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Evidence for Holobaraminic Status of the Verbenaceae (Verbena Family)

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Evidence for Holobaraminic Status of the Verbenaceae (Verbena Family) OPEN ACCESS JCTS Report SERIES B Evidence for Holobaraminic Status of the Verbenaceae (Verbena Family) R.W. Sanders Core Academy of Science, Dayton, TN 37321 Abstract The baraminic status of the Verbenaceae s. str. was investigated using baraminic distance correlations, bootstrap analysis, and multidimensional scaling coordinates. Data were obtained from personal observations and the literature for 80 characters in 25 genera of the Verbenaceae and 19 genera representing 8 near and far outgroup families. Results showed continuity among members of the Verbenaceae and discontinuity of the Verbenaceae from all remaining families, supporting the holobaraminic status of the family. The hypothesis that baramins approximate taxonomic families is corroborated, as is the distinction of the Verbenaceae s. str. from the Lamiaceae, including the subfamilies that have been transferred from the Verbenaceae s. l. to the Lamiaceae by recent cladistic and molecular analyses. Editor: J.W. Francis Received January 26, 2012; Accepted September 13, 2016; Published September 26, 2016 Introduction although a molecular classification at the tribal level has been published recently (Marx et al. 2010; Yuan et al. 2010; O’Leary To date, statistical baraminology has been applied to the study et al. 2012; Lu-Irving & Olmstead 2013). Generally, the family of only 28 families of plants (Wood 2008b, pp. 149-222, 237-239). is characterized by opposite, simple leaves; flowers in elongate or In all cases, these studies were based on publicly available data condensed (head-like) spikes; a weak development of bilateral sets in the conventional literature or online databases. However, floral symmetry; arching corolla tubes; an abruptly flattened and this work needs to be expanded to include plant families for which flaring “trumpet bell” of the corolla; and a fruit with two or four there are no published morphological data sets that are adequate “stones” or nutlets that are sometimes fused (Figure 1). for the method. Because I have been studying the Verbenaceae s. It is of interest that the Verbenaceae are not divided at the str. (Lamiales) for many years and have considerable experience subfamily rank. This is because, as currently circumscribed, the with the group (e.g., Sanders 2001), I provide here a data set newly family contains only the name-bearing subfamily that traditionally compiled from my own observations and numerous published was associated with several other subfamilies as the Verbenaceae descriptions and morphological studies to use in baraminology. s. l. These other subfamilies were transferred to the Lamiaceae The Verbenaceae as currently accepted in the conventional (mint family) in 1992 (Cantino 1992a, 1992b) on the basis of literature comprise perhaps 500 to 1,000 species in about 35 genera several anatomically cryptic, but profound, differences which, and are typical of Lamiales (= Scrophulariales = Bignoniales), the though known since the 1930s, were not widely recognized until order that includes mints, snapdragons, foxgloves, speedwells, cladistic studies brought them to light. Details of this taxonomic plantains, gesneriads, shrimp-plants, and trumpet vines among history and the relevant traits are elaborated by Sanders (2001). others (viz., the “advanced” angiosperms with tubular, bilaterally The hope is that determination of the boundary around the symmetrical flowers and superior, 2-chambered ovaries). The baramin(s) of the plants included in the Verbenaceae will allow genera and species of Verbenaceae share a number of similarities further investigation into a number of vexing issues that emerge that suggest them to be closely related, but are not clearly when young-age assumptions are applied to these plants:the differentiated into infrafamilial groups. As a result, there has origin of chemical defenses and prickles (i.e., natural evil), pre- been no consensus for a tribal classification based on morphology, Flood biogeography and diversity, post-Flood dispersal and ©2016 The author. This article is open access and distributed under a Creative Commons Attribution License, which allows unrestricted use, distribution, and repro- duction in any medium as long as the original author and medium are credited. Citation: Sanders. 