Common Name: BORAGE FAMILY
Habit: Annual, perennial herb, or shrub, often bristly or sharp-hairy. Stem: prostrate-decumbent to erect. Leaf: basal and/or cauline, simple, generally alternate, sometimes opposite, especially at base. Inflorescence: cymes, arranged singly or in groups of 2--5, generally coiled in flower, generally elongating in fruit. Flower: bisexual, generally radial; sepals 5, free or fused at least at base; corolla 5-lobed, salverform, funnel-shaped, rotate, or bell-shaped, appendages (often called "fornices") 0 or 5 at top of tube, when present often differentially pigmented, alternate stamens; stamens epipetalous; ovary superior, 4-lobed, style 1, entire or minutely 2-lobed (2-branched). Fruit: nutlets 1--4, when > 1, all similar (often called "homomorphic") or 1 or 2 dissimilar in size and/or shape from the others (often called "heteromorphic"), free (fused), smooth to roughened, prickly or bristly or not.
Genera In Family: +- 90 genera, +- 1600--1700 species: mostly temperate, especially western North America, Mediterranean; some cultivated (Borago, Echium, Myosotis, Symphytum). Toxicity: Many genera may be TOXIC from pyrrolizidine alkaloids or accumulated nitrates. Note: Sometimes still treated in broader sense of TJM2 (e.g., APG IV 2016 Bot J Linn Soc 181:1--20), but recent evidence (Luebert et al. 2016) supports segregation, for our flora, of the families Ehretiaceae, Heliotropiaceae, Hydrophyllaceae, Lennoaceae, and Namaceae.
eFlora Treatment Author: Michael G. Simpson, C. Matt Guilliams, Kristen Hasenstab-Lehman & Ronald B. Kelley
Scientific Editor: Bruce G. Baldwin, C. Matt Guilliams, Kristen Hasenstab-Lehman, David J. Keil, Ronald B. Kelley, Robert W. Patterson, Thomas J. Rosatti & Michael G. Simpson

Habit: Annual. Stem: branches generally ascending to erect, minutely strigose (hairs distally oriented) and/or soft- to rough-bristly. Leaf: generally sessile, basal and cauline; basal generally not well-developed, often withering at flowering, cauline generally alternate (proximal opposite), distal reduced; generally strigose and/or soft- or rough-bristly, largest bristles generally bulbous-based. Inflorescence: generally terminal, raceme-like or generally spike-like cymes, in groups of 1--5s(>5), generally coiled in bud, generally elongated in fruit; bracts generally 0, flowers also mostly without subtending bracts; pedicel 0--1.5(3) mm in fruit. Flower: generally unscented; calyx generally lengthening in fruit, appressed to spreading (or recurved) in fruit, persistent or not at maturity, lobes separate to base; corolla deciduous, tube generally = calyx in flower, limb 0.5--11 mm diam, generally rotate, occasionally +- funnelform, white, appendages generally present, 5; anthers included; ovary generally 4-lobed; receptacle elongate, terminating in style and stigma. Fruit: nutlets 1--4, similar or less commonly dissimilar, if dissimilar single different nutlet positioned toward inflorescence axis, nutlets symmetric (or asymmetric), flattened or not, generally gray to brown, often mottled, smooth to tubercled, and/or papillate, dull to shiny, margin rounded, sharp-edged, or a +- flat linear rim or wing; abaxially longitudinal ridge present or not; adaxially grooved above centered attachment scar, groove extending to nutlet tip, attachment scar edges abutted entire length to variably gapped, often fork- or flare-gapped at base; style from receptacle axis extending << or > tip of nutlet(s).
Etymology: (Greek: hidden flowers, from cleistogamous flowers of some South American species, including the type species) Note: Other taxa in TJM2 moved to Eremocarya, Greeneocharis, Johnstonella, Oreocarya. Homostylous (see Oreocarya). The tissue between ovary lobes, interpreted as a modified receptacle, extends to various degrees in fruit, forming what is often called the gynobase, to which the nutlets are laterally attached at maturity, leaving an attachment scar. A single style with a minute stigma arises from the gynobase. The extension of the style tip/stigma relative to the height/length of the nutlet(s) can be important in identification. Species without yellow corolla appendages are generally thought to be self-pollinating, but there is no direct evidence for this. Some species, e.g., Cryptantha ambigua, Cryptantha barbigera, Cryptantha mariposae, are thought to hybridize with co-occurring species. Observation of nutlets, hairs best at 10+ × generally critical for identification. Corolla limb diam much < at end of flowering period, especially noticeable for large-flowered taxa; measurements here attempt to reflect this developmental change, which far exceeds population differences for most taxa. 2n=(12)20, 24 (North American taxa). Some South American species are perennial, cleistogamous, and/or polyploid. One taxon, C. maritima var. pilosa, is found in both North and South America.
eFlora Treatment Author: Michael G. Simpson, Kristen E. Hasenstab-Lehman, Makenzie E. Mabry, and Ronald B. Kelley
Unabridged Reference: Johnston 1925 Contr Gray Herbarium 74:1--125; Simpson & Hasenstab 2009 Crossosoma 35:1--59; Hasenstab-Lehman & Simpson 2012 Syst Bot 37:738--757; Simpson et al. 2013 Madroño 60:24--34; Mabry et al. 2016 Phytotaxa 253:97--130; Simpson & Kelley 2017 Phytotaxa 295:227--236; Simpson et al. 2017 Taxon 66:1406--1420; Mabry & Simpson 2018 Syst Bot 43:53--76; Simpson & Rebman 2021 Phytotaxa 509:185--210; Simpson & Rebman 2022 Madroño 69: in press.