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Family guide for fruits and seeds

J.H. Kirkbride, Jr., C.R. Gunn, and M.J. Dallwitz

Boraginaceae Juss., nom. cons.

Synonyms: Anchusaceae Vest; Buglossaceae Hoffmanns. & Link, nom. illeg.; Cordiaceae R. Br. ex Dumort., nom. cons.; Ehretiaceae Mart., nom. cons.; Heliotropiaceae Schrad., nom. cons.; Onosmaceae Martinov; Sebestenaceae Vent., nom. illeg.; Wellstediaceae Novák

Common name: Borage Family.

Number of genera 129. Number of species 2500.

Angiosperm. Magnoliopsida.

Disseminule a dehisced fruit, or an intact or entire fruit, or a partial fruit (a segment of an entire fruit), or a seed.

Fruits

Pistil(s) compound; 1; 1-pistillate; with carpels nearly separate to base, or carpels united. Fruit pericarpium; simple, or schizocarp; drupe (Tournefortia, Heliotropium), or capsule (not Spjut); microbasarium (Spjut Fig. 38F); loculicidal capsule (Wellstedioideae, Wellstedia but not Spjut); capsule not inflated; capsule without operculum; without persistent central column; with styles(s); at base or nearly basal (for drupe), or at apex (capsule); not within accessory organ(s); 1-seeded to 10 to less than 25-seeded; 1–4-seeded (-8–10 for Trigonotis (Goldberg)); 4 cm long, or 5 cm long; 1–4-carpellate; with carpels united, or separate; with carpels remaining united at maturity, or separating at maturity; with carpels radiating at maturity, or not radiating at maturity; with carpels separating at style; without sterile carpels; not sulcate; in transection terete; apex not beaked; wall leathery; indehiscent, or dehiscent (Wellstedioideae). Dehiscent unit seed(s). Dehiscent regularly; and shedding seeds; without replum. Epicarp black, or brown (all shades), or white, or gray; dull, or shiny; durable; glabrous (without hairs), or not glabrous (with hairs); hairs not glandular; with armature, or without armature; with bristles (plain- or hooked-tipped); without armature glochidiate; smooth, or not smooth; warted, or roughened; without wing(s), or with wing(s); without apical respiratory hole. Mesocarp present, or absent. Endocarp present; not separating from exocarp; thin, or bony, or crustaceous; not splitting into 1-seeded pyrenes; smooth, or not smooth; with sculptures; without wing, or with wing; without operculum; without secretory cavities; without mechanism for seedling escape; without grooves; without longitudinal ridges. Funiculus short; short without seed bearing hooks (retinacula); not persisting in fruit after seed shed.

Seeds

Aril present, or absent; an arillike structure (rarely). Arillike structure falling with seed an elaisome. Seed larger than minute; in transection terete (including sub); not bowl shaped; not nutlike; without winglike beak; without caudate appendage(s); at maturity with food reserves, or without food reserves, or without apparent food reserves; with endosperm; without canavanine. Sarcotesta absent. Testa present; without fleshy or leathery layer over hard layer; surface smooth (assumed); without crease or line separating cotyledons from hypocotyl-radicle; without notch along margin where cotyledons from hypocotyl-radicle tip approach each other; without glands; without bristles; glabrous, or pubescent; with tuft of hairs at one end (coma) (Wellstedia); without glandular pubescence; without wings; without collar; without operculum; colored; monochrome; brown (all shades); thin; not becoming mucilaginous when wetted; surrounding embryo, or surrounding food reserve. Endosperm development nuclear, or cellular; moderate, or scant, or trace; fleshy, or fleshy-soft; with oils (in more archaic genera); without fatty acid containing cyclopropene; without apical lobes; without chlorophyll; without isodiametric faceted surface; without odor.

