused and apically free petals, rotate, between 7.9 and 8.9 mm (mean 8.4 mm) in diameter. The free portion of the petals is between 1.7 and 2.4 mm wide (mean 2.0 mm) and between 3.0 and 3.5 mm long (mean 3.3 mm), the total length of the corolla is unknown. In the holotype, MNHN-14215b, the apices of three petals are folded over suggesting that in life the corolla was reflexed, probably tortuous (Fig. 1a-c). In the second specimen, MNHN-14223, the margins of the petals are broken and folded inward also suggesting a non-flat, reflexed corolla (Fig. Id). Each petal has a conspicuous but not prominent midvein and several secondary veins parallel to the midvein that dichotomize towards the margin (Fig. 1e). The androecium is composed of five stamens that alternate with the petals. The filaments are short (Fig. 1 g) and presumably epipetalous as they remained attached to the shed corolla (Fig. 1a-c). The anthers are between 3.0 and 3.7 mm long (mean 3.3 mm) and between 0.3 and 0.4 mm wide (mean 0.4 mm). The anthers have striated walls and end in a projection of the connective (Fig. If). One of the specimens (MNHN-14223) shows what could be interpreted as the imprint of a two-carpelled gynoecium with separate styles and capitate stigmas (Fig. 2). No fruits or vegetative structures had been associated with these flowers.

Locality. Laminated marly shales of the lower part, Aix-en-Provence Formation.

Age. When S. brongniartii was described, it was assigned a Late Eocene age by de Saporta (1862, 1886). In the earlier part of the 20th century, the flora of Aix-en-Provence was cited as Sannoisien (i.e. Berry, 1916) and/or Aquitanien (i.e. specimen labels in MNHN), which corresponded to the Oligocene-Lower Miocene of North America (Osbom, 1907). Today, with the standardization of the Geological Time Scale, the name "Aquitanien" applies to the earliest stage of the Miocene and not to the Oligocene (Gradstein et al., 2004). New studies on the Aix-en-Provence Fm confirm that the age of the flora is Late Oligocene, Chattian (Chateauneuf & Nury, 1995; Kvacek & Erdei, 2001).

Cantisolanum daturoides

The observation of the only specimen known of this taxon reveals that the fossil is fragmentary and does not have good structural data preserved. In addition, by comparing the original photographs with the fossil in its current state, it can be concluded that a significant amount of pyrite decay has occurred since its original description. The dissection of the fossil and the decay could have obscured or eliminated some features that the original authors observed in the fossil but that today are not evident. For example, the three abortive seeds mentioned in the protologue (Reid & Chandler, 1933) are nowhere to be found today. The lack of diagnostic characters precludes the assignment of this fossil to the Solanaceae or any other higher taxon (Collinson, 1983). (Since this isn't North American, it should go near European Sloanites--but not really sure where it goes.)

North American Solanites

History and Localities

In 1916, Berry described one specimen bearing two flowers as a second species of Solanites naming it S. saportana, a name that ought to be corrected to S. saportanus in order to agree in gender. The specimen was collected from the Holly Springs Sand, Claiborne Formation of Mississippi assigned at the time to the Wilcox Formation. In his original description Berry stated that "... the Wilcox flower described above ... is very similar to Solanites brongniarti, although less completely preserved". Berry (1916) even reproduced some of the figures that Saporta used in his description of S. brongniartii, including those of the extant Saracha and Wintheringia. However, a closer look at the descriptions and the type specimens shows that the only resemblance between S. saportanus and S. brongniartii is in their common pentamery.

Three more species from the Claiborne Formation, were added by Berry in 1930; S. sarrachaformis--to be corrected to S. sarachaformis since the correct name for the extant genus is Saracha and not Sarracha--, S. crassus and S. pusillus. These three species are also based on pentamerous flowers, again with little else in common with Solanites.

Description of S. saportanus

The type specimen of this taxon includes two flowers, one in side view and one in bottom view suggesting that these were close together in life, possibly on the same inflorescence axis (Fig. 3a, b). The flowers are small (5.8 mm diameter) with a shallow floral cup. The calyx is thick, pentamerous, gamosepalous with small rounded to slightly acuminate calyx lobes that show traces of a midvein. The calyx is 2.6 mm in diameter, with each calyx lobe (sepal) measuring between 1.2 and 1.5 mm wide (mean 1.3 mm) and between 0.5 and 0.8 mm (mean 0.65 mm) long. The corolla is also pentamerous with small, coriaceous, concave, acuminate petals whose free portions measure between 1.6 and 2.2 mm (mean 1.42 mm) wide and between 2.3 and 2.7 mm (mean 2.5 mm) long. It is unclear if the petals were completely free, born on the rim of a hypanthium (implying adnation of calyx and corolla), or if they were fused at the bases and free at the tips, where the separation would occur at a level obscured by the calyx. The corolla seems to have been opposite the calyx, with each petal directly in front of a sepal. Berry (1916) reported the existence of stamens in this specimen, however, it was not possible to find evidence of them upon direct examination. There are no traces of a gynoecium. A second specimen (Table 2) does not show additional characters. The holotype of this species is USNM-35990.

Description of S. pusillus

This taxon was described from three specimens, three syntypes (Table 3). One of them represents a flower in face (top) view (Fig. 3c, d), one is a flower in bottom view (Fig. 3e, f) and the third one is a flower in side view (Fig. 3g, h). These flowers are small, from 7.2 to 9.9 mm in diameter (mean 8.7 mm), and have always been found isolated. The flowers are pentamerous, with a shallow, thick, well developed hypanthium, semicircular (U-shape) to cupulate in shape (Fig. 4a). In top view, the rim of the floral cup is very thick, ring-shaped (Figs. 3c, d and 4d-f), sometimes slightly lobed and interpreted as a nectary ring that measures between 1.6 and 4.2 mm in diameter (mean 3.4 mm). The five appendages that compose the perianth cannot be defined as sepals or petals in the type specimens (Fig. 3c h), but additional specimens (Table 3) with both whorls preserved indicate that they represent sepals (Fig. 4c-f). These sepals are triangular in shape, between 1.5 and 2.8 mm wide (mean 2.14 mm), and between 1.5 mm and 3.5 mm long (mean 2.6 mm), with a prominent midvein or keel sometimes with additional smaller ridges at either side (Fig. 3c, d, g, h). In addition, they are covered by abundant filiform hairs (Fig. 4b). The petals alternate with the sepals (Fig. 4c-f), they are membranaceous with some faint veins, small, cucullate ("hooded") and possibly clawed, between 3.2 and 3.4 mm in length (mean 3.3 mm) and 0.9 and 1.5 mm (mean 1.1 mm) in width. There are five stamens, opposite the petals (Fig. 4e, f), each stamen is about 3.4 mm in length with a 0.4 mm long tetrasporangiate anther. The pollen grains appear to be tricolporate (Fig. 5a, b) with striate-rugulate exine (Fig. 5c, d), they are between 16 and 21 p.m in length. In most specimens, the gynoecium is not discernible but in the type specimen in side view, it appears to be superior with one relatively thick style and one stigma (Fig. 3g, h) which would make the flower, perigynous. This species has three syntypes housed at the Smithsonian Institution with numbers USNM-222831, USNM-222832, and USNM-39950.

Description of S. sarachaformis

The type specimen of this taxon is an impression of a flower with no retained organic matter (Fig. 6a, b). The flower measures about 1 cm in diameter. It is pentamerous with a shallow floral cup (Fig. 6a, b). The the perianth segments are fused towards the bases and free towards the tips, they are 3 mm in width and 2.6 mm long. It is not clear if they represent calyx lobes or corolla lobes. There are no traces of stamens or of a gynoecium. Additional specimens comparable to this fossil are in a similarly bad state of preservation (Table 4). The type for this species is USNM-35948.

Description of S. crassus

This taxon is represented only by three syntypes and the three are poorly preserved (Table 5). Although there is still some original matter left, there is almost no structural detail. The impressions seem to be of a membranaceous corolla with the petals fused throughout their length. They are between 8.1 and 16.5 mm (mean 11.7 mm) in diameter. Two of the specimens are pentagonal in outline (Fig. 6c, d, g, h) while the third is circular (Fig. 6e, f).

Age

The fossils assigned the different Solanites species were originally assigned to the Lower Eocene Wilcox Formation (Berry, 1916, 1930). However, after thorough palynological studies, these sediments are assigned today to the Middle Eocene Claiborne Formation (Dilcher, 1971; Potter, 1976).

Cladistic Analysis and Methods for S. brongniartii and S. pusillus

Analysis with S. brongniartii

Because S. brongniartii appeared to have morphological features consistent with Solanaceae and in order to test this initial impression, an appropriate combined morphological and molecular matrix was compiled. For the molecular data, the six cpDNA sequences used by Bremer et al. (2002)-rbcL, ndhF, matK, rpsl6, trnT-trnF, and trnV-were downloaded from GenBank, aligned with CLUSTAL X (Thompson et al., 1997) using a gap-opening penalty of 20.00 and a gap-extension penalty of 5.00, and manually adjusted afterward. A few corrections had to be made to the downloaded data before the alignment could be completed: (1) accession number AJ429683 (trnV) identified in GenBank with the name Dipentodon sinicus was changed to Sanango sp. in accordance with the table provided by Bremer et al. (2002), (2) the two accession numbers AF130223 and AJ238344 reported in Bremer et al. (2002) as rbcL sequences were moved to the ndhF matrix as they correspond to that gene, (3) the inverse (reversed complement) sequence for accession number Z00044 (Nicotiana tabacum chloroplast genome) was used for the rpsl6 sequence, (4) accession number U73971 for Orobanche racemosa was not used because Bremer et al. (2002) indicated it represents a pseudogene. After alignment, 5 leading positions from matK, 22 from ndhF, 22 trailing positions from matK, and 50 from trnT-trnF were removed as they were present in only one or two taxa. Conversely, 272 trailing positions from ndhF were trimmed because they were not confidently aligned. The rbcL, rpsl6 and trnV matrices were not modified. Each of the six genes was subjected to the simple indel-coding method of Simmons and Ochoterena (2000) as implemented in the program GapCoder (Young & Healy, 2003). These six marker and six indel matrices were combined into a "molecular only" matrix of 132 taxa and 15147 characters (6899 informative) which is, in principle, equivalent to the Bremer et al. (2002) matrix. This matrix was analyzed to corroborate the Bremer et al. (2002) results using the same parameters and search strategies as for the combined "total evidence" matrix (see below).

A morphological matrix (Table 6) based on the asterid taxa used by Bremer et al. (2002) was constructed using family descriptions by Cronquist (1981) and Takhtajan (1997) and digital photographs available at www.plantsystematics.org/. The matrix has 111 taxa and 23 characters:

1. Flower sexuality: bisexual = 0; unisexual = 1.

2. Flower size (considered as corolla diameter): less than 1 cm = 0; more than 1 cm = 1.

3. Corolla symmetry: actinomorphic = 0; zygomorphic = 1.

4. Corolla shape [additive]: narrow: salverform/tubular = 0; widening: campanulate/ urceolate/infundibiliform = 1; open: rotate (flat)/reflexed/lobed = 2.

