Free Standard AU & NZ Shipping For All Book Orders Over $80!
Register      Login
Australian Systematic Botany Australian Systematic Botany Society
Taxonomy, biogeography and evolution of plants
L. A. S. JOHNSON REVIEW

Review of the systematics of Scrophulariaceae s.l. and their current disposition

David C. Tank A , Paul M. Beardsley B , Scot A. Kelchner B and Richard G. Olmstead A C
+ Author Affiliations
- Author Affiliations

A Department of Biology, Box 355325, University of Washington, Seattle, WA 98195-5325, USA.

B Department of Biological Sciences, Box 8007, Idaho State University, Pocatello, ID 83209-8007, USA.

C Corresponding author. Email: olmstead@u.washington.edu

Australian Systematic Botany 19(4) 289-307 https://doi.org/10.1071/SB05009
Submitted: 14 April 2005  Accepted: 3 January 2006   Published: 25 August 2006

Abstract

Recent molecular phylogenetic studies in Lamiales have shown that the large group traditionally recognised as Scrophulariaceae is not monophyletic. Efforts to reconstruct the phylogeny of this large clade and to revise its classification to reflect that phylogeny have resulted in seven monophyletic groups, comprised mostly of members of Scrophulariaceae s.l., recognised as families in recent angiosperm classifications. These are Scrophulariaceae s.s., Orobanchaceae, Veronicaceae (cf. Plantaginaceae), Phrymaceae, Calceolariaceae, Linderniaceae, and Stilbaceae. Sampling completeness at the genus level varies from group to group, but is quite good for many. A few individual genera formerly assigned to Scrophulariaceae do not fit into any existing clade recognised at family rank and are left, at present, unassigned to family. In addition to the recognition of several clades comprised primarily of former members of Scrophulariaceae s.l., several groups previously recognised as families are now included within some of these clades. For example, Scrophulariaceae s.s. includes Buddlejaceae and Myoporaceae, and Veronicaceae includes Callitrichaceae, Globulariaceae, Hippuridaceae, and Plantaginaceae. The clades now recognised as families often are not easily diagnosed, but in many cases are more consistent with certain functional traits and geographical patterns. Examples include Orobanchaceae, which comprises all of the parasitic plants (hemiparasites and holoparasites) and Scrophulariaceae s.s., which is predominantly a southern hemisphere group.


Acknowledgments

We thank A. Wolfe and P. Kornhall for providing prepublication manuscripts, J. Bennett and S. Mathews for providing unpublished results of their PHYA trees of Orobanchaceae, and three anonymous reviewers whose comments improved this manuscript. This work was supported in part by University of Washington, Biology Department Molecular Systematics Fellowships to PMB and DCT, United States National Science Foundation grants BSR-9107827 and DEB-9509804 to RGO, DEB-0105048 to PMB and RGO, DEB 0412653 to DCT and RGO and a Smithsonian Senior Fellowship to RGO.


References


Albach DC, Chase MW (2001) Paraphyly of Veronica (Veroniceae; Scrophulariaceae): evidence from the internal transcribed spacer (ITS) sequences of nuclear ribosomal DNA. Journal of Plant Research 114, 9–18.
Crossref |
open url image1

Albach DC, Chase MW (2004) Incongruence in Veroniceae (Plantaginaceae): evidence from two plastid and a nuclear ribosomal DNA region. Molecular Phylogenetics and Evolution 32, 183–197.
Crossref | GoogleScholarGoogle Scholar | PubMed | open url image1

Albach DC, Soltis PS, Soltis DE, Olmstead RG (2001) Phylogenetic analysis of asterids based on sequences of four genes. Annals of the Missouri Botanical Garden 88, 163–212.
Crossref |
open url image1

Albach DC, Martinez-Ortega M, Fischer MA, Chase MW (2004a) Evolution of Veroniceae: a phylogenetic perspective. Annals of the Missouri Botanical Garden 91, 275–302. open url image1

Albach DC, Martinez-Ortega M, Fischer MA, Chase MW (2004b) A new classification of the tribe Veroniceae—problems and a possible solution. Taxon 53, 429–452. open url image1

