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OPEN ACCESS JCTS Report SERIES B

Evidence for Holobaraminic Status of the ( )

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 , including the 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 (Wood 2008b, pp. 149-222, 237-239). is characterized by opposite, simple ; 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 families for which flaring “trumpet bell” of the corolla; and a 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. () for many years and have considerable experience 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 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 that includes mints, snapdragons, foxgloves, speedwells, cladistic studies brought them to light. Details of this taxonomic plantains, gesneriads, shrimp-plants, and trumpet 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 Jacaranda 1/110 Phyrmaceae Phryma 1/1 * Antirrhinum 4/220 Penstemon Scrophularia 3/350 Avicenniaceae 1/1 Lamiaceae s. l. 6/230 Figure 1. Portion of of Lamiaceae s. str. 3/170 jamaicensis, illustrating family characteristics mentioned in text. Photo by J.M. Garg, Wikimedia ex Verbenaceae s. l. 3/60 Commons. Verbenaceae s. str. 23/35 Petreeae 2/2 diversification, extent of design in homoplasy and its role in post- Citharexyleae 4/8 Flood adaptation. Coelocarpum Raphithamnus Methods Verbeneae 13/16 Taxa. A total of 44 genera in nine families are sampled in Casselia the analysis. The ingroup includes 25 of the 35 genera and all tribes of the Verbenaceae. Seven other families of Lamiales, represented by 17 genera, were selected as near outgroups (or extended ingroup). Based on cladistic morphological studies Lampaya (Cantino 1982) and molecular systematics (Soltis et al. 2005), Pitraea the Boraginaceae (Boraginales or Solanales) were chosen as the 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 6/9 secondary chemistry (3 characters), general growth form (1 Goniostachyum character), stem morphology (1), epidermal anatomy (5), morphology (2), inflorescence morphology (6), morphology (26), floral anatomy (8), anatomy (4), embryology (16), 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=; 1=; 2=; 3=; 4=; 30. 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. 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=/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 with long-elliptic depres- 2=Stachytarpheta type; 3=Bouchea type; 4=Priva type; sions for calyces; 2=clavate, spongy, with rounded 5=Petrea type; 6=Acanthaceae type; 7=Martynia type; impressions for flower bases. 8=Borage type. 15. Inflorescence , shape:0=±subulate/linear; 1=el- 43. , no. carpels: 0=2; 1=4. liptic/ovate; 2=rotund/reniform; 3=foliar. 44. Gynoecium, no. functional carpels: 0=2; 1=1 (adaxi- 16. Inflorescence bracts, persistence:0=persistent in fruit; al); 2=4. 1=caducous in fruit. 45. Gynoecium, no. locules: 0=2; 1=4; 2=1; 3=8. 17. Flower arrangement: 0=altermate; 1=decussate; 46. Carpel margin: 0=not intruded; 1=intruded forming 2=whorled; 3=decussate dichasia; 4=alternate mono- false septa & fused completely to carpel wall; 2=in- chasial. truded forming false septa & fused to carpel wall only 18. presence: 0=well developed; 1=subsessile; at base & apex; 3=intruded forming false septa & free 2=sessile. from carpel wall. 19. Calyx merosity: 0=5; 1=4; 2=2. 47. Carpel medial wall: 0=not invaginated or intruded; 20. Calyx consistency in flower: 0=sepaloid; 1=intruded forming false septa; 2=partially intruded but 1=paleaceous; 2=petaloid. not the primary source of a false septum. 21. Calyx shape: 0=rotate; 1=tubular; 2=compressed. 48. Style base position: 0=terminal; 1=partially intruded 22. Calyx size: 0=shorter than corolla; 1=about equaling into lobed ovary; 2=gynobasic. corolla; 2=longer than corolla. 49. Style persistence: 0=persistent into post-anthesis; 23. Calyx, post-anthesis growth: 0=none; 1=accrescent, 1=caducous after anthesis; 2=persistent into mature but not greatly enlarging; 2=accrescent and enveloping/ fruit. covering fruit. 50. Style branch course: 0=straight; 1=recurved. 24. Calyx adhesion in fruit: 0=not adherent; 1=adherent. 51. Style branch insertion: 0=terminal; 1=lateral 25. Calyx substance in fruit: 0=herbaceous; 1=withering; (oblique). 2=membranous & fragile, often fragmenting; 3=thick- 52. Style branches, relative size: 0=branches equal; ened, succulent; 4=corneous. 1=adaxial longer; 2=abaxial longer. 26. Calyx lobes, orientation: 0=divergently spreading; 53. Style branch shape: 0=terete/globose; 1= branch(es) 1=± erect. flattened. 27. Calyx lobes, shape: 0=deltate; 1=oblong; 2=indupli- 54. Style branches, posture: 0=divaricate; 1=ascending; cate -like; 3=irregular/erose; 4=subulate/terete. 2=fused into globular mass; 3=fused into funnel. 28. Calyx lobes, size: 0=equaling tube; 1=shorter than 55. , presence of stigmatoid tissue: 0=not modify- tube; 2=longer than tube. ing style-branch structure; 1=forming glandular mass 29. Calyx lobes, venation: 0=reticulate veined; on abaxial branch; 2=forming glandular mass on both 1=1-veined; 2=3-veined; 3=5-9 parallel-veined. branches.