5. Corolla aestivation: valvate = 0; convolute/contorte = 1; imbricate/quincuncial = 2.

6. Corolla merosity: trimerous = 0; tetramerous = 1; pentamerous = 2; hexamerous = 3; octamerous = 4.

7. Corolla fusion [additive]: free petals (polypetalous) = 0; fused at the base (basally gamopetalous) = 1; fused with lobes (gamopetalous) = 2; fully fused, no lobes (gamopetalous) = 3; fused distally (calyptra) = 4.

8. Corolla texture: membranaceous = 0; coriaceous = 1.

9. Corolla persistence: persistent = 0; caducous/deciduous = 1.

10. Petal midvein: absent = 0; present = 1.

11. Apices of petals: straight = 0; tortuous = 1; involute = 2; revolute = 3.

12. Stamen number relative to petals: haplostemonous = 0; diplostemonous = 1; anisomerous (less than petals) = 2; numerous = 3.

13. Stamen cycle orientation: alternate with petals (altemipetalous) = 0; opposite to petals (antepetalous) = 1; centrifugal/centripetal = 2.

14. Stamen attachment: to gynoecium = 0; to petals (epipetalous) = 1; to receptacle = 2.

15. Filament vs. anther: short = 0; long = 1.

16. Anther surface: striated = 0.

17. Connective projection: absent = 0; present = 1.

18. Anther dehiscence: longitudinal slits = 0; poricidal = 1.

19. Stamens vs. corolla lobes (non flat corolla): exserted = 0; included = 1.

20. Carpel number: 2 = 1; 3 = 2; 4 = 3; 5 = 4; more than 5 = 5.

21. Ovary: apocarpous = 0; syncarpous = 1.

22. Number of styles: 0 = 0; 1 = 1; 2 = 2; 3 = 3; 4 = 4; 5 = 5; more than 5 = 6.

23. Stigma shape: simple = 0; capitate = 1; clavate = 2; lobed (carpel number) = 3; lingulate = 4.

The final, total evidence matrix has 133 taxa and 15,170 characters of which 6920 are informative (Table 7). Four different sets of analyses were performed based on this matrix. The first, "total evidence", includes all the characters and taxa. The matrix was analyzed 10 times using TNT (Goloboff et al., 2003, 2008). Each analysis consisted of 1000 replications of subtree pruning and regrafting (SPR) on randomly generated Wagner trees holding up to 10 trees per replication, followed by a round of tree bisection-reconnection (TBR) (rseed 0; mult = spr replic 1000 hold 10; bbreak = tbr). Standard bootstrap values were calculated on 1000 TBR replications of 5 runs each, holding up to 5 trees per replication and keeping the consensus only.

The second, "without gynoecium characters" uses the same matrix and the same parameters as the first analysis but the character state for characters 20 (carpel number), 22 (style number), and 23 (stigma shape) in Solanites brongniartii has been changed to "?" in order to test one interpretation of the gynoecium in the fossil as being bistylar.

The third, "without fossil", excludes the taxon Solanites brongniartii. Its objective is to determine the effect of the fossil in the strict consensus. This matrix has 132 taxa and 15,170 characters of which 6,920 are informative. The analysis used the same parameters as the "total evidence" analysis.

The fourth, "character-by-character", used the total evidence matrix with one morphological character turned off. Two morphological characters, 16-anther surface and 21-ovary, were excluded from all the analyses because they are uninformative. The remaining 21 characters were turned off one-at-a-time and one run of 1000 SPR replications holding up to 10 trees per replication followed by a round of TBR (rseed 0; mult = spr rep lie 1000 hold 10; bbreak = tbr) was performed in each case.

The resulting trees were evaluated in Winclada version 1.00.08 (Nixon, 2002) where strict consensuses were constructed for each analysis.

Analysis with S. pusillus

Based again on an a priori assessment of available morphological characters, S. pusillus seemed a poor fit with Solanaceae (Table 8). As indicated by the fossil's major morphological characteristics, a different and appropriate data matrix consistent with the fossil's distribution of characters was constructed using the aligned data matrix of Richardson et al. (2000) downloaded from the American Journal of Botany supplementary data site (http://ajbsupp.botany.org/v87/)-This matrix includes 66 taxa (14 outgroups) and 2807 characters: 1428 rbcL, 1363 trnL and 16 representing trnL indels.

For the morphological matrix, that of Calvillo-Canadell (2000) which has 18 taxa and 26 characters was selected and obtained from the author (Table 8). This matrix includes two outgroups, Vitis (Vitaceae) and Mortonia (Celastraceae) and two fossil flowers assigned to Rhamnaceae, Nahinda axamilpensis from the Oligocene and Coahuilanthus belindae from the Cretaceous (Calvillo-Canadell & Cevallos-Ferriz, 2007). The characters are:

1. Petal width (mm): 0.15 = 0; 0.30-0.60 = 1; 0.90+ = 2.

2. Petal length (mm): 0.50-0.90 = 1; 1.00-1.80 = 2; 3+ = 3.

3. Sepal width (mm): 0.30-0.70 = 1; 0.90-1.50 = 2; 2+ = 3.

4. Sepal length (mm): 0.70-1.10 = 0; 1.20-1.40= 1; 1.50-1.80 = 2; 2+= 3.

5. Ring diameter (mm) (insertion of other floral parts area): 0.70-1.10 = 0; 1.20-1.50= 1; 2.00-2.10 = 2; 3+ = 3.

6. Pedicel length (mm): 0.30-1.70 = 0; 2.00-3.00 = 1; 3.50-5.00 = 2.

7. Ovary length: 0.20-0.30 = 0; 0.40-0.60 = 1.

8. Flower length (mm): 1.50-3.90 = 0; 4.00-4.80 = 1; 5.00-6.81 = 2.

9. Claw in petal: absent = 0; short = 1; long = 2.

10. Petal shape: cucullate = 0; obovate = 1; ovate concave = 2; linear= 3; urceolate = 4; valvate = 5; imbricate = 6.

11. Petal apex: no emarginate = 0; emarginate = 1.

12. Sepal shape: triangular = 0; triangular deltate = 1; triangular reflexed = 2; triangular lobed = 3; triangular acute = 4; triangular ovate = 5; triangular indexed = 6.

13. Keel: absent = 0; not prominent = 1; prominent = 2.

14. Enlarged sepal apex: absent = 0; present = 1.

15. Fruit type: datpe = 0; unilocular capsule = 1; trilocular capsule = 2; schizocarp = 3; berry = 4; samara = 5.

16. Fruit shape: globose = 0; subglobose = 1; obovoid = 2; elipsoid = 3; ovoid = 4.

17. Pyrene number: none = 0; two pyrene = 2; three pyrene = 3.

18. Locule number: two = 2; three = 3.

19. Petal presence: always present = 1; frequently present = 2; rarely present = 3.

20. Ovary type: inferior = 0; superior = 1; seminferior = 2; semisuperior = 3.

21. Floral cup shape: hemisphaeric = 0; campanulate = 1; obconic = 2; patelliform = 3; urceolate = 4.

22. Flower sex: perfect = 0; imperfect = 1; perfect and poligamous = 2.

23. Stamen size: longer than petals = 0; shorter than petals = 1; shorter than sepals = 2.

24. Floral part number: 4 or 6 = 0; 5 = 1; 4 or 5 = 2; 3 to 5 = 3.

25. Corolla vs. calyx: same size = 0; corolla bigger than calyx = 1; corolla smaller than calyx = 2.

26. Winged fruit: not winged = 0; winged = 1.

The molecular matrix of Richardson et al. (2000) was reduced by fusing species of the same genus into one single taxon and deleting genera not present in the morphological matrix. In addition, five sequences were added to the matrix: (1) trnL sequence for Adolphia infesta (AY460408, Aagesen et al., 2005), (2) trnL sequence for Vitis vinifera (EF179097, Rossetto et al., 2007), (3) rbcL sequence for V. aestivalis (L01960, Albert et al., 1992), (4) trnL sequence for Mortonia greggii (DQ217437, Islam et al., 2006), and (5) rbcL sequence for M. greggii (AY935727, Zhang & Simmons, 2006). The addition of the five sequences and the reduction in the number of taxa resulted in an alignment adjustment with a consequent change in the trnL indel presence/absence matrix. The new trnL indel matrix was reconstructed using the program GapCoder (Young & Healy, 2003). The final matrix has 19 taxa and 2751 characters (Table 9).

The matrix was analyzed using implicit enumeration on TNT (Goloboff et al., 2003, 2008). The resulting trees were evaluated in Winclada version 1.00.08 (Nixon, 2002) where the strict consensus was constructed. Standard bootstrap values were calculated on 1000 implicit enumeration replications on TNT keeping the consensus only.

Results of Cladistic Analyses

Analysis with S. brongniartii

The phylogenetic analysis of the Bremer et al. (2002) molecular matrix yielded two trees. Although they show the same general backbone as the Bremer et al. (2002) strict consensus of 24 trees, there are some differences (not shown). The first is in the relationship of the outgroups; in our analysis Vitis and Dipentodon are not sister groups, instead Vitis is sister to the Asteridae. Within the Asteridae, the four clades, Comales, Ericales, Lamiids and Campanulids are each monophyletic and have the same relationships as in the Bremer et al. (2002) consensus tree: (Cornales(Ericales(Lamiids, Campanulids))). Within Ericales and Campanulids, the basal polytomies shown in Bremer et al. (2002) are resolved in our analysis. And in the Lamiids, some relationships among major clades have changed, for example, in our analysis, the Icacinaceae resolves as monophyletic with Oncotheca moving to a position as sister to the Gariyales instead of a clade Oncotheca-Apodytes and a second clade of the rest of the Icacinaceae as in Bremer et al. (2002). Another major difference is the relationship of the Lamiid clades that in Bremer et al. (2002) is (Gentianales (Vahliaceae Boraginaceae) (Lamiales Solanales including Convolvulaceae)) but in our analysis is ((Convolvulaceae Boraginaceae) Lamiales) (Vahliaceae (Solanales Gentianales)). Except for the position of Convolvulaceae, the differences pertain to families unplaced to order and to the relationships among orders, not to membership to those orders.

The analysis of the total evidence matrix resulted in 83 most parsimonious trees (MPTs). The strict consensus of these trees (Fig. 7) shows that the fossil taxon, Solanites brongniartii, is found in a polytomy in the first node of the Euasteridae along with eight individual terminals and nine clades. This placement is supported by a 90% bootstrap value but it is also suggestive of the "floating" nature of the fossil taxon. In the strict consensus a change in character 14, attachment of the stamens, from attachment to the receptacle (free stamens) to attachment to the petals (epipetaly) supports this group; however, optimization in each of the MPTs does not result in this change at this node and in fact, no morphological character defines this group.