Embryo differentiated from food reserve; well developed; 1 per seed; nearly filling testa (trace or scanty food reserve), or partially filling testa (with food reserve); 0.4–1 times the length of food reserve; at one end of seed not extending into a depression or cup; foliate, or linear (linear in Ehretia); with spatulate cotyledons, or investing cotyledons (partially); straight, or C-shaped (slightly); parallel to seed length; without coleorhiza; without simmondsin; without stomata; not green; with 2 or more cotyledons. Cotyledons 2 (in Amsinckia so deeply bifid as to appear 4); well developed; as wide as hypocotyl-radicle, or somewhat to significantly wider than hypocotyl-radicle; 1–2.7 times wider than hypocotyl-radicle; partially concealing hypocotyl-radicle, or not concealing hypocotyl-radicle (assumed); foliaceous, or not foliaceous; thick; once-folded (Cordia); smooth, or ruminate; with apices entire, or bifid (Amsinckia); with margins separate; equal in size; not punctate dotted. Hypocotyl-radicle moderately developed, or well developed; coiled; not thickened.

Distribution

Cosmopolitan (esp. western North America & Mediterranean region). New World, Old World. North America, Middle America, South America, Europe, Africa, Asia Major, Asia Minor, southeastern Asia, Australia, Oceania.

Notes

Fruits: Cordioideae: hard or fleshy drupe with locular stone; Ehretioideae: drupe with 2, 2-seeded or 1-seeded stones, or separating into 4 segments; Heliotropoideae" 2 rarely 1 2- seeded, separating nutlets, or 4, 1-seeded separating nutlets; Boraginoideae: (1-)4 distinct, smooth or often variously ornamented nutlets individually attached to gynobase, separate or separating carpels having a common style, and at maturity each carpel disarticulated into discrete seed-containing half-carpels (mericarps); Wellstedioideae: (Wellstedia, 2 species) 1–2-seeded, locular capsule, seed comose, endosperm nil.

Weed information

1 or more USA state noxious weeds.

USA states and territories with listed noxious weeds: Alaska (AK), California (CA), Colorado (CO), Connecticut (CT), Massachusetts (MA), Montana (MT), Nevada (NV), New Hampshire (NH), Oregon (OR), South Dakota (SD), Washington (WA), Wyoming (WY).

USA state and territory noxious weeds: -- Anchusa arvensis (L.) M. Bieb.: USA state noxious weed: WA●. -- Anchusa officinalis L.: USA state noxious weed: OR●, WA●. -- Buglossoides arvensis (L.) I. M. Johnst.: USA state noxious weed: WA°. -- Cynoglossum officinale L.: USA state noxious weed: CO●°, MT●°, NV●, OR●, WA●, WY●°. -- Echium plantagineum L.: USA state noxious weed: OR●.-- Echium vulgare L.: USA state noxious weed: WA●. -- Lappula echinata Gilib. (=Lappula squarrosa (Retz.) Dumort.): USA state noxious weed: AK°. -- Myosotis scorpioides L.: USA state noxious weed: CTª, MAª, NHª.-- Symphytum asperum Lepech.: USA state noxious weed: CA●. -- Symbols: ªaquatic weed; ●terrestrial weed; °weed in seed. -- Last updated September 2008.

Listed seeds

ASOA listed seeds, ISTA listed seeds.