JCTS B: Life Sciences www.coresci.org/jcts Volume 6:83 56. no. per carpel: 0=2; 1=4; 2=1; 3=many. 68. Chalazal haustoria : 0=developed ; 1=not developed. 57. Ovule, endothelium: 0=completely lining integument; 69. Microphylar haustoria: 0=developed; 1=not devel- 1=lacking from micropylar end; 2=absent. oped; 2=strongly developed as large bulb; 3=strongly 58. Ovule, nucellus: 0=tenuinucellar; 1=crassinucellar. developed as branched mycelioid. 59. Ovule attachment: 0=margin of true septa; 1=margin 70. Embryo axis: 0=straight; 1=curved. of false septa; 2=lateral faces of false septa; 3=enlarged 71. Embryo shape: 0=spatulate; 1=invested. placenta on true septum. 72. Endosperm: 0=all or most used up; 1=a distinct fleshy 60. Ovule orientation/position: 0=pendulous/apical; 1=in- layer. termediate; 2=erect/basal. 73. Endo-/Pericarp cohesion: 0=separating in fruit; 61. Ovule structure: 0=orthotropous; 1=hemianatropous; 1=connate in fruit. 2=anatropous; 3=campylotropous. 74. Endo-/Pericarp ornamentaion: 0=none; 1=reticulate; 62. Ovule, raphe orientation: 0=epitropous; 1=apo- 2=striations only; 3=muricate/tuberculate. tropous; 2=not applicable; 3=variable. 75. Endo-/Pericarp, commissure walls: 0=flat; 1=con- 63. Funiculus origin: 0=whole marginal carpel bundle; cave; 2=not applicable. 1=branch of marginal carpel bundle. 76. Fruiting unit: 0=2 fused 1/2 carpels; 64. Funiculus fusion: 0=free from wall; 1=fused to near 1=single carpel; 2=1, 1/2 carpel; 3=4 fused 1/2 carpels. apex to wall. 77. Fruit type: 0=single drupe; 1=schizocarp; 2=schizocar- 65. Obturator (extension of septum basal margin): pic drupe/multi-drupes; 3=indehiscent nut; 4=. 0=not developed; 1=developed. 78. Fruit apex: 0=truncate/rounded; 1=emarginate; 2=long, 66. Embryology type: 0=onograd; 1=asterad or chenopo- tapered rostrate; 3=short abrupt rostrate. diad. 79. Mesocarp: 0=lacking (dry); 1=leathery; 2=fleshy; 67. Micropyle position: 0=up; 1=down; 2=intermediate or 3=juicy. variable. 80. Chromosome base no.: 0=5; 1=6; 2=7; 3=8/9; 4=11/12; 5=17; 6=19.