A closer look to the 83 MPTs reveal that Solanites brongniartii floated between only six different positions (Fig. 8). These are: (1) as sister to Vahlia in 13 trees (Fig. 8a) supported by a change from a widening to an open corolla (ch. 4) and a change from one to two styles (ch. 22), (2) as sister to Oncotheca in 8 trees (Fig. 8b) supported by a short filament (ch. 15) and the presence of a connective projection (ch 17), (3) in a polytomy with Oncotheca and Apodytes in 5 trees (Fig. 8c) supported by the same characters as in the previous position as Apodytes is not present in the morphological matrix, (4) nested within Aquifoliales as sister to a clade composed of Helwingia and Phyllonoma in 19 trees (Fig. 8d), this position is supported by the membranaceous texture of the corolla (ch. 8) shared with Phyllonoma (ambiguous for Helwingia) and the petal midvein (ch. 10) shared with Helwingia (ambiguous for Phyllonoma), (5) nested within Solanales as sister to the Montiniaceae (Fig. 8e) in 19 trees supported by a change from widening to open corolla (ch. 4), and (6) as sister to Olea in 19 trees (Fig. 8f) supported by a short filament (ch. 15) and a projection of the connective (ch. 17).

The analysis "without gynoecium characters" resulted in 27 MPTs (not shown). These trees are a subset of the 83 trees found when S. brongniartii is interpreted as having a bicarpellate gynoecium with separate styles and capitate stigmas. In these trees, S. brongniartii floated between two positions: as sister to Olea in 19 trees and as sister to Oncotheca in 8 trees. In both cases, the relationships are supported by a short filament (ch. 15) and the presence of a connective projection (ch 17).

The analysis of the combined matrix without the fossil resulted in 19 MPTs (L = 56322, CI = 32, RI = 48). As expected, the strict consensus (not shown) is more resolved than in the analysis including the fossil. However, polytomies abound. For example, the Asterales/Apiales/Dipsacales/Quintinia-Paracryphia/Polyosma-Tribeles-EremosyneEscallonia polytomy (Fig. 7) is present in the consensus with or without the effect of the fossil. The same occurs with Apodytes, Oncotheca, Icacinaceae (remaining 3 taxa) and Garryales, their relationships remain uncertain when the fossil is excluded. When the fossil is not included, the higher-nested lamiid groups form a monophyletic group with a basal polytomy of the structure: VahlialLamiales (31 taxa)/Solanales-Gentianales/ Convolvulaceae-Boraginaceae. With the fossil, this monophyletic group collapses and so does the Solanales, but the Lamiales only lose the first two diverging taxa, the remaining 29 still form a clade. In summary, the addition of the fossil resulted in only about 5 significant node collapses in the strict consensus.

The "character by character" analysis revealed that turning off ten of the characters had no impact on the results as the MPTs obtained were the same obtained when all characters are included (Table 10). In most of the other cases, removal of one character resulted in a subset of the 83 MPTs obtained when all characters were included. The only two characters whose removal resulted in new trees are corolla fusion (ch. 7) and stamen attachment (ch.14). The analysis with stamen attachment removed (ch. 14) is also the only in which a larger set of trees was found (Table 10).

Analysis with S. pusillus

The phylogenetic analysis of the combined matrix resulted in two trees whose only difference is the position of the fossil taxon Solanites pusillus as either sister to Condalia or to Krugiodendon (Fig. 9). The backbone of the cladogram is more similar to the rbcL cladogram of Richardson et al. (2000) than to their trnL or combined rbcL/ trnL tree. In the analysis the Rhamnaceae is resolved as monophyletic with a bootstrap value of 87, although within the Rhamnaceae the support for the different clades is generally low. The two fossils described by Calvillo-Canadell and Cevallos-Ferriz (2007), Nahinda axamilpensis and Coahuilanthus belindae are found well nested in different clades, as sisters to Colubrina and Sageretia respectively.

Discussion

The study of the type specimens of the fossil species once assigned to Solanaceae has shed important light into the identities of these fossils. One of the most important conclusions is that the five fossil species of the genus Solanites do not belong to the same or even related taxa. For example, S. pusillus and S. saportana do not have a gamopetalous corolla, one of the diagnostic characters of the genus (Fig. 3). The corolla is gamopetalous in S. crassus, but it is not rotate and it does not have corolla lobes (Fig. 6C-H). In S. sarachaformis, it is not clear if the impression represents a calyx or a floral cup, but it is not rotate (Fig. 6A). In the case of the stamens, they have short filaments and long fusiform anthers in the type (Fig. 1) while in S. pusillus, the only other species with known anthers, they have long filaments and globose anthers (Fig. 4e, f). Not only does each of the North American species differ significantly from the type species, they also differ from one another indicating that none of the five species belong in the same genus.

Affinities of S. brongniartii

In gross morphology, S. brongniartii does show some solanaceous characters such as the pentamerous rotate corolla with reflexed corolla lobes (petals) and the stamens with long anthers and short filaments. (Fig. 1). These characters are typical of the genus Solanum itself and they are so conspicuous in the fossil that it was originally assigned to the genus Solanum (de Saporta, 1861). However a closer look reveals that there are structural differences which result in the fossil not fitting completely into the genus. For example, the anthers in the genus Solanum typically have poricidal dehiscence, in the fossil there is no evidence of such pores, instead, the apices of the anthers end in a connective projection. Longitudinal dehiscence is found elsewhere in the Solanaceae, including in some species of Solanum (Carrizo Garcia et al., 2008), but in these instances the anthers lack a connective projection.

Another character present in S. brongniartii, under one interpretation, is a two styled and two stigma-bearing gynoecium, but this condition is not found in the Solanaceae. In the Solanaceae, the ovary is typically composed of two completely fused carpels, with one style and one stigma. The character "two carpel, two style", if present and not a preservation-based illusion, would call into question the assignment of S. brongniartii to the Solanaceae. However, when the gynoecium is perhaps more conservatively coded as "unknown" (the character states for three gynoecium characters are changed to "?"), S. brongniartii is also not placed close to the Solanaceae.

With respect to the two-styled-two stigma bearing interpretation of the fossil, the Apocynaceae is a family in which the combination divided styles, tortuous corolla and capitate stigmas occur, and all are possibly present in S. brongniartii. However, the fossil taxon does not demonstrate various specializations characteristic of that family either (appendages, outgrowths, etc.), especially in the stamens. The same could be said about possible ties to Boraginaceae, another family that the fossil taxon superficially resembles.

The phylogenetic analysis in which S. brongniartii was included, did not place the fossil in a defined position (Fig. 7). Instead it produced six different placements (Fig. 8). Interestingly, none of them related to the Apocynaceae or to the Boraginaceae. However one of the alternatives relates to the Solanales, not as sister to the Solanaceae but as sister to the Montiniaceae (Fig. 8e). This result highlights the mosaic nature of this fossil taxon that combines characters that today are found in different taxa.

One conclusion that can be drawn from the phylogenetic analyses is that S. brongniartii is a member of the Euasteridae (Fig. 7). Five of the six alternatives place the fossil taxon in the Lamiid clade while only one places it in the Campanulid clade, within the Aquifoliales (Fig. 8). A closer look to the characters that support the different placements reveal that they tend to be the same, for example, a change from a widening corolla (campanulate, infundibiliform) to an open corolla supports the Solanites-Montininaceae clade as well as the Solanites-Vahlia clade. The same occurs with the short filament and the connective projection; they support the clade Solanites-Oncotheca as well as the clade Solanites-Olea. This suggests that these characters are highly homoplasic and that the rest of the characters do not contribute significantly to the establishment of the relationships of Solanites brongniartii.

This idea is confirmed for at least ten characters as their removal did not change the number of trees obtained in the analysis or the positions of Solanites in those trees (Table 10). For nine other characters, removal resulted in a subset of the original trees (Table 10), suggesting once again that Solanites has a combination of characters that is not found in extant taxa. For these reasons, it is suggested that S. brongniartii be accepted as a bona fide euasterid. Many characters of S. brongniartii are still unknown; discovery of those character states can shed more light into its relationships but until more specimens are discovered, and more refined phylogenetic analyses are performed --for example by breaking down highly polymorphic taxa into more discrete units--the best placement for the fossil is as insertae sedis within the Euasteridae.

Affinities of Cantisolanum daturoides

The direct examination of the only specimen known of this taxon revealed that the fossil is now fragmentary without well preserved structural characters. In addition, by comparing the original photographs with the fossil in its current state, it can be concluded that a significant amount of pyrite decay has occurred since its original description. The dissection of the fossil and the decay may well have obscured or eliminated some features that the original authors observed in the fossil but that today are not evident. For example, the three abortive seeds mentioned in the protologue (Reid & Chandler, 1933) are nowhere to be found today. The lack of diagnostic characters precludes the assignment of this fossil to the Solanaceae or any other higher taxon (Collinson, 1983).

Affinities of North American Solanites

Affinities of Solanites crassus, sarachaformis, and saportanus

The study of the type specimens of these fossil species once assigned to Solanaceae has shed important light into the identities of these fossil taxa and revealed characters that are inconsistent with affinities with Solanaceae. In S. crassus the corolla is gamopetalous, but it is not rotate and it does not have corolla lobes (Fig. 6c-h). S. sarachaformis, has few characters, as only the outline of the flower is preserved with no other structural details and it is not possible to distinguish if the imprint is that of a calyx or a floral cup. But, judging from the shape and depth of the imprint in the sediment, it seems these perianth parts were somewhat fleshy, as a thin, delicate tissue would have left a flatter less topologically defined imprint. It is possible that S. sarachaformis represents the external mold of other taxa that, when better preserved could be identified as different Solanites species. The imprint, however, could not have been made by a S. brongniartii-like flower because the size and shape of the lobes is not consistent with the dimensions and form of S. brongniartii.

The remaining two species, S. saportanus and S. pusillus, show more structural details than the previous North American ones, allowing for more critical comparisons. In gross morphology and general view, these two species seem very similar, but a closer look reveals important differences. Both species have a shallow, thick, cupulate floral cup, but that of S. pusillus (affinities discussed in 2. separately below) clearly shows the insertion of the sepals at its rim, while in S. saportanus the sepals are continuous with the hypanthium with no clear attachment point. The most striking difference between these two species is, however, in the petals: in S. saportanus they are concave, coriaceous and are opposite to the sepals/calyx lobes, while in S. pusillus they are cucullate, presumably thin, and alternate with the sepals. This difference is not evident from the type specimens because in those of S. pusillus, the petals are not preserved. But even so, the sepal morphology alone can be used to distinguish between specimens of both fossil species. Sepals are triangular with a keel in S. pusillus as opposed to acute with no keel in S. saportanus. And in S. saportanus, there are too few characters preserved to reliably assess its affinities. However, the characters that are preserved (for example, S. saportana like S. pusillus, does not have a gamopetalous corolla, one of the diagnostic characters of the genus [Fig. 3]), illustrate that S. saportanus does not belong in the genus Solanites as this fossil does not fit the definition of that genus.