2016. Evidence for holobaraminic status of the Verbenaceae (Verbena family). Journal of Creation Theology and Science Series B: Life Sciences 6:81-90. Table 1. Genera and higher taxa sampled for the baraminological analysis of the Verbenaceae in the context of the classification used in this paper. In the representation numbers, the numerator is the number of genera sampled and the denominator is the number of genera in the higher taxon. In the Lamiaceae, the genera in the category “ex Verbenaceae s. l.” represent three subfamilies that were placed in the Verbenaceae before 1992 but have been transferred to the Lamiaceae. Higher Taxa Genera sampled Representation Outgroup—Boraginales: Boranginaceae 2/150 Cordioideae Cordia Boraginoideae Anchusa Lamiales Martyniaceae Proboscidea 1/5 Bignoniaceae Jacaranda 1/110 Phyrmaceae Phryma 1/1 Scrophulariaceae* Antirrhinum 4/220 Mimulus Penstemon Scrophularia Acanthaceae Elytraria 3/350 Justicia Thunbergia Avicenniaceae Avicennia 1/1 Lamiaceae s. l. 6/230 Figure 1. Portion of inflorescence of Stachytarpheta Lamiaceae s. str. Lamium 3/170 Mentha jamaicensis, illustrating family characteristics Salvia mentioned in text. Photo by J.M. Garg, Wikimedia ex Verbenaceae s. l. Clerodendrum 3/60 Holmskioldia Commons. Petitia Verbenaceae s. str. 23/35 Petreeae Petrea 2/2 Xolocotzia diversification, extent of design in homoplasy and its role in post- Citharexyleae Citharexylum 4/8 Flood adaptation. Coelocarpum Duranta Raphithamnus Methods Verbeneae Bouchea 13/16 Taxa. A total of 44 genera in nine families are sampled in Casselia Diostea the analysis. The ingroup includes 25 of the 35 genera and all Dipyrena tribes of the Verbenaceae. Seven other families of Lamiales, Glandularia represented by 17 genera, were selected as near outgroups (or Junellia extended ingroup). Based on cladistic morphological studies Lampaya Neosparton (Cantino 1982) and molecular systematics (Soltis et al. 2005), Pitraea the Boraginaceae (Boraginales or Solanales) were chosen as the Priva principal outgroup, being represented by two genera in different Stachytarpheta subfamilies (Table 1). Tamonia Verbena Characters. An holistic data set of 80 characters representing Lantaneae Aloysia 6/9 secondary chemistry (3 characters), general growth form (1 Goniostachyum character), stem morphology (1), epidermal anatomy (5), leaf Lantana morphology (2), inflorescence morphology (6), flower morphology Lippia (26), floral anatomy (8), pollen anatomy (4), embryology (16), Nashia Riedelia fruit morphology (7), and chromosomes (1) is detailed in Table 2. Data were compiled primarily from 30 years of personal *Note that the Scrophulariaceae are treated traditionally here, but recent observations and general references on Verbenaceae (Troncoso classifications based on molecular data have reapportioned most of its genera 1974; Cronquist 1981; Sanders 2001; Atkins 2004). Numerous with segregate or adjacent families. However, those realignments are not relevant to the results presented here. other references provided additional data, especially for either JCTS B: Life Sciences www.coresci.org/jcts Volume 6:82 Table 2. Characters and character states used to generate the data matrix. Data sources cited below. 1. Habit: 0=tree; 1=shrub; 2=herb; 3=liana; 4=vine; 30. Petal connation, pattern: 0=equally 5-lobed; 1=weak- 5=subshrubs. ly 2 upper + 3 lower (weakly bilabiate); 2=strongly 2 + 2. Stem: 0=round; 1=square. 3 (strongly bilabiate); 3=± 4 equal lobes. 3. Hairs, cell structure: 0=unicellular only; 1=uniseriate 31. Stamen number, fertile: 0=5; 1=4; 2=2. common; 2=biseriate; 3=multiseriate. 32. Stamen number, sterile: 0=0; 1=1; 2=2. 4. Hairs, posture: 0= straight; 1=proximally geniculate; 33. Stamen lengths: 0=equal lengths; 1=abaxial pair lon- 2=distally hooked. ger; 2=adaxial pair longer. 5. Glands: 0=”sessile”; 1=stalked. 34. Anther, shape: 0=truncate; 1=sagittate; 2=divaricate. 6. Gland head, no. of cells: 0=1-4; 1=5-20; 2=complex; 35. Anther, connective tissue: 0=placentoid tissue intrud- 3=Not applicable. ing into thecae; 1=no placentoid tissue. 7. Stomata: 0=only anomocytic; 1=diacytic & diallelocyt- 36. Anther, connective length: 0=not exceeding thecae; ic common. 1=longer than thecae. 8. Flavones present: 0=6-oxy; 1=6-oxy absent. 37. Anther, connective size: 0=not dilated; 1=dilated. 9. Iridoids, non-seco-iridoid types: 0=present; 1=absent. 38. Anther, connective appendage: 0=glandular append- 10. Oligosaccharides: 0=present; 1=absent. age lacking; 1=glandular appendage present on abaxial 11. Leaf arrangement: 0=decussate; 1=alternate; 2=ro- anther pair. sette; 3=decussate below, alternate into inflorescence. 39. Pollen, aperture no.: 0=3; 1=6; 2=2; 3=4; 4=fused. 12. Leaf blade: 0=unlobed; 1=lobed or dissected; 2=re- 40. Pollen, aperture structure: 0=colpate; 1=colporate; duced, scale=like. 2=not applicable. 13. Inflorescence type: 0= spiciform thyrse; 1= axillary/ 41. Pollen, colpus ornamentation: 0=exine thickened ad- terminal cyme; 2=raceme/spike; 3=capitulum; 4=heli- jacent to colpi; 1=exine not thickened adjacent to colpi; coid cyme; 5=terminal/axillary thyrse. 2=not applicable. 14. Inflorescence axis structure: 0=twiggy w/o depres- 42. Pollen type: 0=general for Lamiales; 1=Verbena type; sions for flowers; 1=thickened
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