Sources covering many characters in many taxa: Atkins 2004; Cantino 1982, 1992a; Cronquist 1981; Dahlgren 1975; Harley et al. 2004; Sanders 2001, unpublished personal observations; Troncoso 1974; Wagenitz 1992; Watson & Dallwitz 2000. Character sets and/or limited taxa indicated in parentheses after citations: Abu-Asab & Cantino 1987 (3-7), 1992, 1993, 1994 (39-42), 2004 (Phryma); Ahmad 1978 (3-6 Acanthaceae); Bhandari et al. 1976 (43-72 Scrophularia); Bonzani et al. 2007 (3-7 Mentha); Buurman 1977 (39-42 Bignoniaceae); Cantino 1990 (3-7), 2004 (Phryma); Chadwell et. al 1992 (39-42 Phryma); Chuang & Heckard 1991 (62 Scrophulariaceae); Cooper 1941 (43-79 Phyrma); Daniel 1998 (39-42 Acanthaceae), 2000 (80 Acanthaceae), 2006 (80 Acanthaceae); Dasti et al. 2003 (7 Boraginaceae); Díez 1994 (39-42 Anchusa); Erdtman 1945, 1952 (39-42); Fischer 2004 (Scrophulariaceae); Fischer et al. 2004 (Bignoniaceae); Galati & Strittmatter 1999 (56-71 Jacaranda); Gentry & Tomb 1979 (39-42 Bignoniaceae); Goldblatt & Gentry 1979 (80 Bignoniaceae); Guşuleac 1937 (62); Hilger et al. 2004 (39, 42, 80 Anchusa); Ihlenfeldt 2004 (Proboscidea); Jensen et al. 1975 (9); Junell 1934 (43-72); Lawrence 1937 (30-32, 43-65 Boraginaceae); Long 1970 (Acanthaceae); McDade et al. 2008 (39-42 Acanthaceae), 2009 (Acanthaceae); Mifsud 2010 (39-42 Anchusa); Mohan Ram & Wadhi 1965 (43-47, 56-72 Acanthaceae); Nowicke & Ridgway 1973 (39-42 Cordia); Schrock & Palser 1967 (29, 31, 32, 43-72 Scrophulariaceae); Raj 1983 (39-42); Ramana et al. 1983 (18, 19, 23, 29, 30-32, 43-65, 67 Phryma); Roubik 2003 (39-42); Rueda 1994 (Petrea & Xolocotzia); Siedo 2006 (51, 80 Aloysia); Souza 2008(43-79 Cordia); Stewart & Canne-Hilliker 1998 (62 Scrophulariaceae); Thieret 1972 (Phryma), 1977 (Proboscidea); Tieghem 1907 (56-71); Trigo 1993 (39-42); Ventrella & Marinho 2008 (3-6 Cordia); Whipple 1972 (Phryma). limited character sets or limited taxa or both and are cited in Table Results 2. Characters were chosen to ensure variation among genera of Verbenaceae and between genera of Verbenaceae and outgroups, All 80 characters and all taxa had a relevance value greater than as well as fixed differences between Verbenaceae and outgroups. 0.95. The baraminological distance correlations resulted in three It is my belief that apportioning the variation into these characters distinct clusters, the Verbenaceae, the remaining Lamiales, and captures any significant differences and similarities among the the Boraginaceae (Figure 2). The Boraginaceae were partially Verbenaceae and similar families. Furthermore, I argue that these negatively correlated (none with bootstrap values >90%) with 80 characters provide a holistic assessment of variation. the Verbenaceae and neither positively nor negatively correlated Analysis. The data matrix (44 taxa × 80 characters; Table 3) was with the other Lamiales, except for some negative correlation analyzed using the BDISTMDS program (Robinson & Cavanaugh with Phyrma. The Lamiales consisted of two subclusters, the 1998; Wood 2005, 2008a) to obtain baraminic distances (BDIST), Lamiaceae and the remaining families (“scrophularian families”), baraminic distance correlations (BDC), correlation bootstrap which were united by positive correlations (none with bootstrap values, and multidimensional scaling coordinates (MDS) for all values >90%) among four pairs of genera. The Lamiaceae were pairs of taxa. The coordinates were displayed using the MAGE not positively correlated with the Verbenaceae, and only negatively software package (Richardson & Presley 2006). correlated with Petrea and Xolocotzia (none with bootstrap values >90%). The scrophularian families (except the Phyrmaceae and Avicenniaceae) were nearly consistently negatively correlated

JCTS B: Life Sciences www.coresci.org/jcts Volume 6:84 Table 3. Data matrix used in the analysis of the Verbenaceae. Letters indicate different combinations of states in variable taxa.

Genus 00000000011111111112222222222333333333344444444445555555555666666666677777777778 12345678901234567890123456789012345678901234567890123456789012345678901234567890

Petrea G101011000002010100202114012011A100110010501030000000B20101001001010000010010015 Xolocotzia 11010A10000020101002021140120110100110010501030000000B20101001001010000010010015 Citharexylum A1000100000020A01000101041011111110110010100110000000120101221001010000000000026 Raphithamnus A1000B00000020001000101131L11111110010010100110000021110101221001010000100000023 Duranta A1A00F00000020011000102131011111110110010112310000000020101221001010000000000023 Coelocarpum 51000000000020001100101001011110110111010100110000020110101221001010000000002023 Casselia K1000010000020000100101001011110110111010100120000100F20101221111010000001032113 Tamonia K1000010000020000100101001411110110111010101120000A00F20101221111010000011000113 Dipyrena 1100001000F020000100101011011110110001010100110000020B20101221001010000001002313 Pitraea 2102001000002000210010200141111111011101010011000102111010122100101000000A102311 Diostea 100000100000200022001010013111111A0000010101010001100220101221001010000100022010 Lampaya 1100A0100000201002001020010111101A0000010101010000020110101221001010000001122123 Bouchea 21000010000021000200101001011110100110010301010001021110101221001010000000121200 Stachytarpheta K1A000100000211002A0101001011122120110010201010000000220101221001010000001021203 Priva K10200100000200001A010200101111011011101040AA10000021110101221001010000001101001 Neosparton 2000001000023011?1001010010111111000?0010101010000A002201012210010100001110110J3 Glandularia 2100A0A0000A200002A0102001411110100011010100110000010110101221001010000001021100 Junellia H10000000001200002A010F00141111010011A0101001100000B0110101221001010000001021100 Verbena 210000A0000A20000FA01010010111101001100101001100000B01101012210010100000010J1102 Aloysia 11000010000A20013210101001411110100000010101010000A00020101221001010000000021A03 Goniostachyum 11000010000030F032211010B11A1110100000010101010000100020101221001010000000021A01 Riedelia 11000010000030A032211010B11A1110100000010101010000100020101221001010000000121NAC Lippia Q10A0010000030F032212010B122111010000A010101010000100020101221001010000000021A0C Nashia 1101001000003010321110102131111010000001010101000000002010122100101000000002001C Lantana 110AAA100000JBMA32F11021213A111010000001010101000010002010122100101000001MA10CP4 Petitia A1100110000050103110101001012310000000001000030010000000102221000010200010030024 Clerodendrum N1100FA0000010A040001010010A211010000000100011211000010010211100001020000002B124 Holmskioldia 111001A0000010003100002040312210100000001000110010020100102001000010200103021001 Lamium 21100A10000000003100101001002210100000001000112210000100102221000010200100021103 Mentha 2110AA1000000000310010100101231010000010100011221000010010222100001020100A021104 Salvia F110AA100000B000G1001010010122221201001010001122100B010010222100001020100C021104 Avicennia 01100111000020101200101001022G10A10000011600030000000100202002000010310100234313 Thunbergia 4010A0100000201010001000013B3110110000421600000000001300200220000010000010214302 Justicia 2010A2100000D0103100100001D0222011001001160000000000010B200220000010000010214302 Elytraria 2010A21000B020101100100001422210100000011600100000000103000220000010000110214304 Antirrhinum K0101310003020G00100101001022211120000001000000020100223103123000020000110214303 Scrophularia K110131000CA500030001000010B2211120000011000000000000223103123000020000110214303 Mimulus 2110131000B02010G0001020210122101200000A1000000000001123103123000020000110214302 Penstemon F110A310000000A030001010010B2211120000001000000020000223103123000020000110214303 Phyrma 21101000000020001100102041411210110000001001200000010122100211001010000010213002 Jacaranda A0100210000150013000100011012211120000001000000000001123003223000010000010214003 Proboscidea 20101110000020211000100011012211120000422700030000001023101120000000000013114213 Anchusa 20A0000111F040100100101001022000001000C10800001100000100001110000101A00001121103 Cordia A0A010011110G001400010F0413120000010000A0800001000001100011110000100000101020023

