Boraginaceae
Subtribe Amsinckiinae Systematics

Michael G. Simpson
San Diego State University
San Diego, California, U.S.A.

      This is a web site on the plant family Boraginaceae (sensu Weigend et al. 2014, Luebert et al. 2016), subtribe Amsinckiinae Brand, A. in H.G.A. Engler, Nat. Pflanzenr. 97: 204. 27 Feb 1931. The Amsinckiinae was referred to as subtribe Cryptanthinae by others (Brand 1931 :22, Ovchinnikova 2009, Hasenstab-Lehman and Simpson 2012), but recent nomenclatural opinion is that Cryptanthinae Brand is illegitimate, and that the earliest available name for the clade is Amsinckiinae Brand. Recent molecular work has demonstrated that the taxa of subtribe Amsinckiinae form a natural, monophyletic group (Schwarzer 2007, Hasenstab 2009, Hasenstab-Lehman and Simpson 2012) nested within tribe Cynoglosseae (Långström and Chase 2002, Weigend et al. 2010, Nazaire and Hufford 2012, Cohen 2014, Weigend et al. 2013, Chacón et al. 2016). Note a recent biogeographic study of the family (Chacón et al. 2017).
     Recent molecular phylogenetic studies (Weigend et al. 2013, Otero et al. 2014, Cohen 2014, Chacón et al. 2016, Simpson et al. 2017) have demonstrated the need for an expansion of our Amsinckiinae clade to include some former New World members of Cynoglossum s.l., these now classified as the genera Adelinia and Andersonglossum (Cohen 2015), the Mexican monotypic Oncaglossum pringlei, and the monotypic Dasynotus daubenmirei. We suggest that the Amsinckiinae include these taxa, as accepted by Chacón et al. 2016.
      The Amsinckiinae Working Group and collaborators are working on the phylogenetic relationships of the taxa of this subtribe. The subtribe currently includes a total of 13 genera. We have proposed a number of nomenclatural changes (see Amsinckia, Cryptantha s.l., Cryptantha s.s., Eremocarya, Greeneocharis, Harpagonella, Johnstonella, Oreocarya, Pectocarya, and Plagiobothrys) and anticipate more in the future; see Simpson et al. 2017. Our biogeographic analyses (Guilliams et al., 2017) of the Amsinckiinae support the occurence of 18 dispersal events from North America to South America.
      We also cite a more distant outgroup to the Amsinckiinae: Microula.

Amsinckiinae Working Group

Boraginales Working Group

Participants of the 2016 Boraginales Working Group, Bonn, Germany

Key to the Genera of the Amsinckiinae
(modified from Kelley et al. 2012 and Cohen 2015):