Albach DC, Meudt HM, Oxelman B (2005a) Piecing together the ‘new’ Plantaginaceae. American Journal of Botany 92, 297–315. open url image1

Albach DC, Utteridge T, Wagstaff SJ (2005b) Origin of Veroniceae (Plantaginaceae, formerly Scrophulariaceae) on New Guinea. Systematic Botany 30, 412–423.
Crossref | GoogleScholarGoogle Scholar | open url image1

Angiosperm Phylogeny Group II (2003) An update of the Angiosperm Phylogeny Group classification for the orders and families of flowering plants: APG II. Botanical Journal of the Linnean Society , 399–436.
Crossref | GoogleScholarGoogle Scholar | open url image1

Archibald JK, Mort ME, Wolfe AD (2005) Phylogenetic relationships within Zaluzianskya (Scrophulariaceae s.s., tribe Manuleeae): Classification based on DNA sequences from multiple genomes and implications for character evolution and biogeography. Systematic Botany 30, 196–215.
Crossref | GoogleScholarGoogle Scholar | open url image1

Backlund M, Oxelman B, Bremer B (2000) Phylogenetic relationships within the Gentianales based on ndhF and rbcL sequences, with particular reference to the Loganiaceae. American Journal of Botany 87, 1029–1043.
Crossref | PubMed |
open url image1

Barker WR (1982) Evolution and biogeography of arid Australian Scrophulariaceae. In ‘Evolution of the flora and fauna of arid Australia’. (Eds WR Barker, PJM Greenslade) pp. 341–350. (Peacock Publications: Adelaide)

Barringer K (1983) Monopera, a new genus of Scrophulariaceae from South America. Brittonia 35, 111–114.
Crossref |
open url image1

Barringer K (1993) Five new tribes in the Scrophulariaceae. Novon 3, 15–17.
Crossref |
open url image1

Beardsley PM, Olmstead RG (2002) Redefining Phrymaceae: the placement of Mimulus, tribe Mimuleae and Phryma. American Journal of Botany 89, 1093–1102. open url image1

Beardsley PM, Barker WR (2005) Patterns of evolution in Australian Mimulus and related genera (Phrymaceae–Scrophulariaceae): a molecular phylogeny using chloroplast and nuclear sequence data. Australian Systematic Botany 18, 61–73.
Crossref | GoogleScholarGoogle Scholar | open url image1

Beardsley PM, Schoenig SE, Whittall JB, Olmstead RG (2004) Patterns of evolution in western North American Mimulus (Phrymaceae). American Journal of Botany 91, 474–489. open url image1

Bellini R (1907) Criteri per una nuova classificazione delle Personatae (Scrophulariaceae et Rhinanthaceae). Annals of Botany 6, 131–145. open url image1

Bello MA, Chase MW, Olmstead RG, Rønsted N, Albach DC (2002) The pàramo endemic Aragoa is the sister genus of Plantago (Plantaginaceae, Lamiales): evidence from plastid rbcL and nuclear ribosomal ITS sequence data. Kew Bulletin 57, 585–597. open url image1

Bennett J, Mathews S (2006) Phylogeny of the parasitic plant family Orobanchaceae inferred from Phytochrome A. American Journal of Botany In press , open url image1

Bentham G (1846) Scrophulariaceae. In ‘Prodromus systematis naturalis regni vegetabilis’. (Ed. A deCandolle) pp. 180–586. (Victoris Masson: Paris)

Bentham G (1876) Scrophularineae. In ‘Genera plantarum’. (Eds G Bentham, JD Hooker) pp. 913–980. (Reeve and Co.: London)

Boeshore I (1920) The morphological continuity of Scrophulariaceae and Orobanchaceae. Contributions of the Botanical Laboratory of the University of Pennsylvania 5, 139–177. open url image1

Bremer B, Olmstead RG, Struwe L, Sweere JA (1994) rbcL sequences support exclusion of Retzia, Desfontainia, and Nicodemia from the Gentianales. Plant Systematics and Evolution 190, 213–230.
Crossref | GoogleScholarGoogle Scholar | open url image1