A=01 B=02 C=03 D=04 E=05 F=12 G=13 H=15 J=23 K=25 L=34 M=012 N=013 P=123 Q=125

(10% with bootstrap values >90%) with the Verbenaceae. vertices, respectively. The Verbenaceae were separated from the All genera of the Verbenaceae were positively correlated (82% Lamiaceae, the nearest outgroup, by distances of 0.250 (Verbena– with bootstrap values >90%) with each other except for Petrea and Mentha) and 0.275 (Verbena –Salvia), yielding correlations of only Xolocotzia with Glandularia, Pitraea, Priva, and Verbena. These 0.246 and 0.230, respectively. Within the Verbenaceae, a distance two sets of genera represent the ends of a variation spectrum in of 0.438 separated the variational extremes of Pitreae from Petrea the family. and Xolocotzia (correlation of 0.298), whereas adjacent members Results of the MDS analysis are given in Figure 3. The stress in of the Verbenaceae were separated by a maximum distance of dimension 3 was 0.300 with minimum stress of 0.048 in dimension 0.250 (Lantana- Xolocotzia) with a correlation of 0.701. 11. In the three-dimensional projection of the MDS results, the Verbenaceae were separated in a planar array on one side. On Discussion the opposite side of the origin, the outgroups formed a triangular planar array with the Boraginaceae well separated at one vertex These results support the holobaraminic status of the and the Lamiaceae and scrophularian families at the other two Verbenaceae s. str. and previous baraminological studies

JCTS B: Life Sciences www.coresci.org/jcts Volume 6:85 Figure 2. Baraminic distance correlation (BDC) bootstrap results for the Verbenaceae, as calculated by BDISTMDS (relevance cutoff 0.95). Closed squares indicate significant, positive BDC; open circles indicate significant, negative BDC. Black symbols indicate bootstrap vales >90% in a sample of 100 pseudoreplicates. Grey symbols represent bootstrap values ≤90%. frequently placing the holobaramin boundary at the conventional str. would happen, especially due to lack of critical anatomical taxonomic rank of family (Wood 2008b). It is of interest that characters in previous classifications, given the distances of the these results detected the discontinuity between the Verbenaceae Lamiaceae and Verbenaceae in the present data. Furthermore, s. str. and genera previously placed in the Verbenaceae s. l. these data also clearly distinguished the segregate families (represented by Clerodendrum, Holmskioldia, Petitia in this Avicenniaceae and Phyrmaceae, which have been included in the study), a distinction which has been recognized widely only Verbenaceae widely in past classifications. since 1992 on the basis of several morphological and molecular Even though a previous morphological cladistics analysis (and cladistic analyses (Cantino 1992a, 1992b; Wagstaff & Olmstead hence morphological data set) did not exit (before the present 1997; Wagstaff et al. 1998; Harley et al. 2004; Soltis et al. 2005). research was completed in 2011; see Sanders 2010), O’Leary et al That is, as long as the correct characters are included, the present (2012) mapped 21 morphological characters onto their molecular baraminological phenetic methods retrieve the separation without phylogeny of the Verbenaceae. They optimized the placement relying on cladistic methods or biomolecular data. of the characters to minimize the evolutionary steps without BDISTMDS also clearly separates the Verbenaceae from the altering the molecular results. All their characters are included traditionally conceived Lamiaceae (once thought to be derived in my character assignments except styles long vs. short and from or share a most recent common ancestor with Verbenaceae; inflorescence of terminal vs. axillary vs. terminal plus axillary Dalhgren 1975; Cronquist 1981; Sanders 2001). However, it is thyrses. They found the inflorescence character to be the most easy to see how former misalignments among the Verbenaceae homoplastic of their characters. I had intuitively noticed that and s. str., former members of Verbenaceae s. l., and Lamiaceae s. did not bother to distinguish the character states as such. They

