Amsinckiinae

Adelinia Amsinckia Amsinckiopsis Andersonglossum Cryptantha Dasynotis Eremocarya Greeneocharis Harpagonella Johnstonella Oncaglossum Oreocarya Pectocarya Plagiobothrys Simpsonanthus Sonnea
Adelinia Amsinckia Amsinckiopsis Andersonglossum Cryptantha Dasynotus Eremocarya Greeneocharis Harpagonella Johnstonella Oncaglossum Oreocarya Pectocarya Plagiobothrys Simpsonanthus Sonnea

Systematics of Amsinckiinae (Boraginaceae): The Popcorn Flowers


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

      This is a website on the plant family Boraginaceae (sensu Weigend et al. 2014, Luebert et al. 2016, Hasenstab-Lehman 2017), subtribe Amsinckiinae, many members of which are commonly known as "popcorn flowers". The Amsinckiinae was referred to as subtribe Cryptanthinae (orginally Eucryptantheae Brand 1931:22) by others (Ovchinnikova 2009, Hasenstab-Lehman and Simpson 2012), but recent nomenclatural opinion is that Cryptanthinae Brand is illegitimate (given it was described originally with a "Eu-" prefix), and that the earliest available name for the clade is Amsinckiinae (originally Amsinckieae; Brand, 1931:204). Recent molecular work has demonstrated that the taxa of subtribe Amsinckiinae form a natural, monophyletic group (Schwarzer 2007,Hasenstab-Lehman and Simpson 2012) nested within tribe Cynoglosseae (Långström and Chase 2002, Weigend et al. 2010, Nazaire and Hufford 2012, Weigend et al. 2013, Cohen 2014, Chacón et al. 2016). There is a recent biogeographic study of the Boraginaceae, including the Amsinckiinae (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 the monotypic Dasynotus daubenmirei and some former New World members of Cynoglossum s.l., the latter now classified as the genera Adelinia (monotypic; Cohen 2015), Andersonglossum (Cohen 2015), and Oncaglossum (monotypic; Sutory' 2010). We suggest that the Amsinckiinae include these taxa, as accepted by Chacón et al. 2016. A recent study (Sutory' 2019) suggests that a newly named genus, Ailuroglossum (2 species), native to China, may be a close relative of Adelinia, Andersonglossum, and Oncaglossum, and may warrant placement in or near the Amsinckiinae.

      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 16 genera, 292 species, and 335 minimum ranked taxa (including varieties and subspecies; see table below). We have proposed a number of nomenclatural changes 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. See Balderas et al. 2019 (pdf; png) for a poster of nutlet diversity in the Amsinckiinae.


AMSINCKIINAE WORKING GROUP

BORAGINALES WORKING GROUP


SUMMARY OF TAXONOMIC DIVERSITY:
GENERA-16 SPECIES MIN. RANKED TAXA
Adelinia 1 1
Amsinckia 15 17
Amsinckiopsis 1 2
Andersonglossum 3 3
Cryptantha 99 111
Dasynotus 1 1
Eremocarya 2 3
Greeneocharis 2 3
Johnstonella 17 19
Harpagonella 2 2
Oncaglossum 1 1
Oreocarya 63 72
Pectocarya 12 13
Plagiobothrys 68 84
Simpsonanthus 1 1
Sonnea 2 2
     
TOTAL: 290 335



KEY TO GENERA OF THE AMSINCKIINAE (after Guilliams et al., 2020):