Bremer K, Backlund A, Sennblad B, Swenson U, Andreasen K, Hjertson M, Lundberg J, Backlund M, Bremer B (2001) A phylogenetic analysis of 100+ genera and 50+ families of euasterids based on morphological and molecular data with notes on possible higher level morphological synapomorphies. Plant Systematics and Evolution 229, 137–169.
Crossref | GoogleScholarGoogle Scholar | open url image1

Bremer B, Bremer K, Heidari N, Erixon P, Olmstead RG, Anderberg AA, Kallersjo M, Barkhordarian E (2002) Phylogenetics of asterids based on 3 coding and 3 non-coding chloroplast DNA markers and the utility of non-coding DNA at higher taxonomic levels. Molecular Phylogenetics and Evolution 24, 274–301.
Crossref | GoogleScholarGoogle Scholar | PubMed | open url image1

Briggs BG (2001) Lawrence Alexander Sidney Johnson 1926–1997. Historical Records of Australian Science 13, 475–494.
Crossref | GoogleScholarGoogle Scholar | open url image1

Brühl P (1920) On the systematic position of Lindenbergia, Lehmann. Journal of the Department of Science, University of Calcutta 2, 11–16. open url image1

Burtt BL (1962) Studies on the Gesneriaceae of the old world XXIV: tentative keys to the tribes and genera. Notes from the Royal Botanic Garden Edinburgh 24, 205–220. open url image1

Carlquist S (1986) Wood anatomy of Stilbaceae and Retziaceae: ecological and systematic implications. Aliso 11, 299–316. open url image1

Carlquist S, Hoekman D (1986) Wood anatomy of Myoporaceae: ecological and systematic considerations. Aliso 11, 317–334. open url image1

Chadwell TB, Wagstaff SJ, Cantino PD (1992) Pollen morphology of Phryma and some putative relatives. Systematic Botany 17, 210–219.
Crossref |
open url image1

Cronquist A (1981) ‘An integrated system of classification of flowering plants.’ (Columbia University Press: New York)

Crosswhite FS, Kawano S (1970) Pennellianthus (Scrophulariaceae)—a new genus of Japan and USSR. American Midland Naturalist 83, 358–367.
Crossref |
open url image1

Dahlgren R (1975) ‘Dicotelydonernes taxonomi: Fabanae–Lamianae.’ (Akademisk forlag: Köbenhavn)

Don G (1838) Scrophularineae. In ‘A general history of the dichladymous plants, comprising complete descriptions of the different orders’. (Eds JG Rivington, F Rivington) pp. 501–640. (Rivington et al.: London)

Fischer E (1992) Systematik der afrikanischen Lindenieae (Scrophulariaceae). Tropische und Subtropische Pflanzenwelt 82, 1–365. open url image1

Fischer E (2004) Scrophulariaceae. In ‘The families and genera of vascular plants’. (Ed. K Kubitzki) pp. 333–432. (Springer: Berlin)

Ghebrehiwet M, Bremer B, Thulin M (2000) Phylogeny of the tribe Antirrhineae (Scrophulariaceae) based on morphological and ndhF sequence data. Plant Systematics and Evolution 220, 223–239.
Crossref | GoogleScholarGoogle Scholar | open url image1

Hallier H (1903) Ueber die abgrenzung und verwandtschaft der einzelnen sippen bei den Scrophulariaceen. Bulletin de l’Herbier Boissier 3, 181–207. open url image1

Hedberg O (1955) A taxonomic revision of the genus Sibthorpia L. Botaniska Notiser 108, 161–183. open url image1

Hempel AL, Reeves PA, Olmstead RG, Jansen RK (1995) Implications of rbcL sequence data for higher-order relationships of the Loasaceae and the anomalous aquatic plant Hydrostachys (Hydrostachyaceae). Plant Systematics and Evolution 194, 25–37.
Crossref | GoogleScholarGoogle Scholar | open url image1