S. pusillus on the other hand, is relatively abundant in the flower collections from the Claiborne Formation, allowing for a better assessment of its identity.

Affinities of S. pusillus

The combination of characters present in this taxon: small pentamerous flowers with cupulate hypanthium ending in a thick presumably nectariferous rim, triangular sepals with a prominent midvein or keel, thin clawed cucullate petals alternating with the sepals, and stamens opposite to the petals is not found in the Solanaceae or even in the Asteridae, but in the Rhamnaceae, a family that is today classified with the Order Rosales in the Rosid clade (APG, 1998, 2003). The gross morphology of genera such as Colubrina, Scutia or Ziziphus (Fig. 4g-i) is very similar to that of S. pusillus. The pollen morphology of the fossil is also consistent with Rhamnaceae: tricolporate pollen with rugulate to striate-rugulate exine is present in Ziziphus (Nasri-Ayachi & Nabli, 1995), Paliurus (Schirarend, 1996), Hovenia (Zhang & Chen, 1992), and Sageretia (Perveen & Qaiser, 2005); but not in Rhamnus, Frungula (Punt et al., 2003), or Colubrina (Zhang & Chen, 1992).

The phylogenetic analysis conducted in this study yielded two trees, one with S. pusillus as sister to Condalia and as sister to Krugiodendron in the other, well nested in Tribe Rhamneae (Fig. 9). Flowever S. pusillus has a few characters that distinguish it from its putative closer relatives, for example its pubescence, a character not present in Krugiodendron or in Condalia (Medan et ah, 2004). Pubescence is not a common feature among Rhamnaceae but it can be found in other genera such as Adolphia, Discaria, Kentrorhamnus, Retanilla, or Trevoa (Medan & Aagesen, 1995). Only with more detailed studies, can the position of S. pusillus within the Rhamnaceae be more confidently established. Flowever, the recognition of S. pusillus as a member of the Rhamnaceae is a significant step towards a better understanding of the fossil history of that family. The abundance of specimens of this taxon (Table 3), the good degree of preservation and the previous observations made on this taxon (i.e. Crepet, 1974, 1979, 1984) allow for a more thorough study of the biology of this taxon, study that will be presented in a separate paper. Suffice to say that S. pusillus is a bona fide member of the Rhamnaceae and therefore not a member of the Solanaceae as was originally suggested (Berry, 1930). It is also noteworthy that leaves of Rhamnaceae have been reported from the same deposits by Jones and Dilcher (1980).

Conclusion

Of all the fossil species studied, only S. brongniartii and S. crassus can be considered to have possible, but not definite, affinities with the order Solanales, however not with the Solanaceae but with other families of the Order Solanales. Nonetheless, there are not sufficient characters in these taxa to support the possibility of affinities with the family Solanaceae. After careful revision of the types and in some cases additional specimens of fossil taxa once described as Solanaceae, the conclusion to reach is that none of the fossils studied here belong in the Solanaceae. One of the species, S. pusillus, was confidently assigned to the Rhamnaceae, and three more -S. sarachaformis, S. saportana and Cantisolanum daturoides- were shown to have too few characters preserved to confidently assign them to any family.

DOI 10.1007/s12229-014-9134-2

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Thompson, J. D., T. J. Gibson, F. Plewniak, F. Jeanmougin & D. G. Higgins. 1997. The CLUSTAL X windows interface: flexible strategies for multiple sequence alignment aided by quality analysis tools. Nucleic Acids Research. 25(24): 4876-4882.

Young, N. D. & J. Healy. 2003. GapCoder automates the use of indel characters in phylogenetic analysis. BMC Bioinformatics. 4: 6.

Zhang, Y.-L. & Y.-L. Chen. 1992. A study on pollen morphology of tribe Rhamneae (Rhamnaceae) in China. Acta Phytotaxonomica Sinica. 30(1): 73-81.

Zhang, L.-B. & M. P. Simmons. 2006. Phylogeny and delimitation of the Celastrales inferred from nuclear and plastid genes. Systematic Botany. 31(1): 122-137.

Marcela Millan (1), William Crepet (1,2)

(1) L. H. Bailey Hortorium, Department of Plant Biology, 412 Mann Library Building, Cornell University, Ithaca, NY 14853, USA

(2) Author for Correspondence; e-mail: wlc1@comell.edu

Published online: 16 May 2014

Table 1 Fossils assigned to Solanaceae

Fossil taxon                     Age             Locality

Flowers
  Solanites brongniartii         Oligocene       Aix-en-Provence,
    de Saporta, 1862                               France
  Solanites saportanus           Early Eocene    Claiborne, TN, USA
    Berry, 1916
  Solanites pusillus Berry,      Early Eocene    Claiborne, TN, USA
    1930
  Solanites sarachaformis        Early Eocene    Claiborne, TN, USA
    Berry, 1930
  Solanites crassus Berry,       Early Eocene    Claiborne, TN, USA
   1930
Fruits
  Cantisolanum daturoides        Early Eocene    London clay, England
    Reid & Chandler, 1933
  Physalis pliocaenica           Late Miocene    Stare Gliwice, Poland
    Szafer, 1947
Seeds                              (Tortonian)
  Solanispermum reniforme        Eocene          Lower Bagshot, UK
    Chandler, 1957
  Solanum arnense Chandler,      Eocene          Lower Bagshot, UK
   1962
Pollen
  Datura cf. D. discolor         Late Eocene     Florissant, CO, USA
    Leopold & Clay-Poole, 2001

Leaves
  Solandra haeliadum             Eocene          Salcedo, Italy
    Massalongo, 1851

Fossil taxon                     Reference

Flowers
  Solanites brongniartii         de Saporta, 1862
    de Saporta, 1862
  Solanites saportanus           Berry, 1916
    Berry, 1916
  Solanites pusillus Berry,      Berry, 1930
    1930
  Solanites sarachaformis        Berry, 1930
    Berry, 1930
  Solanites crassus Berry,       Berry, 1930
   1930
Fruits
  Cantisolanum daturoides        Reid &
    Reid & Chandler, 1933        Chandler, 1933
  Physalis pliocaenica           Szafer, 1961
    Szafer, 1947
Seeds
  Solanispermum reniforme        Chandler, 1962
    Chandler, 1957
  Solanum arnense Chandler,      Chandler, 1962
   1962
Pollen
  Datura cf. D. discolor         Leopold &
    Leopold & Clay-Poole, 2001   Clay-Poole,
                                 2001
Leaves
  Solandra haeliadum             Massalongo, 1851
    Massalongo, 1851

Table 2 Specimens identified as S. saportanus

Housing       Collector         Locality          Formation
institution

USNM          Berry             Holly Springs     Claiborne
FMNH          Dilcher et al.    Bovay Clay Pit    Claiborne

Housing       State    Specimen #
institution

USNM          MS       35990
FMNH          MS       UF8222/UF8222'

Table 3 Specimens identified as S. pusillus

Housing        Collector         Locality
institution

USNM           Berry             Holly Springs or La Grable (a)
USNM           Berry
USNM           Berry
FMNH           Dilcher et al.    Bolden Pit
FMNH           Dilcher et al.    Bovay Clay Pit
FMNH           Dilcher et al.    Bovay Clay Pit
FMNH           Dilcher et al.    Bovay Clay Pit
FMNH           Dilcher et al.    Bovay Clay Pit
FMNH           Dilcher et al.    Warman Clay Pit
UCPC           Crepet            Warman Clay Pit
UCPC           Crepet            Puryear
UCPC           Crepet            Puryear
UCPC           Crepet            Puryear
UCPC           Crepet            Puryear
IUPC           Crepet            Miller

Housing        Formation    State    Specimen #
institution

USNM           Claiborne    TN       222831
USNM           Claiborne    TN       222832
USNM           Claiborne    TN       39950
FMNH           Claiborne    MS       UF8214/UF8214'
FMNH           Claiborne    MS       UF8223/UF8223'
FMNH           Claiborne    MS       UF8226
FMNH           Claiborne    MS       UF49566/UF49566'
FMNH           Claiborne    MS       UF49568/UF49568'
FMNH           Claiborne    TN       UF33557/UF33557
UCPC           Claiborne    TN       UCPC (5)
UCPC           Claiborne    TN       UCPC "P5" (18)
UCPC           Claiborne    TN       UCPC "P" (3)
UCPC           Claiborne    TN       UCPC "P10" (22)
UCPC           Claiborne    TN       UCPC "P" (13)
IUPC           Claiborne    TN       IUPC M2203 (b)

(a) Berry (1930) did not identify the locality for each specimen, but
noted that only one specimen came from La Grable

(b) this specimen was examined via photographs archived in the
paleobotanical laboratory collection

Table 4 Specimens identified as S. sarachaformis

Housing institution    Collector         Locality          Formation

USNM                   Berry             Holly Springs     Claiborne
FMNH                   Dilcher et al.    Bovay Clay Pit    Claiborne

Housing institution    State    Specimen #

USNM                   TN       35948
FMNH                   MS       UF49569/UF49569'

Table 5 Specimens identified as S. crassus

Housing institution    Collector    Locality         Formation

USNM                   Berry        Holly Springs    Claiborne
USNM                   Berry        Holly Springs    Claiborne
USNM                   Berry        Holly Springs    Claiborne

Housing institution    State    Specimen #

USNM                   TN       39949a
USNM                   TN       39949b
USNM                   TN       39949c

Table 6 Matrix of morphological characters including the fossil taxon,
Solanites brongniartii. A-[234], B-[12345], C-[23456], D-[01], E-
[12], F-[02], G-[2345], H-[345], I-[124], J-[123], K-[012], L-[013],
M-[13], N-[01234], O-[014], P-[123456], Q-[34], R-[03], S-[23], T-
[45], U-[1234], V-[0123], W-[14], X-[24], Y-[15]