JCTS B: Life Sciences www.coresci.org/jcts Volume 6:86 Figure 3. Three-dimensional MDS applied to Verbenaceae baraminic distances and the stress of k-dimensional MDS on the same baraminic distance matrix plotted as a function of the number of dimensions (k). Members that are labeled are mentioned in the text. also found style length to be homoplastic and mostly differentiated than some of economic or agricultural importance) survived genera within Verbeneae. Likewise, I had concluded that the outside the ark and have an unknown number and diversity of character was too variable to be of importance for baraminology. survivors. Other criteria need to be developed to address these The present research supports the position that ancestral questions. Verbenaceae s. str. as a baramin, were created on Creation Day On the other hand, data that support the baramin at the family 3. Sanders (2013) also published data on the history of level (or possibly higher level) can help begin to unravel natural flowering plants that appears to support that conclusion. However, evil in the form of prickles and some chemical defenses. This the fossil data are less clear. Sanders based the pre-Flood fossil is because prickles and liver toxins are limited to a few closely record of the Verbenaceae on the report in the online version of related species within the Lantana, which now is seen as Benton’s The Fossil Record 2 (Benton & Benton (1993-2006), a very small subset of the entire baramin. This would suggest which simply lists the family name, leaf material, and the range that these traits developed only during post-Flood diversification. in the geologic column. Therefore, Sanders could not verify Volatile compounds as defensive natural evil do not characterize the identification of the cited material or know if it belongs the whole family, but do characterize one whole tribe, the to Verbenaceae s.str. or to a genus that has been transferred to Lantaneae. The molecular phylogenies cited above (Marx et Lamiaceae s.l. If the material that the Bentons cite belongs to al. 2010; Yuan et al. 2010; O’Leary et al. 2012; Lu-Irving & Lamiaceae rather than Verbenaceae, then Sanders’ work would Olmstead 2013) all support the Lantaneae as a late development suggest that the Verbenaceae s. str. is subbaraminic. in the Verbenaceae accompanied by a rapid diversification of However, these data do not resolve whether any modern genus or species. In fact, Lu-Irving and Olmstead (2013) substantiate the even tribe was part of the Verbenaceae archaebaramin or whether long-held suspicion that the core genera of the Lantaneae are not they, instead, originated as part of the post-Flood (Cenozoic) monophyletic (see Sanders 2001). Rather the species of the tribe diversification of the Verbenaceae. This is because plants (other cannot be easily placed in genera and appear to have originated