Hilu KW, Borsch T, Müller K, Soltis DE, Soltis PS , et al. (2003) Angiosperm phylogeny based on matK sequence information. American Journal of Botany 90, 1758–1776. open url image1

Hoggard RK, Kores PJ, Molvray M, Hoggard GD, Broughton DA (2003) Molecular systematics and biogeography of the amphibious genus Littorella (Plantaginaceae). American Journal of Botany 90, 429–435. open url image1

Hong DY , Yang H , Jin CL , Holmgren NH (1998) Scrophulariaceae. In ‘Flora of China’. (Eds ZY Wu, PH Raven) pp. 1–212. (Science Press: Beijing)

Hufford L, McMahon M (2004) Morphological evolution and systematics of Synthyris and Besseya (Veronicaceae): a phylogenetic analysis. Systematic Botany 29, 716–736.
Crossref | GoogleScholarGoogle Scholar | open url image1

Judd WS, Olmstead RG (2004) A survey of tricolpate (eudicot) phylogenetic relationships. American Journal of Botany 91, 1627–1644. open url image1

Kelchner SA (2003) Phylogenetic structure, biogeography, and evolution of Myoporaceae. PhD thesis, The Australian National University, Canberra.

Kornhall P, Bremer B (2004) New circumscription of the tribe Limoselleae (Scrophulariaceae) that includes the taxa of the tribe Manuleeae. Botanical Journal of the Linnean Society 146, 453–467.
Crossref | GoogleScholarGoogle Scholar | open url image1

Kornhall P, Heidari N, Bremer B (2001) Selagineae and Manuleeae, two tribes or one? Phylogenetic studies in the Scrophulariaceae. Plant Systematics and Evolution 228, 199–218.
Crossref | GoogleScholarGoogle Scholar | open url image1

Lersten NR, Beaman JM (1998) First report of oil cavities in Scrophulariaceae and reinvestigation of air spaces in leaves of Leucophyllum frutescens. American Journal of Botany 85, 1646–1649.
Crossref |
open url image1

Lersten NR, Curtis JD (2001) Idioblasts and other unusual internal foliar secretory structures in Scrophulariaceae. Plant Systematics and Evolution 227, 63–73.
Crossref | GoogleScholarGoogle Scholar | open url image1

Li HL (1948) A revision of the genus Rehmannia. Tiawania 1, 71–82. open url image1

Lu AM (1990) A preliminary cladistic study of the families of the superorder Lamiiflorae. Botanical Journal of the Linnean Society 103, 39–57. open url image1

Mabberley DJ (1997) ‘The plant-book: a portable dictionary of the vascular plants.’ (Cambridge University Press: Cambridge)

Marloth R (1898) Charadrophila Marloth nov. gen. Botanische Jahrbücher für Systematik und Pflanzengeographie 26, 358–359. open url image1

McDade LA, Moody ML (1999) Phylogenetic relationships among Acanthaceae: evidence from noncoding trnL–trnF chloroplast DNA sequences. American Journal of Botany 86, 70–80.
Crossref |
open url image1

Melchior H (1964) ‘Syllabus der pflanzenfamilien II.’ (Gebrüder Borntraeger: Berlin)

Molau U (1988) Scrophulariaceae—part I. Calceolarieae. Flora Neotropica Monograph 47, 1–325. open url image1

Moldenke HN (1971) ‘A fifth summary of the Verbenaceae, Avicenniaceae, Stilbaceae, Dicrastylidaceae, Symphoremaceae, Nyctanthaceae, and Eriocaulaceae of the world as to the valid taxa, geographic distribution and synonomy.’ (The William Paterson College of New Jersey: Wayne, NJ)

Müller K, Borsch T, Legendre L, Porembski S, Theisen I, Barthlott W (2004) Evolution of carnivory in Lentibulariaceae and the Lamiales. Plant Biology 6, 477–490.
Crossref | GoogleScholarGoogle Scholar | PubMed | open url image1

Nakai T (1949) Classes, ordinae, familiae, subfamilieae, tribus, genera nova quae attinent ad plantas Koreanas. Journal of Japanese Botany 24, 8–14. open url image1