Character                                       1

Paeoniaceae Paeonia                             0    1    0    2
Vitaceae Vitis                                  0    0    0    --
Acanthaceae Avicennia Lamiales                  0    0    D    E
Acanthaceae Acanthus Lamiales                   0    1    D    F
Actinidiaceae Actinidia Ericales                D    1    0    2
Adoxaceae Viburnum Dipsacales                   0    0    D    F
Alseuosmiaceae Alseuosmia Asterales             D    1    0    0
Apiaceae Apium Apiales                          D    0    D    2
Apocynaceae Alstonia Gentianales                0    1    0    D
Aquifoliaceae Ilex Aquifoliales                 1    0    0    2
Araliaceae Aralia Apiales                       D    0    0    2
Aralidiaceae Aralidium Apiales                  1    ?    0    2
Argophyllaceae Argophyllum Asterales            0    ?    0    2
Asteraceae Helianthus Asterales                 D    D    D    F
Balsaminaceae Impatiens Ericales                0    0    1    0
Bignoniaceae Jacaranda Lamiales                 0    1    1    0
Boraginaceae Borago                             0    D    0    D
Boraginaceae Pholisma                           0    0    0    1
Bruniaceae Brunia                               0    0    0    2
Byblidaceae Byblis Lamiales                     0    ?    0    2
Calyceraceae Boopis Asterales                   0    ?    D    0
Campanulaceae Campanula Asterales               D    1    D    D
Caprifoliaceae Lonicera Dipsacales              0    ?    1    F
Cardiopteridaceae Cardiopteris Aquifoliales     D    0    0    0
Clethraceae Clethra Ericales                    0    0    0    2
Columelliaceae Columellia                       0    ?    1    2
Columelliaceae Desfontainia                     0    1    0    0
Convolvulaceae Ipomea Solanales                 D    1    0    1
Comaceae Cornus Comales                         D    0    0    F
Cyril laceae Cyril la Ericales                  0    ?    0    2
Diapensiaceae Diapensia Ericales                0    ?    0    2
Dipentodontaceae Dipentodon                     0    0    0    2
Dipsacaceae Dipsacus Dipsacales                 0    ?    D    D
Ebenaceae Diospyros Ericales                    0    1    0    2
Ebenaceae Lissocarpa Ericales                   0    ?    0    0
Eremosynaceae Eremosyne                         0    0    0    2
Ericaceae Erica Ericales                        0    0    0    1
Escalloniaceae Escallonia                       0    D    0    0
Eucommiaceae Eucommia Garryales                 1    0    0    --
Fouqueriaceae Fouqueria Ericales                0    1    0    0
Garryaceae Aucuba Garryales                     1    ?    0    2
Garryaceae Garry'a Garryales                    1    0    0    --
Gelsemiaceae Gelsemium Gentianales              0    ?    0    1
Gentianaceae Gentiana Gentianales               D    ?    0    ?
Gesneriaceae Streptocarpus Lamiales             0    1    1    0
Goodeniaceae Scaevola Asterales                 0    ?    1    0
Griseliniaceae Griselinia Apiales               1    0    0    2
Grubbiaceae Grubbia Comales                     0    0    0    --
Helwingiaceae Helwingia Aquifoliales            1    0    0    2
Hydrangeaceae Hydrangea Comales                 D    1    0    ?
Icacinaceae Icacina                             D    ?    0    2
Lamiaceae Lamium Lamiales                       D    ?    1    0
Lecythidaceae Barringtonia Ericales             0    1    1    2
Lentibulariaceae Pinguicula Lamiales            0    0    1    0
Loasaceae Loasa Comales                         0    1    0    2
Loganiaceae Logania Gentianales                 0    ?    0    1
Maesaceae Maesa Ericales                        0    0    0    0
Marcgraviaceae Marcgravia Ericales              0    ?    0    --
Martyniaceae Proboscidea Lamiales               0    1    1    1
Melanophyllaceae Melanophylla Apiales           0    ?    0    2
Menyanthaceae Menyanthes Asterales              0    1    0    2
Montianiaceae Kaliphora Solanales               1    0    0    2
Montianiaceae Montinia Solanales                1    0    0    2
Morinaceae Morina Dipsacales                    ?    ?    D    0
Myrsinaceae Myrsine Ericales                    0    0    0    F
Oleaceae Olea Lamiales                          0    0    0    2
Oncothecaceae Oncotheca                         0    0    0    1
Orobanchaceae Cyclocheilon Lamiales             0    ?    1    1
Orobanchaceae Lindenbergia Lamiales             0    1    1    0
Paracryphiaceae Paracryphia                     D    0    0    2
Pedal iaceae Sesamum Lamiales                   0    ?    1    0
Pentaphragmataceae Pentaphragma Asterales       D    0    0    2
Pentaphylacaceae Pentaphylax Ericales           0    ?    0    2
Phellinaceae Phelline Asterales                 1    ?    0    2
Phrymaceae Phryma Lamiales                      0    0    1    0
Phyllonomaceae Phyllonoma Aquifoliales          0    0    0    2
Pittosporaceae Pittosporum Apiales              D    0    0    2
Plantaginaceae Antirrhinum Lamiales             0    1    1    0
Plantaginaceae Globularia Lamiales              0    0    1    0
Plantaginaceae Plantago Lamiales                D    0    0    2
Plocospermataceae Plocosperma Lamiales          1    ?    0    1
Polemoniaceae Polemonium Ericales               0    1    0    0
Primulaceae Primula Ericales                    0    1    0    2
Roridulaceae Roridula Ericales                  0    ?    0    ?
Rousseaceae Carpodetus Asterales                0    0    0    2
Rousseaceae Roussea Asterales                   0    1    0    0
Rubiaceae Luculia Gentianales                   D    1    0    0
Sapotaceae Manilkara Ericales                   0    0    0    1
Sarraceniaceae Sarracenia Ericales              0    1    0    2
Scrophulariaceae Buddleja Lamiales              0    ?    0    0
Scrophulariaceae Myoporum Lamiales              0    ?    D    F
Scrophulariaceae Scrophularia Lamiales          0    1    1    F
Solanaceae Nicotiana Solanales                  0    1    0    K
Sphenocleaceae Sphenoclea Solanales             0    0    0    1
Stilbaceae Stilbe Lamiales                      0    ?    0    0
Stylidiaceae Donatia Asterales                  0    ?    0    2
Stylidiaceae Stylidium Asterales                0    ?    1    0
Styracaceae Styrax Ericales                     D    ?    0    E
Symplocaceae Symplocos Ericales                 D    1    0    E
Temstroemiaceae Temslroemia Ericales            0    1    0    2
Tetrachondraceae Tetrachondra Lamiales          0    ?    0    2
Tetrameristaceae Pelliciera Ericales            0    1    0    2
Tetrameristaceae Tetramerista Ericales          0    ?    0    2
Theaceae Schima Ericales                        0    1    0    2
Theophrastaceae Theophrasta Ericales            0    1    0    2
Torricelliaceae Torricellia Apiales             1    0    0    1
Tribelaceae Tribeles                            0    0    0    1
Vahliaceae Vahlia                               0    0    0    2
Valerianaceae Valeriana Dipsacales              D    ?    D    D
Verbenaceae Verbena Lamiales                    0    1    D    D
Solanites brongniartii                          0    0    0    2

Character                                       5

Paeoniaceae Paeonia                             --   A    0    0    ?
Vitaceae Vitis                                  0    2    0    1    1
Acanthaceae Avicennia Lamiales                  2    1    1    ?    ?
Acanthaceae Acanthus Lamiales                   E    2    2    ?    ?
Actinidiaceae Actinidia Ericales                2    2    D    0    ?
Adoxaceae Viburnum Dipsacales                   F    2    2    ?    ?
Alseuosmiaceae Alseuosmia Asterales             0    J    2    ?    ?
Apiaceae Apium Apiales                          0    2    0    0    ?
Apocynaceae Alstonia Gentianales                K    E    2    ?    ?
Aquifoliaceae Ilex Aquifoliales                 F    1    1    1    0
Araliaceae Aralia Apiales                       F    N    0    1    1
Aralidiaceae Aralidium Apiales                  2    2    0    ?    ?
Argophyllaceae Argophyllum Asterales            0    2    1    1    ?
Asteraceae Helianthus Asterales                 0    2    2    0    0
Balsaminaceae Impatiens Ericales                --   2    2    0    9
Bignoniaceae Jacaranda Lamiales                 F    2    2    ?    ?
Boraginaceae Borago                             K    J    2    ?    ?
Boraginaceae Pholisma                           ?    A    S    ?    ?
Bruniaceae Brunia                               2    E    1    ?    ?
Byblidaceae Byblis Lamiales                     1    2    1    0    ?
Calyceraceae Boopis Asterales                   0    J    2    ?    9
Campanulaceae Campanula Asterales               0    2    2    9    ?
Caprifoliaceae Lonicera Dipsacales              F    2    2    ?    ?
Cardiopteridaceae Cardiopteris Aquifoliales     2    2    1    ?    ?
Clethraceae Clethra Ericales                    2    S    D    0    ?
Columelliaceae Columellia                       2    I    1    ?    0
Columelliaceae Desfontainia                     E    2    2    1    ?
Convolvulaceae Ipomea Solanales                 D    E    3    0    ?
Comaceae Cornus Comales                         F    E    0    ?    ?
Cyril laceae Cyril la Ericales                  E    S    D    1    ?
Diapensiaceae Diapensia Ericales                E    2    2    ?    1
Dipentodontaceae Dipentodon                     0    S    0    ?    ?
Dipsacaceae Dipsacus Dipsacales                 2    2    2    ?    ?
Ebenaceae Diospyros Ericales                    K    V    2    1    0
Ebenaceae Lissocarpa Ericales                   1    1    2    1    ?
Eremosynaceae Eremosyne                         0    2    0    ?    ?
Ericaceae Erica Ericales                        E    E    K    0    ?
Escalloniaceae Escallonia                       F    E    0    0    0
Eucommiaceae Eucommia Garryales                 --   --   --   --   --
Fouqueriaceae Fouqueria Ericales                2    2    2    ?    ?
Garryaceae Aucuba Garryales                     0    1    0    1    ?
Garryaceae Garry'a Garryales                    --   1    --   --   --
Gelsemiaceae Gelsemium Gentianales              2    2    2    0    ?
Gentianaceae Gentiana Gentianales               1    E    2    0    ?
Gesneriaceae Streptocarpus Lamiales             2    2    2    ?    ?
Goodeniaceae Scaevola Asterales                 0    2    2    ?    ?
Griseliniaceae Griselinia Apiales               2    2    0    ?    ?
Grubbiaceae Grubbia Comales                     --   1    --   --   --
Helwingiaceae Helwingia Aquifoliales            0    K    0    ?    ?
Hydrangeaceae Hydrangea Comales                 K    E    0    ?    ?
Icacinaceae Icacina                             0    V    K    ?    ?
Lamiaceae Lamium Lamiales                       2    2    2    ?    ?
Lecythidaceae Barringtonia Ericales             2    J    0    1    1
Lentibulariaceae Pinguicula Lamiales            2    2    2    ?    ?
Loasaceae Loasa Comales                         0    2    K    ?    ?
Loganiaceae Logania Gentianales                 2    E    2    ?    ?
Maesaceae Maesa Ericales                        ?    2    2    1    0
Marcgraviaceae Marcgravia Ericales              --   E    O    1    1
Martyniaceae Proboscidea Lamiales               2    2    2    0    ?
Melanophyllaceae Melanophylla Apiales           2    2    0    1    ?
Menyanthaceae Menyanthes Asterales              F    2    2    ?    ?
Montianiaceae Kaliphora Solanales               0    1    0    1    ?
Montianiaceae Montinia Solanales                2    K    0    1    1
Morinaceae Morina Dipsacales                    2    2    2    ?    ?
Myrsinaceae Myrsine Ericales                    K    2    2    1    0
Oleaceae Olea Lamiales                          K    1    2    ?    ?
Oncothecaceae Oncotheca                         2    2    2    ?    ?
Orobanchaceae Cyclocheilon Lamiales             2    2    2    0    ?
Orobanchaceae Lindenbergia Lamiales             2    2    2    ?    ?
Paracryphiaceae Paracryphia                     2    2    0    ?    0
Pedal iaceae Sesamum Lamiales                   2    2    2    ?    ?
Pentaphragmataceae Pentaphragma Asterales       0    2    E    1    ?
Pentaphylacaceae Pentaphylax Ericales           2    2    0    ?    ?
Phellinaceae Phelline Asterales                 0    J    0    1    ?
Phrymaceae Phryma Lamiales                      ?    2    2    0    ?
Phyllonomaceae Phyllonoma Aquifoliales          0    E    0    0    ?
Pittosporaceae Pittosporum Apiales              2    2    D    ?    ?
Plantaginaceae Antirrhinum Lamiales             F    E    2    ?    ?
Plantaginaceae Globularia Lamiales              2    2    2    ?    ?
Plantaginaceae Plantago Lamiales                2    1    0    0    ?
Plocospermataceae Plocosperma Lamiales          2    2    2    ?    ?
Polemoniaceae Polemonium Ericales               1    J    2    ?    ?
Primulaceae Primula Ericales                    2    N    2    0    ?
Roridulaceae Roridula Ericales                  E    2    1    ?    ?
Rousseaceae Carpodetus Asterales                0    S    0    0    1
Rousseaceae Roussea Asterales                   0    E    2    1    0
Rubiaceae Luculia Gentianales                   K    E    2    ?    ?
Sapotaceae Manilkara Ericales                   2    U    2    ?    ?
Sarraceniaceae Sarracenia Ericales              2    2    0    ?    1
Scrophulariaceae Buddleja Lamiales              F    E    2    ?    ?
Scrophulariaceae Myoporum Lamiales              2    2    2    ?    ?
Scrophulariaceae Scrophularia Lamiales          F    2    2    ?    ?
Solanaceae Nicotiana Solanales                  K    E    J    ?    ?
Sphenocleaceae Sphenoclea Solanales             2    2    2    0    1
Stilbaceae Stilbe Lamiales                      1    2    2    ?    9
Stylidiaceae Donatia Asterales                  2    2    0    9    ?
Stylidiaceae Stylidium Asterales                2    2    2    ?    ?
Styracaceae Styrax Ericales                     F    2    K    0    ?
Symplocaceae Symplocos Ericales                 2    N    1    1    ?
Temstroemiaceae Temslroemia Ericales            2    2    0    0    ?
Tetrachondraceae Tetrachondra Lamiales          2    1    2    ?    ?
Tetrameristaceae Pelliciera Ericales            2    2    0    9    1
Tetrameristaceae Tetramerista Ericales          2    E    0    ?    ?
Theaceae Schima Ericales                        2    2    D    0    ?
Theophrastaceae Theophrasta Ericales            2    E    2    1    ?
Torricelliaceae Torricellia Apiales             0    2    0    ?    ?
Tribelaceae Tribeles                            1    2    0    1    ?
Vahliaceae Vahlia                               2    2    0    ?    ?
Valerianaceae Valeriana Dipsacales              2    K    2    ?    ?
Verbenaceae Verbena Lamiales                    2    I    2    ?    9
Solanites brongniartii                          0    2    E    0    1