JCTS B: Life Sciences www.coresci.org/jcts Volume 6:87 in a starburst pattern as predicted by Wood and Murray (2003, implications. In: Harley, R.M. and T. Reynolds, eds. Advances p.174). in Labiate Science. Royal Botanic Gardens, Kew, U.K., pp. 97- Sanders (2001, 2012) noted that the prevalence of homoplasy 112. in Lantana and the whole family prevented the application of Abu-Asab, M.S. and P.D. Cantino. 1993. Phylogenetic cladistics of morphological data. O’Leary et al. (2012) and Lu- implications of pollen morphology in tribe Ajugeae (Labiatae). Irving and Olmstead (2013) substantiated the morphological Systematic 18:100-122. homoplasy using molecular data. The traits used to delimit Abu-Asab, M.S. and P.D. Cantino. 1994. Systematic species, genera, and even tribes pop up erratically on different implications of pollen morphology in subfamilies lineages. These data along with the conclusion of the previous and Pogostemonoideae (Labiatae). Annals of the Missouri paragraph suggest that this homoplasy is a result of rapid post- Botanical Garden 81:653-686. Flood diversification rather than interbaraminic modularity or Ahmad, K.J. 1978. Epidermal hairs of Acanthaceae. Blumea pre-Flood design to fit pre-Flood environments. 24:101-117. There is one other line of evidence to suggest that the origin Atkins, S. 2004. Verbenaceae. In: Kubitzki, K. and J.W. of current species of the Verbenaceae starburst was post-Flood, Kadereit, eds. The Families and Genera of Vascular Plants, perhaps late post-Flood about the time of the Pleistocene Ice Volume VII:Lamiales (excluding the Acanthaceae including Advance (reckoned here as much as a few hundred years after Avicenniaceae). Springer Verlag, Berlin, pp. 449-468. the Flood). Olmstead (2013) used a molecular phylogeny to Benton, M, and M.J. Benton. 1993-2006. The Fossil Record analyze transoceanic disjunctions in the Verbenaceae. There are Website, Accessed November 25, 2012 at http://www. six disjunctions from the New World to the Old World, and five fossilrecord.net/fossilrecord/index.html. Based on Benton, M. are in the starburst section of the family. Four of these resulted J. (ed.) 1993. The Fossil Record 2. London, England:Chapman in one (in one case two) species disjunct, with the sister species & Hall. remaining in the New World. Thus, these disjunctions appear to Bhandari, N.N., F. Bouman, and S. Natesh. 1976. Ovule ontogeny be recent long-distance dispersals across the ocean each followed and coat structure of Scrophularia himalensis Royle. by the origin of one to two species. Botanische Jahrbücher für Systamatik, Pflanzengeschichte und Therefore, I present the hypothesis that, at least, the current Plfanzengeographie 95:535-548. species of the tribe Lantaneae are all the result of rapid post-Flood Bonzani, N.E., M. Costaguta, and G.E. Barboza. 2007. Estudios diversification. As a corollary, the defensive liver toxins and anatómicos en especies de Mentha (Fam. Lamiaceae) de prickles in Lantana are a post-Flood development from genetic Argentina. Arnaldoa 14:77-96. potential provided at creation but not expressed until well after Buurman, J. 1977. Contribution to the pollen morphology of the Fall and the Flood. I will seek additional criteria by which this the Bignoniaceae, with special reference to the tricolpate type. hypothesis can be tested. Pollen et Spores 19:447-520. Cantino, P.D. 1982. Affinities of the Lamiales. Systematic Conclusion Botany 7:237-248. Cantino, P.D. 1990. Phylogenetic significance of stomata and This study is a first attempt of statistical baraminology to resolve in the Labiatae and Verbenaceae. Journal of the the baraminic status of the Verbenaceae and the first attempt to Arnold Arboretum 71:323-370. assemble a data set from personal research for a baraminological Cantino, P.D. 1992a. Evidence for a polyphyletic origin of the analysis of a plant group. The results support the holobaraminic Labiatae. Annals of the Missouri Botanical Garden 79:361- status of the Verbenaceae s.str. They are also suggestive that the 379. current species of the tribe Lantaneae, centered around Lantana, Cantino, P.D. 1992b. Toward a phylogenetic classification of the diversified during the post-Flood recovery, and that many features Labiatae. In: Harley, R.M. and T. Reynolds, eds., Advances in of natural evil in these plants is a form of recently developed Labiate Science. Royal Botanic Gardens, Kew, U.K., pp. 27-37. mediated design. It is hoped that future studies will provide Cantino, P.D. 2004. . In: Kubitzki, K. and J.W. further insights into the timing and extent of diversification in this Kadereit, eds. The Families and Genera of Vascular Plants, interesting plant family. Volume VII:Lamiales (excluding the Acanthaceae including Avicenniaceae). Springer Verlag, Berlin, pp. 323-326. Acknowledgments Chadwell, T.B., S.J. Wagstaff, and P.D. Cantino. 1992. Pollen morphology of Phryma and some putative relatives. Systematic Todd C. Wood provided encouragement and critical comments Botany 17:210-219. on the drafts of the paper. Chuang, T.I. and L.R. Heckard. 1991. Generic realignment and synopsis of subtribe Castillejinae (Scrophulariaceae-Tribe References Pediculareae). Systematic Botany 16:644-666. Cooper, D.C. 1941. Macrosporogenesis and the development of Abu-Asab, M.S. and P.D. Cantino. 1987. Phylogenetic the seed of . American Journal of Botany implications of leaf anatomy in subtribe Melittidinae (Labiatae) 28:755-761. and related taxa. Journal of the Arnold Arboretum 68:1-34. Cronquist, A. 1981. An Integrated System of Classification of Abu-Asab, M.S. and P.D. Cantino. 1992. Pollen morphology Flowering Plants. Columbia University Press, New York, NY. in subfamily Lamioideae (Labiatae) and its phylogenetic Dahlgren, R. 1975. A system of classification of the angiosperms