Neizgoda CJ, Tomb AS (1975) Systematic palynology of the tribe Leucophylleae (Scrophulariaceae) and selected Myoporaceae. Pollen et Spores , 495–516. open url image1

Nickrent DL , Duff RJ , Colwell AE , Wolfe AD , Young ND , Steiner KE , dePamphilis CW (1998) Molecular phylogenetic and evolutionary studies of parasitic plants. In ‘Molecular systematics of plants II, DNA sequencing’. (Eds PS Soltis, DE Soltis, JJ Doyle) pp. 211–241. (Kluwer: Boston)

Nie Z-L, Sun H, Beardsley PM, Olmstead RG, Wen J (2006) Evolution of biogeographic disjunction between eastern Asia and eastern North America in Phryma L. (Phrymaceae). American Journal of Botany , open url image1

Olmstead RG, Reeves PA (1995) Evidence for the polyphyly of the Scrophulariaceae based on chloroplast rbcL and ndhF sequences. Annals of the Missouri Botanical Garden 82, 176–193.
Crossref |
open url image1

Olmstead RG, Michaels HJ, Scott KM, Palmer JD (1992) Monophyly of the Asteridae and identification of their major lineages inferred from DNA sequences of rbcL. Annals of the Missouri Botanical Garden 79, 249–265.
Crossref |
open url image1

Olmstead RG, Bremer B, Scott KM, Palmer JD (1993) A parsimony analysis of the Asteridae sensu lato based on rbcL sequences. Annals of the Missouri Botanical Garden 80, 700–722.
Crossref |
open url image1

Olmstead RG, Kim KJ, Jansen RK, Wagstaff SJ (2000) The phylogeny of the Asteridae sensu lato based on chloroplast ndhF gene sequences. Molecular Phylogenetics and Evolution 16, 96–112.
Crossref | GoogleScholarGoogle Scholar | PubMed | open url image1

Olmstead RG, dePamphilis CW, Wolfe AD, Young ND, Elisons WJ, Reeves PA (2001) Disintegration of the Scrophulariaceae. American Journal of Botany 88, 348–361.
PubMed |
open url image1

Oxelman B, Backlund M, Bremer B (1999) Relationships of the Buddlejaceae s. l. investigated using parsimony jackknife and branch support analysis of chloroplast ndhF and rbcL sequence data. Systematic Botany 24, 164–182.
Crossref |
open url image1

Oxelman B, Kornhall P, Olmstead RG, Bremer B (2005) Further disintegration of the Scrophulariaceae. Taxon 54, 411–425. open url image1

Oyama RK, Baum DA (2004) Phylogenetic relationships of North American Antirrhinum (Veronicaceae). American Journal of Botany 91, 918–925. open url image1

dePamphilis CW, Young ND, Wolfe AD (1997) Evolution of plastid gene rps2 in a lineage of hemiparasitic and holoparasitic plants: many losses of photosynthesis and complex patterns of rate variation. Proceedings of the National Academy of Sciences USA 94, 7367–7372.
Crossref | GoogleScholarGoogle Scholar | open url image1

Pan KY, Li ZY, Wang YZ (2002) Floral organogenesis of Titanotrichum oldhamii (Gesneriaceae). Acta Botanica Sinica 44, 895–902. open url image1

Pennell FW (1919) Scrophulariaceae of the southeastern United States. Proceedings. Academy of Natural Sciences of Philadelphia 71, 224–291. open url image1

Pennell FW (1921) Scrophulariaceae of the west gulf states. Proceedings. Academy of Natural Sciences of Philadelphia 73, 459–536. open url image1

Pennell FW (1935) The Scrophulariaceae of eastern temperate North America. Academy of Natural Sciences of Philadelphia Monographs 1, 1–650. open url image1

Rahmanzadeh R, Muller K, Fischer E, Bartels D, Borsch T (2005) The Linderniaceae and Gratiolaceae are further lineages distinct from the Scrophulariaceae (Lamiales). Plant Biology 7, 67–78.
Crossref | GoogleScholarGoogle Scholar | PubMed | open url image1