Character                                       10

Paeoniaceae Paeonia                             0    0    3    2    2
Vitaceae Vitis                                  0    --   0    1    2
Acanthaceae Avicennia Lamiales                  0    0    0    0    1
Acanthaceae Acanthus Lamiales                   ?    0    2    0    1
Actinidiaceae Actinidia Ericales                0    0    3    E    1
Adoxaceae Viburnum Dipsacales                   0    3    0    0    1
Alseuosmiaceae Alseuosmia Asterales             0    0    0    0    E
Apiaceae Apium Apiales                          0    2    0    0    2
Apocynaceae Alstonia Gentianales                9    L    0    0    1
Aquifoliaceae Ilex Aquifoliales                 0    F    0    0    1
Araliaceae Aralia Apiales                       0    0    0    0    2
Aralidiaceae Aralidium Apiales                  0    0    0    0    2
Argophyllaceae Argophyllum Asterales            0    ?    0    0    2
Asteraceae Helianthus Asterales                 0    0    0    0    1
Balsaminaceae Impatiens Ericales                ?    0    0    --   2
Bignoniaceae Jacaranda Lamiales                 0    R    2    0    1
Boraginaceae Borago                             0    0    0    0    1
Boraginaceae Pholisma                           0    0    0    ?    1
Bruniaceae Brunia                               ?    0    0    0    E
Byblidaceae Byblis Lamiales                     9    0    0    0    E
Calyceraceae Boopis Asterales                   0    0    0    0    1
Campanulaceae Campanula Asterales               1    0    0    0    D
Caprifoliaceae Lonicera Dipsacales              1    0    0    0    1
Cardiopteridaceae Cardiopteris Aquifoliales     ?    0    0    0    1
Clethraceae Clethra Ericales                    0    2    1    D    2
Columelliaceae Columellia                       0    0    2    0    1
Columelliaceae Desfontainia                     0    0    0    0    1
Convolvulaceae Ipomea Solanales                 1    0    0    0    1
Comaceae Cornus Comales                         ?    0    0    0    2
Cyril laceae Cyril la Ericales                  0    0    D    0    2
Diapensiaceae Diapensia Ericales                1    0    1    0    1
Dipentodontaceae Dipentodon                     ?    0    0    0    2
Dipsacaceae Dipsacus Dipsacales                 1    0    2    0    1
Ebenaceae Diospyros Ericales                    0    3    M    0    1
Ebenaceae Lissocarpa Ericales                   ?    0    1    D    1
Eremosynaceae Eremosyne                         ?    0    0    0    2
Ericaceae Erica Ericales                        0    R    1    D    2
Escalloniaceae Escallonia                       1    R    0    0    2
Eucommiaceae Eucommia Garryales                 --   --   --   --   2
Fouqueriaceae Fouqueria Ericales                0    0    M    --   2
Garryaceae Aucuba Garryales                     0    0    0    0    2
Garryaceae Garry'a Garryales                    --   --   0    --   2
Gelsemiaceae Gelsemium Gentianales              0    0    0    0    1
Gentianaceae Gentiana Gentianales               1    0    0    0    1
Gesneriaceae Streptocarpus Lamiales             ?    0    2    0    1
Goodeniaceae Scaevola Asterales                 1    0    0    0    D
Griseliniaceae Griselinia Apiales               1    0    0    0    2
Grubbiaceae Grubbia Comales                     --   --   1    0    2
Helwingiaceae Helwingia Aquifoliales            1    0    0    0    2
Hydrangeaceae Hydrangea Comales                 1    0    L    K    2
Icacinaceae Icacina                             ?    0    0    0    E
Lamiaceae Lamium Lamiales                       0    0    2    0    1
Lecythidaceae Barringtonia Ericales             0    0    3    2    2
Lentibulariaceae Pinguicula Lamiales            0    0    2    ?    1
Loasaceae Loasa Comales                         0    2    M    E    E
Loganiaceae Logania Gentianales                 1    0    0    0    1
Maesaceae Maesa Ericales                        0    0    0    1    1
Marcgraviaceae Marcgravia Ericales              0    --   3    --   E
Martyniaceae Proboscidea Lamiales               0    0    2    0    1
Melanophyllaceae Melanophylla Apiales           0    2    0    0    2
Menyanthaceae Menyanthes Asterales              0    0    0    0    1
Montianiaceae Kaliphora Solanales               ?    3    0    0    2
Montianiaceae Montinia Solanales                ?    0    0    0    2
Morinaceae Morina Dipsacales                    1    0    2    ?    1
Myrsinaceae Myrsine Ericales                    0    R    0    1    1
Oleaceae Olea Lamiales                          ?    0    F    0    1
Oncothecaceae Oncotheca                         ?    ?    0    0    1
Orobanchaceae Cyclocheilon Lamiales             0    ?    2    0    1
Orobanchaceae Lindenbergia Lamiales             1    0    2    0    1
Paracryphiaceae Paracryphia                     ?    ?    D    D    2
Pedal iaceae Sesamum Lamiales                   ?    0    F    0    1
Pentaphragmataceae Pentaphragma Asterales       ?    0    0    0    1
Pentaphylacaceae Pentaphylax Ericales           0    0    0    0    2
Phellinaceae Phelline Asterales                 0    2    0    0    2
Phrymaceae Phryma Lamiales                      ?    0    2    0    1
Phyllonomaceae Phyllonoma Aquifoliales          ?    0    0    0    2
Pittosporaceae Pittosporum Apiales              1    0    0    0    2
Plantaginaceae Antirrhinum Lamiales             1    0    F    0    1
Plantaginaceae Globularia Lamiales              ?    0    F    0    1
Plantaginaceae Plantago Lamiales                1    0    0    ?    2
Plocospermataceae Plocosperma Lamiales          ?    0    0    0    1
Polemoniaceae Polemonium Ericales               ?    0    0    0    1
Primulaceae Primula Ericales                    0    0    0    1    1
Roridulaceae Roridula Ericales                  ?    0    0    0    1
Rousseaceae Carpodetus Asterales                ?    0    0    0    2
Rousseaceae Roussea Asterales                   0    3    0    0    2
Rubiaceae Luculia Gentianales                   ?    0    0    0    1
Sapotaceae Manilkara Ericales                   0    F    M    1    1
Sarraceniaceae Sarracenia Ericales              0    0    M    2    2
Scrophulariaceae Buddleja Lamiales              0    0    0    0    1
Scrophulariaceae Myoporum Lamiales              ?    0    F    0    1
Scrophulariaceae Scrophularia Lamiales          1    0    F    0    1
Solanaceae Nicotiana Solanales                  ?    R    F    0    1
Sphenocleaceae Sphenoclea Solanales             ?    0    0    0    1
Stilbaceae Stilbe Lamiales                      0    0    0    0    1
Stylidiaceae Donatia Asterales                  0    0    2    ?    0
Stylidiaceae Stylidium Asterales                0    0    2    ?    0
Styracaceae Styrax Ericales                     1    0    M    0    E
Symplocaceae Symplocos Ericales                 0    2    M    0    1
Temstroemiaceae Temslroemia Ericales            0    F    M    --   2
Tetrachondraceae Tetrachondra Lamiales          ?    0    0    0    1
Tetrameristaceae Pelliciera Ericales            0    0    0    0    2
Tetrameristaceae Tetramerista Ericales          0    ?    0    0    2
Theaceae Schima Ericales                        0    0    M    1    E
Theophrastaceae Theophrasta Ericales            0    0    0    1    1
Torricelliaceae Torricellia Apiales             ?    ?    0    0    2
Tribelaceae Tribeles                            ?    ?    0    0    2
Vahliaceae Vahlia                               ?    0    0    0    2
Valerianaceae Valeriana Dipsacales              1    0    2    0    1
Verbenaceae Verbena Lamiales                    0    0    2    0    1
Solanites brongniartii                          1    1    0    0    1