1. Plants perennial; leaves relatively large, generally > 5 cm long; nutlets (3.5–)5–12 mm long
    2. Nutlets not glochidiate......Dasynotus daubenmirei (monotypic)
    2' Nutlets glochidiate
        3. Corollas green to lemon yellow to brown, inflorescences somewhat bracteate; plants of Mexico......Oncaglossum pringlei (monotypic)
        3' Corollas 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
1' Plants annual, biennial, or perennial; leaves relatively small, generally < 5 cm long; nutlets < 5 mm long
    5. Fruits bur-like; sepals in fruit very unequal, upper 2 > others, partly fused, arched over 1 nutlet, with 5 to 10 stout prickles, each with hooked bristles, lower 3 sepals distinct; nutlets 2......Harpagonella
    5’ Fruits not burlike; sepals in fruit ± equal or, if unequal, upper 2 > others, distinct, not arched over 1 nutlet, without prickles but with hooked or straight trichomes; nutlets 1 to 4
        6. Mature nutlets generally 4, widely spreading......Pectocarya
        6' Mature nutlets 1-4, generally erect
            7. Nutlets without ventral groove, generally with ventral ridge distal to attachment scar
                8. Corolla limbs and tubes bright yellow or orange; cotyledons 2-lobed......Amsinckia
                8' Corolla limbs and tubes white to cream colored; cotyledons not lobed
                    9. Cymules glomerate; nutlet attachment scars above nutlet middle......Sonnea
                    9’ Cymules generally elongate (glomerate in Amsinckiopsis); nutlet attachment scars between nutlet base and nutlet middle
                        10. Plant vestiture various, but not coarsely hirsute to hispid......Plagiobothrys
                        10' Plants coarsely hirsute to hispid
                               11. Cymules more or less glomerate; corolla limbs 4–7 mm wide; nutlet abaxial surfaces rugose and papillate......Amsinckiopsis kingii (monotypic)
                               11’ Cymules elongate; corolla limbs 1–3 mm wide; nutlet abaxial surfaces tessellate......Simpsonanthus jonesii (monotypic)
            7' Nutlets with ventral groove extending to apex or sub-apically; attachment scars generally recessed or depressed
                12. Plants annual, generally wider than tall, often rounded to cushionlike; taproots red or purple when dry (often staining herbarium paper); flower bracts present
                      13. Sepals fused proximally, tubular, tube circumscissile in fruit; gynobase < nutlet...... Greeneocharis
                      13' Sepals distinct, intact in fruit; gynobase > nutlet......Eremocarya
                12' Plants annual, biennial, or perennial, generally taller than wide, rarely rounded to cushion–like; taproots usually not red or purple when dry; flower bracts absent or rarely present
                      14. Plants biennial or perennial; leaves basal or tufted; nutlets smooth, rugose, or scabrous, apex broadly rounded to obtuse, ventral groove apex well below nutlet apex......Oreocarya
                      14' Plants annual or rarely biennial or perennial; leaves generally cauline, rarely basal; nutlets smooth, papillate, tuberculate, or muricate, apex narrowly acute to acuminate,
                            ventral groove apex almost to nutlet apex
                            15. Corolla limbs bright yellow or orange; cotyledons 2-lobed......Amsinckia
                            15' Corolla limbs white; cotyledons not lobed (2-lobed in some South American Cryptantha)
                                  16. Plants annual, rarely biennial (possibly perennial in J. racemosa); flowers always chasmogamous; nutlets ovate, triangular-ovate, or lance-ovate, often heteromorphic in size
                                        and sculpturing, margin rounded or angled and then often narrowly ridged or winged, densely or sparsely tuberculate, tubercles usually whitish......Johnstonella
                                  16' Plants annual; flowers chasmogamous or cleistogamous; nutlets lanceolate, lance-ovate, or ovate, usually homomorphic, margin rounded or angled, not ridged or winged
                                        (except C. oxygona, C. cycloptera, and C. pterocarya), smooth or papillate/tuberculate, if tuberculate, tubercles generally not whitish......Cryptantha


CHROMOSOME NUMBERS OF AMSINCKIINAE

TERMINOLOGY USED IN AMSINCKIINAE DESCRIPTIONS AND KEYS

LINKS OF INTEREST


LITERATURE CITED:

Balderis, E., C. M. Guilliams, K. E. Hasenstab-Lehman, M. E. Mabry, and M. G. Simpson. 2019. Amsinckiinae Working Group: Nutlet diversity of Amsinckiinae
      in western North America. Poster (pdf; png) presented at Botany 2019. Tucson, Arizona.
Brand, A. 1931. Borraginaceae-Borraginoideae-Cryptantheae. Pp. 1-236 in A. Engler (ed.), Das Pflanzenreich. 97. Verlag von Wilhelm Engelmann, Leipzig.
      subtribe Eucryptantheae (p. 22); subtribe Amsinckiiae (p. 204)
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.
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.
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.
Cohen, J. I. 2015. Adelinia and Andersonglossum (Boraginaceae), two new genera from New World species of Cynoglossum. Systematic Botany 40: 611–619.
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.
Guilliams, C. M., K. Hasenstab-Lehman, and B. G. Baldwin. 2020. Nomenclatural changes in western North American Amsinckiinae (Boraginaceae). Novon 28:51-59.
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.
Hasenstab-Lehman, K. E. 2017. Phylogenetics of the borage family: Delimiting Boraginales and assessing closest relatives. Aliso: A Journal of Systematic
      and Evolutionary Botany 31:41-49.
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.
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.
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 .
Mabry, M. E. and M. G. Simpson. 2018. Evaluating the monophyly and biogeography of Cryptantha (Boraginaceae). Systematic Botany 43:53-76.
Nazaire, M. and L. Hufford. 2012. A broad phylogenetic analysis of Boraginaceae: Implications for the relationships of Mertensia.
     Systematic Botany 37:758–783.
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.
Ovchinnikova, S. 2009. On the position of the tribe Eritrichieae in the Boraginaceae system. Botanica Serbica 33:141-146.
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.
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.
Sutory´, K. 2010. Oncaglossum, a new genus of the Boraginaceae, tribe Cynoglosseae, from Mexico. Novon 20:463–469.
Sutory’, K. 2019. Ailuroglossum (Boraginaceae, Cynoglosseae), a new genus endemic to southern China, and a new species. Novon 27:131-136.
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.
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.


HOW TO CITE THIS WEBSITE:
Simpson, M. G. 2007 onwards. Systematics of Amsinckiinae (Boraginaceae): The popcorn flowers. Website: https://plants.sdsu.edu/amsinckiinae
or
Amsinckiinae Working Group [Current year]. Systematics of Amsinckiinae (Boraginaceae): The popcorn flowers. Website: https://plants.sdsu.edu/amsinckiinae




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