Rahn K (1996) A phylogenetic study of the Plantaginaceae. Botanical Journal of the Linnean Society 120, 145–198.
Crossref | GoogleScholarGoogle Scholar | open url image1

Reeves PA, Olmstead RG (1998) Evolution of novel morphological and reproductive traits in a clade containing Antirrhinum majus (Scrophulariaceae). American Journal of Botany 85, 1047–1056.
Crossref |
open url image1

Reeves PA, Olmstead RG (2003) Evolution of the TCP gene family in asteridae: cladistic and network approaches to understanding regulatory gene family diversification and its impact on morphological evolution. Molecular Biology and Evolution 20, 1997–2009.
Crossref | GoogleScholarGoogle Scholar | PubMed | open url image1

Reveal JL (1996) Newly required suprageneric names in Magnoliophyta. Phytologia 79, 68–76. open url image1

Rønsted N, Chase MW, Albach DC, Bello MA (2002) Phylogenetic relationships within Plantago (Plantaginaceae): evidence from nuclear ribosomal ITS and plastid trnL–F sequence data. Botanical Journal of the Linnean Society 139, 323–338.
Crossref | GoogleScholarGoogle Scholar | open url image1

Rønsted N, Bello MA, Jensen SR (2003) Aragoside and iridoid glucosides from Aragoa cundinamarcensis. Phytochemistry 64, 529–533.
Crossref | GoogleScholarGoogle Scholar | PubMed | open url image1

Rossow RA (1985) Melospermeae, nueva tribu de Scrophulariaceae. Parodiana 3, 365–396. open url image1

Rourke JP (2000) A review of generic concepts in the Stilbaceae. Bothalia 30, 9–15. open url image1

Rourke JP (1993) Thesmophora, a new genus of Stilbaceae from South Africa. Edinburgh Journal of Botany 50, 89–95. open url image1

Rouy G (1909) Conspectus des tribus et des genres de la famille des Scrophularicées. Revue Generale de Botanique , 194–207. open url image1

Schauer (1846) Phrymaceae. In ‘Prodromus systematis naturalis regni vegetabilis’. (Ed. A deCandolle) p. 520. (Victoris Masson: Paris)

Schneeweiss GM, Colwell A, Park J-M, Jang C-G, Stuessy TF (2004) Phylogeny of holoparasitic Orobanche (Orobanchaceae) inferred from nuclear ITS sequences. Molecular Phylogenetics and Evolution 30, 465–478.
Crossref | GoogleScholarGoogle Scholar | PubMed | open url image1

Scotland RW, Sweere JA, Reeves PA, Olmstead RG (1995) Higher-level systematics of Acanthaceae determined by chloroplast DNA sequences. American Journal of Botany 82, 266–275.
Crossref |
open url image1

Smith JF, Wolfram JC, Brown KD, Carroll CL, Denton DS (1997a) Tribal relationships in the Gesneriaceae: evidence from DNA sequences of the chloroplast gene ndhF. Annals of the Missouri Botanical Garden 84, 50–66.
Crossref |
open url image1

Smith JF, Brown KD, Carroll CL, Denton DS (1997b) Familial placement of Cyrtandromoea, Titanotrichum and Sanango, three problematic genera of the Lamiales. Taxon 46, 65–74.
Crossref |
open url image1

Solereder H (1909) Uber die gattung Rehmannia. Berichte der Deutschen Botanischen Gesellschaft 27, 390–404. open url image1

Soltis DE, Soltis PS, Chase MW, Mort ME, Albach DC , et al. (2000) Angiosperm phylogeny inferred from 18S rDNA, rbcL, and atpB sequences. Botanical Journal of the Linnean Society 133, 381–461.
Crossref | GoogleScholarGoogle Scholar | open url image1

Spangler RE, Olmstead RG (1999) Phylogenetic analysis of Bignoniaceae based on the cpDNA gene sequences rbcL and ndhF. Annals of the Missouri Botanical Garden 86, 33–46.
Crossref |
open url image1