Character                                       15

Paeoniaceae Paeonia                             1    ?    0    0    --
Vitaceae Vitis                                  1    ?    0    0    --
Acanthaceae Avicennia Lamiales                  1    ?    0    0    0
Acanthaceae Acanthus Lamiales                   1    9    9    0    1
Actinidiaceae Actinidia Ericales                1    9    0    D    --
Adoxaceae Viburnum Dipsacales                   1    ?    0    0    0
Alseuosmiaceae Alseuosmia Asterales             1    ?    0    0    0
Apiaceae Apium Apiales                          1    ?    0    0    --
Apocynaceae Alstonia Gentianales                0    ?    ?    0    1
Aquifoliaceae Ilex Aquifoliales                 1    ?    ?    0    --
Araliaceae Aralia Apiales                       1    9    0    0    --
Aralidiaceae Aralidium Apiales                  1    9    0    0    --
Argophyllaceae Argophyllum Asterales            1    ?    0    0    --
Asteraceae Helianthus Asterales                 0    ?    1    0    1
Balsaminaceae Impatiens Ericales                0    ?    0    ?    1
Bignoniaceae Jacaranda Lamiales                 1    ?    ?    0    1
Boraginaceae Borago                             0    9    9    0    1
Boraginaceae Pholisma                           0    9    0    0    1
Bruniaceae Brunia                               1    9    D    0    --
Byblidaceae Byblis Lamiales                     0    ?    0    1    --
Calyceraceae Boopis Asterales                   1    ?    0    0    ?
Campanulaceae Campanula Asterales               1    ?    0    0    0
Caprifoliaceae Lonicera Dipsacales              1    ?    0    0    0
Cardiopteridaceae Cardiopteris Aquifoliales     0    ?    0    0    ?
Clethraceae Clethra Ericales                    1    ?    0    1    --
Columelliaceae Columellia                       0    ?    0    0    --
Columelliaceae Desfontainia                     0    ?    0    0    1
Convolvulaceae Ipomea Solanales                 1    ?    ?    0    1
Comaceae Cornus Comales                         1    ?    ?    0    0
Cyril laceae Cyril la Ericales                  1    ?    0    0    --
Diapensiaceae Diapensia Ericales                1    ?    0    0    --
Dipentodontaceae Dipentodon                     1    ?    0    0    --
Dipsacaceae Dipsacus Dipsacales                 1    ?    0    0    0
Ebenaceae Diospyros Ericales                    D    ?    0    D    --
Ebenaceae Lissocarpa Ericales                   0    ?    1    0    1
Eremosynaceae Eremosyne                         1    ?    0    0    --
Ericaceae Erica Ericales                        1    ?    0    1    1
Escalloniaceae Escallonia                       1    ?    0    0    1
Eucommiaceae Eucommia Garryales                 0    ?    1    0    --
Fouqueriaceae Fouqueria Ericales                1    ?    0    0    0
Garryaceae Aucuba Garryales                     1    ?    0    0    --
Garryaceae Garry'a Garryales                    1    ?    0    0    0
Gelsemiaceae Gelsemium Gentianales              1    ?    0    0    1
Gentianaceae Gentiana Gentianales               1    ?    0    D    ?
Gesneriaceae Streptocarpus Lamiales             1    ?    0    0    0
Goodeniaceae Scaevola Asterales                 1    ?    ?    0    1
Griseliniaceae Griselinia Apiales               1    ?    0    0    --
Grubbiaceae Grubbia Comales                     1    ?    1    0    --
Helwingiaceae Helwingia Aquifoliales            1    ?    0    0    --
Hydrangeaceae Hydrangea Comales                 1    ?    0    ?    0
Icacinaceae Icacina                             1    ?    ?    0    --
Lamiaceae Lamium Lamiales                       1    ?    ?    0    1
Lecythidaceae Barringtonia Ericales             1    ?    0    D    --
Lentibulariaceae Pinguicula Lamiales            1    ?    0    0    1
Loasaceae Loasa Comales                         1    ?    0    0    --
Loganiaceae Logania Gentianales                 1    ?    0    0    1
Maesaceae Maesa Ericales                        0    ?    0    0    0
Marcgraviaceae Marcgravia Ericales              1    ?    0    0    --
Martyniaceae Proboscidea Lamiales               1    ?    0    0    1
Melanophyllaceae Melanophylla Apiales           0    ?    0    0    --
Menyanthaceae Menyanthes Asterales              1    ?    ?    0    --
Montianiaceae Kaliphora Solanales               0    ?    1    0    --
Montianiaceae Montinia Solanales                1    ?    0    0    --
Morinaceae Morina Dipsacales                    1    ?    0    0    0
Myrsinaceae Myrsine Ericales                    0    ?    0    D    0
Oleaceae Olea Lamiales                          0    ?    1    0    --
Oncothecaceae Oncotheca                         0    ?    1    0    ?
Orobanchaceae Cyclocheilon Lamiales             1    ?    0    0    1
Orobanchaceae Lindenbergia Lamiales             1    ?    ?    0    1
Paracryphiaceae Paracryphia                     1    ?    0    0    --
Pedal iaceae Sesamum Lamiales                   1    ?    ?    0    1
Pentaphragmataceae Pentaphragma Asterales       0    ?    0    0    --
Pentaphylacaceae Pentaphylax Ericales           1    ?    0    1    --
Phellinaceae Phelline Asterales                 0    ?    0    0    --
Phrymaceae Phryma Lamiales                      1    ?    0    0    1
Phyllonomaceae Phyllonoma Aquifoliales          1    ?    0    0    --
Pittosporaceae Pittosporum Apiales              0    ?    0    D    --
Plantaginaceae Antirrhinum Lamiales             1    ?    0    0    1
Plantaginaceae Globularia Lamiales              1    ?    ?    0    0
Plantaginaceae Plantago Lamiales                1    ?    0    0    --
Plocospermataceae Plocosperma Lamiales          1    ?    0    0    1
Polemoniaceae Polemonium Ericales               1    ?    ?    0    1
Primulaceae Primula Ericales                    0    ?    0    D    --
Roridulaceae Roridula Ericales                  1    ?    0    1    ?
Rousseaceae Carpodetus Asterales                1    ?    0    0    --
Rousseaceae Roussea Asterales                   1    ?    0    0    0
Rubiaceae Luculia Gentianales                   1    ?    ?    0    1
Sapotaceae Manilkara Ericales                   1    ?    0    0    0
Sarraceniaceae Sarracenia Ericales              1    ?    0    0    --
Scrophulariaceae Buddleja Lamiales              1    ?    0    0    1
Scrophulariaceae Myoporum Lamiales              1    ?    0    0    0
Scrophulariaceae Scrophularia Lamiales          1    ?    0    0    0
Solanaceae Nicotiana Solanales                  D    ?    ?    D    D
Sphenocleaceae Sphenoclea Solanales             0    ?    ?    0    1
Stilbaceae Stilbe Lamiales                      0    ?    0    0    1
Stylidiaceae Donatia Asterales                  1    ?    0    0    ?
Stylidiaceae Stylidium Asterales                1    ?    0    0    0
Styracaceae Styrax Ericales                     1    9    1    0    D
Symplocaceae Symplocos Ericales                 1    ?    0    0    0
Temstroemiaceae Temslroemia Ericales            1    ?    0    0    --
Tetrachondraceae Tetrachondra Lamiales          1    ?    ?    0    --
Tetrameristaceae Pelliciera Ericales            1    ?    1    0    --
Tetrameristaceae Tetramerista Ericales          1    ?    0    0    --
Theaceae Schima Ericales                        1    ?    D    0    --
Theophrastaceae Theophrasta Ericales            1    ?    1    0    --
Torricelliaceae Torricellia Apiales             ?    ?    0    0    1
Tribelaceae Tribeles                            1    ?    0    0    ?
Vahliaceae Vahlia                               1    ?    0    0    --
Valerianaceae Valeriana Dipsacales              1    ?    0    0    0
Verbenaceae Verbena Lamiales                    1    ?    ?    0    1
Solanites brongniartii                          0    0    1    ?    --

