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 2013, Weigend et al. 2013, Chacón et al. 2016). Note recent biogeographic study of the family (Chacón et al. 2017).
      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.
      Recent molecular phylogenetic studies (Weigend et al. 2013, Otero et al. 2014, Cohen 2014, Chacón et al. 2016) 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 segregate Oncaglossum pringlei, and the unique (monotypic) Dasynotus daubenmirei. We suggest that the Amsinckiinae include these taxa, as accepted by Chacón et al. 2016.
      We also cite some more distant outgroups to the Amsinckiinae: Hackelia, Microula, and Myosotis.

Amsinckiinae Working Group

Boraginales Working Group

Participants of the 2016 Boraginales Working Group, Bonn, Germany

Key to the Genera of the Amsinckiinae (excluding Adelinia, Andersonglossum, Dasynotus, and Oncaglossum)
(modified from Kelley and Wilken 1993):


1.
Mature nutlets spreading widely
    2. Sepals in fr 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
    2' Sepals in fr ± 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
1' Mature nutlets ± erect
    3. Nutlets with ventral groove, extending to apex or sub-apically; attachment scar gen recessed or depressed (Cryptantha s.l.)
        4. Plants annual, gen wider than tall, often rounded to cushion-like; taproot red or purple when dry (often staining herbarium paper); floral bracts present
            5. Calyx basally fused, tubular, tube circumscissile in fruit; gynobase < nutlet ...... Greeneocharis
            5' Calyx of distinct sepals, intact in fruit; gynobase > nutlet ...... Eremocarya
        4' 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
            6. Plants biennial or perennial; vegetative leaves basal or tufted; nutlets smooth, rugose, or scabrous, apically broadly rounded to obtuse,
               ventral groove apex well below nutlet apex ...... Oreocarya
            6' 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
               7. 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
               7' Plants annual; flowers chasmogamous or cleistogamous; nutlets lanceolate, lance-ovate, or ovate, usually homomorphic, margin rounded or angled, not winged
                  (except C. oxygona, C. pterocarya with generally toothed to undulate wings), smooth or papillate/tuberculate, if tuberculate, tubercles gen. not whitish ...... Cryptantha s.s.
    3' Nutlets without ventral groove, generally with ventral ridge distal to attachment scar
         8. Corolla limb and tube bright yellow or orange; cotyledons 2-lobed ...... Amsinckia
         8' Corolla limb and tubes white to cream colored; cotyledons not lobed ...... Plagiobothrys



List of Technical Terms Used in Keys


Literature Cited

Brand, A. 1925. Drei neue Gattungen dar Cryptantheae. Repertorium Specierum Novarum Regni Vegetabilis 21:249-254. [pdf]
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]
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]
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. and K. E. Hasenstab. 2009. Cryptantha of Southern California. Crossosoma 35:1-59. [pdf]
Weigend, M., M. Gottschling, F. Selvi, and H. H. Hilger. 2010. Fossil and extant western hemisphere Boragineae, and the polyphyly of
     "Trigonotideae" Riedl (Boraginaceae: Boraginoideae). Systematic Botany 35:409-419. [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]