JCTS B: Life Sciences www.coresci.org/jcts Volume 6:88 to be used to demonstrate the distribution of characters. 4:1-219. Botaniska Notiser 128:119-147. Lawrence, J.R. 1937. A correlation of the and the floral Daniel, T.F. 1998. Pollen morphology of Mexican anatomy of certain of the Boraginaceae. American Journal of Acanthaceae:Diversity and systematic significance. Botany 24:433-444. Proceedings of the California Academy of Sciences 50:217-256. Long, R.W. 1970. The genera of Acanthaceae in the Southeastern Daniel, T.F. 2000. Additional chromosome numbers of American United States. Journal of the Arnold Arboretum 51:255-309. Acanthaceae. Systematic Botany 15-25. Lu-Irving, P. and R.G. Olmstead. 2013. Investigating the Daniel, T.F. 2006. Chromosome numbers of miscellaneous evolution of Lantaneae (Verbenaceae) using multiple loci. Malagasy Acanthaceae. Brittonia 58:291-300. Botanical Journal of the Linnean Society 171:103-119. Dasti, A.A., T.Z. Bokhari, S.A. Malik, and R. Akhtar. 2003. Marx, H.E. and six others. 2010. A molecular phylogeny and Epidermal morphology in some members of family classification of Verbenaceae. American Journal of Botany Boraginaceae in Baluchistan. Asian Journal of Plant Sciences 97:1647-1663. 2:42-47. McDade, L.A, T.F. Daniel, and C. Kiel. 2008. Toward a Díez, M.J. 1994. A general survey of pollen types in Anchusa L. comprehensive understanding of phylogenetic relationships (Boraginaceae) in relation to taxonomy. Acta Botanica Gallica among lineages of Acanthaceae s.l. (Lamiales). American 141:233-242. Journal of Botany 95:1136-1152. Erdtman, G. 1945. Pollen morphology and plant taxonomy. McDade, L.A., C.Kiel, and E. Tripp. 2009. Acanthaceae. IV:Labiatae, Verbenaceae, and Avicenniaceae. Svensk Botanisk Version 15 November 2009. http://tolweb.org/ Tidskrift 39:279-285. Acanthaceae/20878/2009.11.15 in The Tree of Life Web Erdtman, G. 1952. Pollen Morphology and Plant Project, http://tolweb.org. Accessed 21 September 2011. Taxonomy:Angiosperms. Almquist and Wiksell, Stockholm. Mifsud, S. 2010. Encyclopaedia of the Wild Plants of Malta. Fischer, E. 2004. Scrophulariaceae. In: Kubitzki, K. and J.W. http://www.maltawildplants.com (2002-2010). Accessed 23, Kadereit, eds. The Families and Genera of Vascular Plants, September 2011. Volume VII:Lamiales (excluding the Acanthaceae including Mohan Ram, H.Y. and M. Wadhi. 1965. Embryology and the Avicenniaceae). Springer Verlag, Berlin, pp. 333-432. delimitation of the Acanthaceae. Phytomorphology 15:201- Fischer, E., I. Theisen, and L.G. Lohmann. 2004. Bignoniaceae. 205. In: Kubitzki, K. and J.W. Kadereit, eds. The Families and Nowicke, J.W. and J.E. Ridgway. 1973. Pollen studies in the Genera of Vascular Plants, Volume VII:Lamiales (excluding genus Cordia (Boraginaceae). American Journal of Botany the Acanthaceae including Avicenniaceae). Springer Verlag, 60:584-591. Berlin, pp. 9-38. O’Leary, N., C.I. Calviño, S. Martínez, P. Lu-Irving, R.G. Olmstead, Galati, B.G. and L.I. Strittmatter. 1999. Ovule ontogeny and M.E. Múlgura. 2012. Evolution of morphological traits in and megasporogensis in Jacaranda mimosifolia D.Don. Verbenaceae. American Journal of Botany 99:1778-1792. (Bignoniaceae). Phytomorphology 49:67-74 Olmstead, R.G. 2013. Phylogeny and biogeography in Solanaceae, Gentry, A.H. and A.S. Tomb. 1979. Taxonomic implications of Verbenaceae and Bignoniaceae:a comparison of continental Bignoniaceae palynology. Annals of the Missouri Botanical and intercontinental diversification patterns. Botanical Journal Garden 66:756-777. of the Linnean Society 171:80-102. Goldblatt, P and A.H. Gentry. 1979. Cytology of Bignoniaceae. Raj, B. 1983. A contribution to the pollen morphology of Botaniska Notiser 132:475-482. Verbenaceae. Review of Palaeobotany and Palynology 39:343- Guşuleac, M. 1937. Über die Orientierung des Ovulums bei den 422. Boraginaceen and Labiaten, nebst Ausblicken auf das System Ramana, R.V., P.S.P. Rao, and L.L. Narayana. 1983. A contribution dieser Familien. Published by the author. to the floral anatomy ofPhyrma leptostachya. Current Science Harley, R.M. and 12 others. 2004. Labiate. In: Kubitzki, K. 52:922-924. and J.W. Kadereit, eds. The Families and Genera of Vascular Richardson, D.C. and B.K. Presley. 2006. MAGE software, v. Plants, Volume VII:Lamiales (excluding the Acanthaceae 6.44. Duke University. Distributed by the author. including Avicenniaceae). Springer Verlag, Berlin, pp. 167-275. Robinson, D.A. and D.P. Cavanaugh. 1998. A quantitative Hilger, H.H., F. Selvi, A. Papini, and M. Bigazzi. 2004. Molecular approach to baraminology with examples from the primates. systematics of Boraginaceae tribe Boragineae based on ITS1 Creation Research Society Quarterly 34:196-208. and trnL sequences, with special reference to Anchusa s.l. Roubik, D. 2003. Pollen and Spores of Barro Colorado Island. Annals of Botany 94:201-212. http://www.stri.si.edu/sites/roubik. Accessed 23 September Ihlenfeldt, H.-D. 2004. Martyniaceae. In: Kubitzki, K. and J.W. 2011. Kadereit, eds. The Families and Genera of Vascular Plants, Rueda, R.M. 1994. Systematics and evolution of the genus Volume VII:Lamiales (excluding the Acanthaceae including Petrea (Verbenaceae). Annals of the Missouri Botanical Garden Avicenniaceae). Springer Verlag, Berlin, pp. 283-288. 81:610-652. Jensen, S.R., B.J. Nielsen, and R. Dahlgren. 1975. Iridoid Sanders, R.W. 2001. The genera of Verbenaceae in the compounds, their occurrence and systematic importance in the Southeastern United States. Harvard Papers in Botany 5:303- angiosperms. Botaniska Notiser 128:148-180. 358. Junell, S. 1934. Zur Gynäceummorphologie und Systematic der Sanders, R.W. 2010. Baraminic status of the Verbenaceae Verbenaceen and Labiaten. Symbolae Botanicae Upsalienses (verbena family). Occasional Papers of the BSG 17:4-5.