Character                                       20

Paeoniaceae Paeonia                             B    0    C    1
Vitaceae Vitis                                  1    1    1    1
Acanthaceae Avicennia Lamiales                  1    1    1    3
Acanthaceae Acanthus Lamiales                   1    1    1    3
Actinidiaceae Actinidia Ericales                G    1    H    0
Adoxaceae Viburnum Dipsacales                   I    1    1    1
Alseuosmiaceae Alseuosmia Asterales             1    1    1    3
Apiaceae Apium Apiales                          1    1    2    0
Apocynaceae Alstonia Gentianales                1    1    E    1
Aquifoliaceae Ilex Aquifoliales                 H    1    D    M
Araliaceae Aralia Apiales                       B    1    P    0
Aralidiaceae Aralidium Apiales                  2    1    3    0
Argophyllaceae Argophyllum Asterales            9    1    1    M
Asteraceae Helianthus Asterales                 1    1    1    3
Balsaminaceae Impatiens Ericales                Q    1    1    0
Bignoniaceae Jacaranda Lamiales                 1    1    1    3
Boraginaceae Borago                             1    1    1    3
Boraginaceae Pholisma                           T    1    1    M
Bruniaceae Brunia                               E    1    J    0
Byblidaceae Byblis Lamiales                     1    1    1    1
Calyceraceae Boopis Asterales                   1    1    1    1
Campanulaceae Campanula Asterales               I    1    1    3
Caprifoliaceae Lonicera Dipsacales              E    1    1    1
Cardiopteridaceae Cardiopteris Aquifoliales     1    1    2    D
Clethraceae Clethra Ericales                    2    1    1    3
Columelliaceae Columellia                       1    1    1    3
Columelliaceae Desfontainia                     4    1    1    1
Convolvulaceae Ipomea Solanales                 1    1    E    0
Comaceae Cornus Comales                         U    1    1    M
Cyril laceae Cyril la Ericales                  U    1    D    R
Diapensiaceae Diapensia Ericales                2    1    1    3
Dipentodontaceae Dipentodon                     2    1    1    0
Dipsacaceae Dipsacus Dipsacales                 1    1    1    R
Ebenaceae Diospyros Ericales                    M    1    W    0
Ebenaceae Lissocarpa Ericales                   3    1    1    2
Eremosynaceae Eremosyne                         1    1    2    1
Ericaceae Erica Ericales                        Q    1    1    M
Escalloniaceae Escallonia                       U    1    1    M
Eucommiaceae Eucommia Garryales                 l    1    1    0
Fouqueriaceae Fouqueria Ericales                2    1    M    0
Garryaceae Aucuba Garryales                     ?    ?    1    1
Garryaceae Garry'a Garryales                    E    1    S    0
Gelsemiaceae Gelsemium Gentianales              1    1    1    0
Gentianaceae Gentiana Gentianales               1    1    1    3
Gesneriaceae Streptocarpus Lamiales             1    1    1    3
Goodeniaceae Scaevola Asterales                 1    1    1    0
Griseliniaceae Griselinia Apiales               2    1    3    0
Grubbiaceae Grubbia Comales                     1    1    1    3
Helwingiaceae Helwingia Aquifoliales            S    1    Q    0
Hydrangeaceae Hydrangea Comales                 B    1    C    D
Icacinaceae Icacina                             U    1    1    3
Lamiaceae Lamium Lamiales                       1    1    1    3
Lecythidaceae Barringtonia Ericales             B    1    1    M
Lentibulariaceae Pinguicula Lamiales            1    1    D    3
Loasaceae Loasa Comales                         G    1    1    0
Loganiaceae Logania Gentianales                 E    1    1    1
Maesaceae Maesa Ericales                        1    1    1    3
Marcgraviaceae Marcgravia Ericales              B    1    0    R
Martyniaceae Proboscidea Lamiales               1    1    1    3
Melanophyllaceae Melanophylla Apiales           E    1    S    0
Menyanthaceae Menyanthes Asterales              1    1    1    3
Montianiaceae Kaliphora Solanales               1    1    ?    0
Montianiaceae Montinia Solanales                1    1    1    M
Morinaceae Morina Dipsacales                    1    1    1    R
Myrsinaceae Myrsine Ericales                    A    1    1    L
Oleaceae Olea Lamiales                          1    l    1    3
Oncothecaceae Oncotheca                         4    1    0    --
Orobanchaceae Cyclocheilon Lamiales             1    1    1    4
Orobanchaceae Lindenbergia Lamiales             1    1    1    M
Paracryphiaceae Paracryphia                     U    1    G    1
Pedal iaceae Sesamum Lamiales                   1    1    1    3
Pentaphragmataceae Pentaphragma Asterales       E    1    1    1
Pentaphylacaceae Pentaphylax Ericales           4    1    1    3
Phellinaceae Phelline Asterales                 u    1    0    3
Phrymaceae Phryma Lamiales                      1    1    1    3
Phyllonomaceae Phyllonoma Aquifoliales          1    1    2    0
Pittosporaceae Pittosporum Apiales              1    1    1    M
Plantaginaceae Antirrhinum Lamiales             1    1    1    R
Plantaginaceae Globularia Lamiales              1    1    1    M
Plantaginaceae Plantago Lamiales                1    1    1    3
Plocospermataceae Plocosperma Lamiales          1    1    1    0
Polemoniaceae Polemonium Ericales               2    1    1    3
Primulaceae Primula Ericales                    4    1    1    1
Roridulaceae Roridula Ericales                  2    1    1    M
Rousseaceae Carpodetus Asterales                A    1    1    1
Rousseaceae Roussea Asterales                   T    1    1    1
Rubiaceae Luculia Gentianales                   I    1    1    M
Sapotaceae Manilkara Ericales                   B    1    1    M
Sarraceniaceae Sarracenia Ericales              4    1    1    3
Scrophulariaceae Buddleja Lamiales              1    1    1    M
Scrophulariaceae Myoporum Lamiales              1    1    1    0
Scrophulariaceae Scrophularia Lamiales          1    1    1    3
Solanaceae Nicotiana Solanales                  1    1    1    3
Sphenocleaceae Sphenoclea Solanales             1    1    0    1
Stilbaceae Stilbe Lamiales                      1    1    1    3
Stylidiaceae Donatia Asterales                  E    1    S    1
Stylidiaceae Stylidium Asterales                1    1    1    3
Styracaceae Styrax Ericales                     U    1    1    M
Symplocaceae Symplocos Ericales                 U    1    1    M
Temstroemiaceae Temslroemia Ericales            X    1    M    3
Tetrachondraceae Tetrachondra Lamiales          1    1    1    1
Tetrameristaceae Pelliciera Ericales            1    1    1    0
Tetrameristaceae Tetramerista Ericales          Q    1    1    R
Theaceae Schima Ericales                        U    1    Y    3
Theophrastaceae Theophrasta Ericales            4    1    1    0
Torricelliaceae Torricellia Apiales             S    1    3    R
Tribelaceae Tribeles                            2    1    1    3
Vahliaceae Vahlia                               E    1    S    1
Valerianaceae Valeriana Dipsacales              2    1    1    R
Verbenaceae Verbena Lamiales                    1    1    1    3
Solanites brongniartii                          1    ?    2    1

Table 7 Composition of the data
matrix for the phylogenetic analysis
of Solanites brongniartii.
char-characters, inf-informative
characters

Partition        # char    # inf     # taxa

Morphology       23        21        111
matK             2194      1334      131 (a)
ndhF             2403      1248      127 (b)
rbcL             1432      493       131
rps16            1453      755       125
trnT-F           2299      1103      129
trnV             2234      935       125
matK indels      471       141       131
ndhF indels      101       26        127
rbcL indels      7         1         131
rps16 indels     766       308       125
trnT-F indels    1054      297       129
trn V indels     733       261       125
Total            15170     6920      133

Table 8 Matrix of morphological characters including the fossil taxa,
Nahinda axamilpensis, Coahuilanthus belindae, and Solanites pusillus.
A-[01], K-Cretaceous, Olig-Oligocene, Eoc-Eocene

Character               1                   5

Vitis                   1    1    1    1    0    2    ?    2    0
Mortonia                2    2    2    0    0    2    ?    2    0
Adolphia                1    2    2    1    1    1    1    2    2
Berchemia               1    2    2    2    0    1    0    0    1
Ceanothus               2    2    2    1    0    2    0    1    2
Colletia                --   --   --   --   --   --   --   --   0
Colubrina               1    2    2    2    2    2    1    2    1
Condalia                --   --   2    1    2    0    0    0    0
Coahuilanthus (K)       1    1    1    0    0    0    0    0    1
Nahinda (Olig)          1    2    2    0    2    2    1    2    1
Gouania                 1    1    1    0    1    1    l    1    1
Karwinskia              1    2    2    1    1    0    0    1    1
Krugiodendron           --   --   2    1    2    1    1    1    --
Paliurus                0    1    1    0    1    0    0    0    1
Reynosia                --   --   2    1    0    1    0    1    --
Rhamnus                 1    1    2    2    0    2    1    2    1
Sageretia               0    2    1    0    1    0    0    0    1
Ziziphus                1    2    2    1    2    1    1    1    1
Solanites pusillus      2    3    3    3    3    ?    ?    --   0
  (Eoc)

Character               10                       15

Vitis                   5    0    6    0    0    4    2    0    2    1
Mortonia                6    0    4    0    0    ?    ?    0    2    2
Adolphia                4    0    4    2    0    0    0    0    3    1
Berchemia               1    0    4    2    0    0    2    0    2    1
Ceanothus               0    0    5    1    0    1    0    3    3    1
Colletia                3    0    2    1    0    2    0    0    3    1
Colubrina               0    0    1    1    0    A    1    0    3    1
Condalia                --   0    1    1    0    0    4    0    2    3
Coahuilanthus (K)       --   0    4    1    0    --   --   --   --   3
Nahinda (Olig)          0    0    1    2    1    --   --   --   --   1
Gouania                 0    0    3    1    0    3    --   0    3    1
Karwinskia              4    1    4    2    0    0    2    0    2    1
Krugiodendron           --   0    4    2    1    0    4    0    2    2
Paliurus                0    1    0    2    0    5    --   --   A    1
Reynosia                --   0    3    1    0    0    4    0    2    2
Rhamnus                 2    1    4    1    0    0    2    3    2    2
Sageretia               0    0    4    2    0    0    3    2    3    1
Ziziphus                0    0    0    2    1    0    0    0    3    2
Solanites pusillus      1    0    0    1    1    ?    ?    ?    ?    1
  (Eoc)

Character               20                       25

Vitis                   1    4    0    0    2    0    0
Mortonia                1    1    0    1    3    0    0
Adolphia                1    1    0    1    1    1    0
Berchemia               1    1    2    0    1    0    0
Ceanothus               2    2    0    0    1    1    0
Colletia                2    1    0    1    0    2    0
Colubrina               2    0    0    1    1    2    0
Condalia                1    0    0    2    1    --   0
Coahuilanthus (K)       3    1    1    --   1    2    0
Nahinda (Olig)          2    0    0    0    1    2    0
Gouania                 0    2    2    1    1    2    1
Karwinskia              1    2    0    0    1    2    0
Krugiodendron           1    2    0    2    1    --   0
Paliurus                1    3    0    0    1    0    1
Reynosia                1    0    0    2    1    --   0
Rhamnus                 1    1    2    1    2    1    0
Sageretia               1    3    0    0    1    2    0
Ziziphus                1    0    0    0    1    2    0
Solanites pusillus      ?    0    0    --   1    ?    ?
  (Eoc)

Table 9 Composition of the
data matrix for the phylogenetic
analysis of Solanites pusillus.
char-characters, inf-informative
characters

Partition      # characters    # informative    # taxa

Morphology     26              25               19
rbcL           1428            107              16
trnL           1195            83               16
trnL indels    102             34               16

Table 10 Number of MPTs and position of the fossil taxon, Solanites
brongniartii, when each morphological character was excluded from the
analysis, *-indicates analyses that produced different topologies from
the 83 obtained when all characters were included.--indicates analysis
not performed (uninformative character)

                                          Position(s) of Solanites
      Character excluded          MPT     brongniartii

1     Flower sexuality            83      same
2     Flower size                 83      same
3     Corolla symmetry            83      same
4     Corolla shape               8       sister to Oncotheca
5     Corolla aestivation         12      sister to Olea or to
                                            Montiniaceae (Solanales)
6     Corolla merosity            83      same
7     Corolla fusion              14 *    various positions within
                                            Solanales, sister to
                                            Olea, Ilex, Vahlia, or a
                                            feMa-Solanales-
                                            Gentianales clade
8     Corolla texture             5       in a polytomy with
                                            Apodytes and Oncotheca
9     Corolla persistence         19      sister to Helwingia-
                                            Phyllonoma (Aquifoliales)
10    Petal midvein               5       in a polytomy with
                                            Apodytes and Oncotheca
11    Apices of petals            83      same
12    Stamen number relative      83      same
        to petals
13    Stamen cycle orientation    83      same
14    Stamen attachment           97 *    same 6 positions plus as
                                            sister to a fhMa-
                                            Solanales-Gentianales
                                            clade, in a polytomy with
                                            Nicotiana and other
                                            Solanales, or in a
                                            polytomy with Hydrolea
                                            and Sphenoclea
15    Filament vs. anther         32      sister to Ilex or to Vahlia
16    Anther surface              --      --
17    Connective projection       51      sister to Ilex, Vahlia or
                                            Montiniaceae (Solan.)
18    Anther dehiscence           83      same
19    Stamen vs. corolla lobes    83      same
20    Carpel number               83      same
21    Ovary                       --      --
22    Number of styles            51      sister to Oncotheca,
                                            Olea, or Montiniaceae
23    Stigma shape                6       sister to Olea
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Author:Millan, Marcela; Crepet, William
Publication:The Botanical Review
Article Type:Report
Geographic Code:4EUFR
Date:Jun 1, 2014
Words:14574
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