ASOA listed seeds: -- Allocarya Greene spp. = Plagiobothrys Fisch. & C. A. Mey. spp. -- Lappula echinata Gilib. = Lappula squarrosa (Retz.) Dumort. -- Amsinckia intermedia Fisch. & C. A. Mey. -- Amsinckia Lehm. spp. -- Amsinckia tessellata A. Gray -- Anchusa azurea Mill. -- Anchusa capensis Thunb. -- Anchusa L. spp. -- Asperugo procumbens L. -- Borago officinalis L. -- Brunnera macrophylla (Adams) I. M. Johnst. -- Buglossoides arvensis (L.) I. M. Johnst. -- Cryptantha Lehm. ex G. Don spp. -- Cynoglossum amabile Stapf & J. R. Drumm. -- Cynoglossum officinale L. -- Cynoglossum L. spp. -- Cynoglossum virginianum L. -- Echium vulgare L. -- Heliotropium curassavicum L. -- Heliotropium europaeum L. -- Heliotropium L. spp. -- Lappula occidentalis var. cupulata (A. Gray) L. C. Higgins -- Lappula occidentalis var. occidentalis (S. Watson) Greene -- Lappula redowskii var. occidentalis (S. Watson) Rydb. = Lappula occidentalis var. occidentalis (S. Watson) Greene -- Lappula redowskii var. texana (Scheele) Brand = Lappula occidentalis var. cupulata (A. Gray) L. C. Higgins -- Lappula Wolf spp. -- Lappula squarrosa (Retz.) Dumort. -- Lappula texana (Scheele) Britton = Lappula occidentalis var. cupulata (A. Gray) L. C. Higgins -- Lithospermum arvense L. = Buglossoides arvensis (L.) I. M. Johnst. -- Lithospermum incisum Lehm. -- Lithospermum ruderale Douglas ex Lehm. -- Lithospermum L. spp. -- Mertensia Roth spp. -- Myosotis alpestris F. W. Schmidt -- Myosotis alpestris hort. = Myosotis sylvatica Hoffm. -- Myosotis arvensis (L.) Hill -- Myosotis scorpioides L. -- Myosotis sylvatica Hoffm. -- Nama hispida A. Gray -- Nemophila maculata Benth. ex Lindl. -- Nemophila menziesii Hook. & Arn. -- Nemophila Nutt. spp. -- Pectocarya penicillata (Hook. & Arn.) DC. -- Phacelia californica Cham. -- Phacelia campanularia A. Gray -- Phacelia minor (Harv.) Thell. -- Phacelia Juss. spp. -- Phacelia tanacetifolia Benth. -- Plagiobothrys figuratus (Piper) I. M. Johnst. ex M. Peck -- Plagiobothrys hispidulus (Greene) I. M. Johnst. = Plagiobothrys scouleri var. hispidulus (Greene) Dorn -- Plagiobothrys scouleri var. hispidulus (Greene) Dorn -- Plagiobothrys Fisch. & C. A. Mey. spp. -- Symphytum asperum Lepech. -- Last updated September 2008.

ISTA listed seeds: -- Alkanna tinctoria Tausch -- Alkanna tuberculata (Forssk.) Meikle, non Greuter = Alkanna tinctoria Tausch -- Anchusa angustissima K. Koch included in Anchusa leptophylla Roem. & Schult. -- Anchusa azurea Mill.f -- Anchusa capensis Thunb.f -- Anchusa incana Ledeb. included in Anchusa leptophylla Roem. & Schult. -- Anchusa italica Retz. = Anchusa azurea Mill. -- Anchusa leptophylla Roem. & Schult. -- Anchusa officinalis L.w -- Anchusa myosotidiflora Lehm. = Brunnera macrophylla (Adams) I. M. Johnst. -- Borago officinalis L.a -- Brunnera macrophylla (Adams) I. M. Johnst.f -- Buglossoides arvensis (L.) I. M. Johnst.w -- Coldenia procumbens L.w -- Cordia alliodora (Ruiz & Pav.) Oken -- Cordia goeldiana Huber -- Cynoglossum amabile Stapf & J. R. Drumm.f -- Cynoglossum officinale L.w -- Echium candicans L. f.f -- Echium fastuosum J. Jacq., non Salisb. = Echium candicans L. f.f -- Echium italicum L.w -- Echium plantagineum L.f -- Echium vulgare L.w -- Heliotropium amplexicaule Vahlw -- Heliotropium arborescens L.f -- Heliotropium europaeum L.w -- Heliotropium indicum L.w -- Heliotropium ovalifolium Forssk.w -- Lappula squarrosa (Retz.) Dumort.w -- Lithospermum arvense L. = Buglossoides arvensis (L.) I. M. Johnst.-- Myosotis alpestris F. W. Schmidt -- Myosotis alpestris hort., non F. W. Schmidt = Myosotis sylvatica Ehrh. ex Hoffm. -- Myosotis arvensis (L.) Hillw -- Myosotis oblongata Link = Myosotis sylvatica Ehrh. ex Hoffm. -- Myosotis palustris Hill = Myosotis scorpioides L. -- Myosotis scorpioides L.f -- Myosotis sylvatica Hoffm.f-- Trichodesma indicum (L.) Sm.w -- Symbols: aagricultural and vegetable seeds (Table 2A Part 1); ttree and shrub species (Table 2A Part 2); fflower, spice, herb, and medicinal seeds (Table 2A Part 3); wweed seeds. -- Last updated September 2008.