Stefanovic S, Olmstead RG (2004) Testing the phylogenetic position of a parasitic plant (Cuscuta, Convolvulaceae, Asteridae): Bayesian inference and the parametric bootstrap on data drawn from three genomes. Systematic Biology 53, 384–399.
Crossref | GoogleScholarGoogle Scholar | PubMed | open url image1

Steiner KE (1996) Chromosome numbers and relationships in tribe Hemimerideae (Scrophulariaceae). Systematic Botany 21, 63–76.
Crossref |
open url image1

Steiner KE, Whitehead VB (2002) Oil secretion and the pollination of Colpias mollis (Scrophulariaceae). Plant Systematics and Evolution 235, 53–66.
Crossref | GoogleScholarGoogle Scholar | open url image1

Sutton DA (1988) ‘A revision of the tribe Antirrhineae.’ (British Museum of Natural History and Oxford University Press: London)

Takhtajan A (1997) ‘Diversity and classification of flowering plants.’ (Columbia University Press: New York)

Thieret JW (1955) The seeds of Veronica and allied genera. Lloydia 18, 37–45. open url image1

Thieret JW (1967) Supraspecific classification in the Scrophulariaceae: a review. SIDA 3, 87–106. open url image1

Van Tieghem MP (1903) Structure de l’étamine chez les Scrofulariacées. Annales des Sciences Naturelles 17, 363–371. open url image1

Vogel S (1974) Ölblumen und ölsammelnde Bienen. Tropische und Subtropische Pflanzenwelt 7, 1–267. open url image1

Wagstaff SJ, Olmstead RG (1997) Phylogeny of Labiatae and Verbenaceae inferred from rbcL sequences. Systematic Botany 22, 165–179.
Crossref |
open url image1

Wagstaff SJ, Bayly MJ, Garnock-Jones PJ, Albach DC (2002) Classification, origin, and diversification of the New Zealand hebes (Scrophulariaceae). Annals of the Missouri Botanical Garden 89, 38–63.
Crossref |
open url image1

Wallander E, Albert VA (2000) Phylogeny and classification of Oleaceae based on rps16 and trnL–F sequence data. American Journal of Botany 87, 1827–1841.
Crossref | PubMed |
open url image1

Wang C-N, Möller M, Cronk MCB (2004) Phylogenetic position of Titanotrichum oldhamii (Gesneriaceae) inferred from four different gene regions. Systematic Botany 29, 407–418.
Crossref | GoogleScholarGoogle Scholar | open url image1

Weber A (1989) Family position and conjectural affinities of Charadrophila capensis Marloth. Botanische Jahrbücher für Systematik und Pflanzengeographie 111, 87–119. open url image1

Wettstein R (1891) Scrophulariaceae. In ‘Die natürlichen pflanzenfamilien’. (Eds A Engler, K Prantl) pp. 39–107. (Wilhelm Engelmann: Leipzig)

Wolfe AD, dePamphilis CW (1998) The effect of relaxed functional constraints on the photosynthetic gene rbcL in photosynthetic and nonphotosynthetic parasitic plants. Molecular Biology and Evolution 15, 1243–1258.
PubMed |
open url image1

Wolfe AD, Elisens WJ, Watson LE, dePamphilis CW (1997) Using restriction-site variation of PCR-amplified cpDNA genes for phylogenetic analysis of tribe Cheloneae (Scrophulariaceae). American Journal of Botany 84, 555–564.
Crossref |
open url image1

Wolfe AD, Datwyler SL, Randle CP (2002) A phylogenetic and biogeographic analysis of the Cheloneae (Scrophulariaceae) based on ITS and matK sequence data. Systematic Botany 27, 138–148. open url image1

Wolfe AD, Randle CP, Liu L, Steiner KE (2005) Phylogeny and biogeography of Orobanchaceae. Folia Geobotanica 40, 115–134. open url image1

Young ND, Steiner KE, dePamphilis CW (1999) The evolution of parasitism in Scrophulariaceae / Orobanchaceae: plastid gene sequences refute an evolutionary transition series. Annals of the Missouri Botanical Garden 86, 876–893.
Crossref |
open url image1