1.
Plants perennial; leaves relatively large, generally >>5 cm long; nutlets (3.5)5-12 mm long
   2. Nutlets glochidiate
       3. Corolla green to lemon yellow to brown, inflorescence somewhat bracteate; plants of Mexicot ...... Oncaglossum pringlei (monotypic)
       3' Corolla white to blue to red to purple, inflorescences ebracteate; plants of the U.S.A and Canada
          4. Cauline leaves petiolate, pedicels not recurved in fruit, stems glabrous ...... Adelinia grandis (monotypic)
          4' Cauline leaves sessile, pedicels recurved in fruit, stems pubescent ...... Andersonglossum
   2' Nutlets not glochidiate ...... Dasynotus daubenmirei (monotypic)
1' Plants annual, bienniel, or perennial; leaves relatively small, generally <<5 cm long; nutlets rarely glochidiate, <5 mm long
   5. Mature nutlets widely spreading
       6. Sepals in fruit very unequal, upper 2 >> others, partly fused, arched over 1 nutlet, ± bur-like, with 5-10 stout spines, each with hooked bristles, lower 3 sepals distinct; nutlets 2 ...... Harpagonella
       6' Sepals in fruit ± equal or, if unequal, upper 2 > others, sepals without spines but with hooked or straight bristles, distinct, not arched over 1 nutlet; nutlets gen 4 ...... Pectocarya
   5' Mature nutlets generally erect
      7. Nutlets with ventral groove, extending to apex or sub-apically; attachment scar gen recessed or depressed
           8. Plants annual, gen wider than tall, often rounded to cushion-like; taproot red or purple when dry (often staining herbarium paper); floral bracts present
              9. Calyx basally fused, tubular, tube circumscissile in fruit; gynobase < nutlet ...... Greeneocharis
              9' Calyx of distinct sepals, intact in fruit; gynobase > nutlet ...... Eremocarya
           8' Plants annual, biennial, or perennial, gen taller than wide, rarely rounded to cushion–like; taproot usually not red or purple when dry; flower bracts absent or rarely present
              10. Plants biennial or perennial; vegetative leaves basal or tufted; nutlets smooth, rugose, or scabrous, apically broadly rounded to obtuse,
                   ventral groove apex well belownutlet apex ...... Oreocarya
              10' Plants annual or rarely biennial or perennial; vegetative leaves gen. cauline, rarely basal; nutlets smooth, papillate, tuberculate, or muricate, apically narrowly acute to acuminate,
                   ventral groove apex almost to nutlet apex
                   11. Corolla limb bright yellow or orange; cotyledons 2-lobed ...... Amsinckia (A. vernicosa)
                   11' Corolla limb white; cotyledons not lobed
                        12. Plants annual, rarely biennial (perennial in J. racemosa); flowers always chasmogamous; nutlets ovate or triangular-ovate, often heteromorphic in size and sculpturing,
                             margin angled (rarely rounded), often narrowly winged, densely or sparsely tuberculate, tubercles usually whitish ...... Johnstonella
                        12' Plants annual; flowers chasmogamous or cleistogamous; nutlets lanceolate, lance-ovate, or ovate, usually homomorphic, margin rounded or angled, not winged
                             (except C. oxygona, C. cycloptera, and pterocarya), smooth or papillate/tuberculate, if tuberculate, tubercles gen. not whitish ...... Cryptantha
      7' Nutlets without ventral groove, generally with ventral ridge distal to attachment scar
         13. Corolla limb and tube bright yellow or orange; cotyledons 2-lobed ...... Amsinckia
         13' Corolla limb and tubes white to cream colored; cotyledons not lobed ...... Plagiobothrys