JCTS B: Life Sciences www.coresci.org/jcts Volume 6:89 Sanders, R.W. 2012. Taxonomy of Lantana sect. Lantana extratropical. Darwiniana 18:295-412. (Verbenaceae):II. Taxonomic revision. Journal of the Botanical Ventrella, M.C. and C.R. Marinho. 2008. Morphology and Research Institute of Texas 6:403-441. histochemistry of glandular trichomes of Cordia verbenacea Sanders, R.W. 2013. The fossil record of angiosperm families in DC. (Boraginaceae) leaves. Revista Brasileira de Botanica relation to baraminology. In: Horstemeyer, M., ed. Proceedings 31:457-467. of the Seventh Conference on Creationism. Creation Science Wagenitz, G. 1992. The Asteridae:Evolution of a concept and Fellowship, Pittsburgh, Pennsylvania. its present status. Annals of the Missouri Botanical Garden Schrock, G.F. and B.F. Palser. 1967. Floral development, 79:209-217. anatomy, and embryology of heterophylla with some Wagstaff. S.J. and R.G. Olmstead. 1997. Phylogeny of Labiatae notes on ten other species of Collinsia and on tenella. and Verbenaceae inferred from rbcL sequences. Systematic Botanical Gazette (Chicago) 128:83-104. Botany 22:165-179. Siedo, S.J. 2006. Systematics of Aloysia (Verbenaceae). Wagstaff. S.J., L.Hickerson, R. Spangler, P.A. Reeves, and R.G. Unpublished dissertation, University of Texas, Austin. Olmstead. 1998. Phylogeny in Labiatae s.l., inferred from Soltis, D.E, P.S. Soltis, P.K. Endress, and M.W. Chase. 2005. cpDNA sequences. Plant Systematics and Evolution 209:265- Phylogeny and Evolution of Angiosperms. Sinauer Associates, 274. Sunderland, MA. Watson, L. and M. J. Dallwitz. 2000. The Families of Flowering Souza, L.A. de. 2008. Morphology and anatomy of the Cordia Plants:Descriptions, Illustrations, Identification, and trichotoma diaspore (Boraginaceae). Brazilian Archives of Information Retrieval. Version:14th December 2000. http:// Biology and Technology 51:761-768. www.biologie.uni-hamburg.de/b-online/delta/angio. Accessed Stewart, H.M. and J.M. Canne-Hilliker 1998. Floral development Jan 15 2010 to Sept 30 2011. of neoscotica, var. borealis, Whipple, H.L. 1972. Structure and systematics of Phyrma and Agalinis purpuea (Scrophulariaceae):Implications for leptostachya L. Journal of the Elisha Mitchell Scientific Society taxonomy and mating system. International Journal of Plant 88:1-17. Science 159:418-439. Wood, T.C. 2005. Visualizing baraminic distances using classical Thieret, J.W. 1972. The Phrymaceae in the Southeastern United multidimensional scaling. Origins (GRI) 57:9-29. States. Journal of the Arnold Arboretum 53:226-233. Wood, T.C. 2008a. BDISTMDS software, v. 1.0. Core Academy Thieret, J.W. 1977. The Martyniaceae in the Southeastern United of Science. Distributed by the author. States. Journal of the Arnold Arboretum 58:25-39. Wood, T.C. 2008b. Animal and plant baramins. CORE Issues in Tieghem, P. van. 1907. Structure du pistil et du fruit des Labiées, Creation 3:1-258. des boragacées et des familles voisines. Annales des Sciences Wood, T.C. and M.J. Murray. 2003. Understanding the Pattern naturelles, Botanique, Série 9 5:321-350. of Life:Origins and Organization of the Species. Broadman and Trigo, M.M. 1993. Contribucíon al estudio polínico de especies Holman Publ., Nashville, Tennessee. ornamentales:Acanthaceae y Verbenaceae. Acta Botanica Yuan, Y., C. Lui, H.E. Marx, and R.G. Olmstead. 2010. An Malacitana 18:135-146. empirical demonstration of using pentatricopeptide repeat (PPR) Troncoso, N.S. 1974. Los géneros de Verbenáceas de Sudamérica genes as plant phylogenetic tools:Phylogeny of Verbenaceae and the Verbena complex. Molecular and Evolution 54:23-35.

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