Accepted genera

Actinocarya Benth. -- Adelocaryum Brand -- Afrotysonia Rauschert -- Alkanna Tausch, nom. cons. -- Amblynotus (A. DC.) I. M. Johnst. -- Amphibologyne Brand -- Amsinckia Lehm., nom. cons. -- Anchusa L. -- Ancistrocarya Maxim. -- Anoplocaryum Ledeb. -- Antiotrema Hand.-Mazz. -- Antiphytum DC. ex Meisn. -- Arnebia Forssk. -- Asperugo L. -- Auxemma Miers -- Borago L. -- Bothriospermum Bunge -- Bourreria P. Browne, nom. cons. -- Brachybotrys Maxim. ex Oliv. -- Brunnera Steven -- Buglossoides Moench -- Caccinia Savi -- Carmona Cav. -- Cerinthe L. -- Chionocharis I. M. Johnst. -- Choriantha Riedl -- Coldenia L. -- Cordia L., nom. cons. prop. -- Cortesia Cav. -- Craniospermum Lehm. -- Cryptantha Lehm. ex G. Don -- Cynoglossopsis Brand -- Cynoglossum L. -- Cynoglottis (Gusul.) Vural & Kit Tan -- Cystostemon Balf. f. -- Dasynotus I. M. Johnst. -- Decalepidanthus Riedl -- Echiochilon Desf. -- Echiostachys Levyns -- Echium L. -- Ehretia P. Browne -- Elizaldia Willk. -- Embadium J. M. Black -- Eritrichium Schrad. ex Gaudin -- Gastrocotyle Bunge -- Gyrocaryum Valdés -- Hackelia Opiz -- Halacsya Dörfl. -- Halgania Gaudich. -- Heliocarya Bunge -- Heliotropium L. -- Heterocaryum A. DC. -- Huynhia Greuter -- Ivanjohnstonia Kazmi -- Ixorhea Fenzl -- Lacaitaea Brand -- Lappula Wolf -- Lasiarrhenum I. M. Johnst. -- Lasiocaryum I. M. Johnst. -- Lepechiniella Popov -- Lepidocordia Ducke -- Lindelofia Lehm. -- Lithodora Griseb. -- Lithospermum L. -- Lobostemon Lehm. -- Macromeria D. Don -- Maharanga A. DC. -- Mairetis I. M. Johnst. -- Mattiastrum (Boiss.) Brand -- Menais Loefl. -- Mertensia Roth, nom. cons. -- Metaeritrichium W. T. Wang -- Microcaryum I. M. Johnst. -- Microula Benth. -- Mimophytum Greenm. -- Moltkia Lehm. -- Moltkiopsis I. M. Johnst. -- Moritzia DC. ex Meisn. -- Myosotidium Hook. -- Myosotis L. -- Neatostema I. M. Johnst. -- Nesocaryum I. M. Johnst. -- Nogalia Verdc. -- Nomosa I. M. Johnst. -- Nonea Medik. -- Ogastemma Brummitt -- Omphalodes Mill. -- Omphalolappula Brand -- Omphalotrigonotis W. T. Wang -- Onosma L. -- Onosmodium Michx. -- Oxyosmyles Speg. -- Paracaryum (A. DC.) Boiss. -- Pardoglossum E. Barbier & Mathez -- Patagonula L. -- Pectocarya DC. ex Meisn. -- Pentaglottis Tausch -- Perittostema I. M. Johnst. -- Plagiobothrys Fisch. & C. A. Mey. -- Pontechium Böhle & Hilger -- Pseudomertensia Riedl -- Psilolaemus I. M. Johnst. -- Pteleocarpa Oliv. -- Pulmonaria L. -- Rindera Pall. -- Rochefortia Sw. -- Rochelia Rchb., nom. cons. -- Rotula Lour. -- Saccellium Bonpl. -- Scapicephalus Ovcz. & Czukav. -- Selkirkia Hemsl. -- Sericostoma Stocks -- Setulocarya R. R. Mill & D. G. Long -- Sinojohnstonia Hu -- Solenanthus Ledeb. -- Stenosolenium Turcz. -- Stephanocaryum Popov -- Suchtelenia Kar. ex Meisn. -- Symphytum L. -- Thaumatocaryon Baill. -- Thyrocarpus Hance -- Tianschaniella B. Fedtsch. ex Popov -- Tiquilia Pers. -- Tournefortia L., nom. cons. prop. -- Trachelanthus Kunze -- Trachystemon D. Don -- Trichodesma R. Br., nom. cons. -- Trigonocaryum Trautv. -- Trigonotis Steven -- Ulugbekia Zakirov -- Valentiniella Speg. -- Wellstedia Balf. f.