List of Technical Terms Used in Keys


Literature Cited

Brand, A. 1931. Borraginaceae-Borraginoideae-Cryptantheae. In A. Engler (ed.), Das Pflanzenreich. Verlag von Wilhelm Engelmann, Leipzig. [pdf]
Chacón, J., F. Luebert, H. Hilger, S. Ovcinnikova, F. Selvi, L. Cecchi, C. M. Guilliams, K. Hasenstab-Lehman, K. K. Sutorý, M. G. Simpson, and M. Weigend. 2016.
      The borage family (Boraginaceae s.str.): A revised infrafamilial classification based on new phylogenetic evidence, with emphasis on the placement of some
      enigmatic genera. Taxon 3: 523-546. [pdf]
Chacón, J., F. Luebert, and M. Weigend. 2017. Biogeographic events are not correlated with diaspore dispersal modes in Boraginaceae. Frontiers in Ecology
     and Evolution 5 (Article 26):1-14. [pdf]
Cohen, J. I. 2014. A phylogenetic analysis of morphological and molecular characters of Boraginaceae: evolutionary relationships, taxonomy, and patterns of
     character evolution. Cladistics 30: 139-169. [pdf]
Cohen, J. I. 2015. Adelinia and Andersonglossum (Boraginaceae), two new genera from New World species of Cynoglossum. Systematic Botany 40: 611–619. [pdf]
Guilliams, M. C., K. Hasenstab-Lehman, M. Mabry, and M. G. Simpson. 2017. Memoirs of a frequent flier: Phylogenomics reveals 18 long-distance dispersals
     between North America and South America in the popcorn flowers (Amsinckiinae, Boraginaceae). American Journal of Botany 104:1717-1728. [pdf]
Hasenstab, K. 2009. Phylogenetic systematics of the genus Cryptantha (Boraginaceae). Master of Science Thesis, San Diego State University,
     San Diego, California. [pdf]
Hasenstab-Lehman, K. E. and M. G. Simpson. 2012. Cat’s eyes and popcorn flowers: Phylogenetic systematics of the genus Cryptantha s.l.
     (Boraginaceae). Systematic Botany 37:738–757. [pdf]
Kelley, R. B., M. G. Simpson, and K. E. Hasenstab. 2012. Cryptantha. Pp. 455-468 in B. G. Baldwin, D. H. Goldman, D. J. Keil, R. Patterson,
   T. J. Rosatti, and D. H. Wilken (eds.), The Jepson Manual: Vascular Plants of California. Second Edition. University of California Press, Berkeley,
   Los Angeles, London. [link]
Långström, E. and M. W. Chase. 2002. Tribes of Boraginoideae (Boraginaceae) and placement of Antiphytum, Echiochilon, Ogastemma and
     Sericostoma: A phylogenetic analysis based on atpB plastid DNA sequence data. Plant Systematics and Evolution 234:137-153. [pdf]
Luebert, F., L. Cecchi, M. W. Frohlich, M. Gottschling, C. M. Guilliams, K. E. Hasenstab-Lehman, H. H. Hilger, J. S. Miller, M. Mittelbach, M. Nazaire, M. Nepi,
      D. Nocentini, D. Ober, R. G. Olmstead, F. Selvi, M. G. Simpson, K. Sutorý, B. Valdés, G. K. Walden, and M. Weigend. 2016. Familial classification of the
      Boraginales. Taxon 65: 502–522 .[pdf]
Mabry, M. E. and M. G. Simpson. 2018. Evaluating the monophyly and biogeography of Cryptantha (Boraginaceae). Systematic Botany 43:53-76. [pdf]
Nazaire, M. and L. Hufford. 2012. A broad phylogenetic analysis of Boraginaceae: Implications for the relationships of Mertensia.
     Systematic Botany 37:758–783. [pdf]
Otero, A., P. Jiménez-Mejías, V. Valcárcel, and P. Vargas. 2014. Molecular phylogenetics and morphology support two new genera (Memoremea and Nihon)
     of Boraginaceae s.s. Phytotaxa 173:241-277. [pdf]
Ovchinnikova, S. 2009. On the position of the tribe Eritrichieae in the Boraginaceae system. Botanica Serbica 33:141-146. [pdf]
Schwarzer, C. 2007. Systematische Untersuchungen an den peruanischen Vertretern der Gattungen Pectocarya D.C. ex Meisn., Amsinckia
     
Lehm., Plagiobothrys Fisch. & C.A. Mey. und Cryptantha Lehm. ex G. Don (Boraginaceae). Dissertation: Biology Institut für Biologie
     Systematische Botanik und Pflanzengeographie, Freie Universität Berlin. [pdf]
Simpson, M. G., C. M. Guilliams, K. E. Hasenstab-Lehman, M. E. Mabry, and L. Ripma. 2017. Phylogeny of the popcorn flowers: Use of genome skimming to
    evaluate monophyly and interrelationships in subtribe Amsinckiinae (Boraginaceae). Taxon 66:1406–1420. [pdf]
Weigend, M., F. Luebert, F. Selvi, G. Brokamp, and H. H. Hilger. 2013. Multiple origins for Hound’s tongues (Cynoglossum L.) and Navel seeds
      (Omphalodes Mill.) – The phylogeny of the borage family (Boraginaceae s.str.). Molecular Phylogenetics and Evolution 68: 604–618. [pdf]
Weigend, M., F. Luebert, M. Gottschling, T. L. P. Couvreur, H. H. Hilger, and J. S. Millerd. 2014. From capsules to nutlets—phylogenetic relationships
     in the Boraginales. Cladistics 30:508-518. [pdf]