References specific to this family

Cronquist page 917.

General references

Corner, E.J.H. 1976. The seeds of Dicots, esp. vol. 2. Cambridge University Press, New York, Cronquist, A. 1981. An integrated system of classification of flowering plants, 1,262 p. Columbia University Press, New York, Gaertner, J. 1788–1805. De fructibus et seminibus plantarum. The Author, Stuttgart, Gunn, C.R. and C.A. Ritchie. 1988. Identification of disseminules listed in the Federal Noxious Weed Act. Techn. Bull. U.S.D.A. 1719:1–313, Gunn, C.R., J.H. Wiersema, C.A. Ritchie, and J.H. Kirkbride, Jr. 1992 and amendments. Families and genera of Spermatophytes recognized by the Agricultural Research Service. Techn. Bull. U.S.D.A. 1796:1–500, Mabberley, D.J. 1987. The plant-book, 706 p. Cambridge University Press, Cambridge, Martin, A.C. 1946. The comparative internal morphology of seeds. Amer. Midl. Naturalist 36:513–660, Spjut, R.W. 1994. A systematic treatment of fruit types. Mem. New York Bot. Gard. 70:1–182, Wood, C.E., Jr. 1974. A student's atlas of flowering plants: Some dicotyledons of eastern North America, 120 pp. Harper and Row, New York.

Illustrations

Acceptable fruit and no seed illustrations. Disseminule illustration(s): fruit, or fruit, or embryo. Fruit illustration(s): Cronquist. Fruit illustration(s): Wood, Jr., Gunn & Ritchie. Embryo illustration(s): Gunn & Ritchie, Martin.

• Seed. 2 of 16. Amsinckia tessellata A. Gray: fruits. • Seed. 3 of 16. Borago officinalis L.: fruits. • Fruit. 4 of 16. Cordia myxa L.: fruit. • Seed. 6 of 16. Cynoglossum officinale L.: fruits. • Fruit. 7 of 16. Echium candicans L.f.: fruiting head. • Seed. 8 of 16. Echium candicans L.f.: fruits. • Fruit and seed. 9 of 16. Lithospermum ruderale Douglas ex Lehm.: seed and fruit with calyx. • Embryo. 10 of 16. Amsinckia calycina (Moris) Chater: embryo. • Embryo. 11 of 16. Bothriospermum tenellum (Hornem.) Fisch. & C.A.Mey.: embryo. • Embryo. 12 of 16. Cordia curassavica Jacq.) Roem. & Schult.: embryo. • Embryo. 13 of 16. Cynoglossum coeruleum Hochst. ex A. DC.: embryo. • Embryo. 14 of 16. Echium plantagineum L.: embryo. • Embryo. 15 of 16. Heliotropium ellipticum Ledeb.: embryo. • Embryo. 16 of 16. Trichodesma zeylanicum (Burm. f.) R. Br.: embryo.


We advise against extracting comparative information from the descriptions. This is much more easily achieved using the DELTA data files or the interactive key, which allows access to the character list, illustrations, full and partial descriptions, diagnostic descriptions, differences and similarities between taxa, lists of taxa exhibiting or lacking specified attributes, and distributions of character states within any set of taxa. See also Guidelines for using data taken from Web publications.


Cite this publication as: ‘J.H. Kirkbride, Jr., C.R. Gunn, and M.J. Dallwitz. 2000 onwards. Family guide for fruits and seeds: descriptions, illustrations, identification, and information retrieval. Version: 12th April 2021. delta-intkey.com’.


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