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

Systematics of Amsinckiinae (Boraginaceae): The Popcorn Flowers

[Amsinckiinae HOME]

Pectocarya DC. ex Meisn.

Pl. Vasc. Gen. 1: 279 (1840)

Gr. pectos, combed, + karua, nut," in reference to the pectinate (comb-like) nutlet margins on some
Cynoglossum L., Sp. Pl.: 134 (1753)
Gruvelia A.DC., Candolle, Prodr. 10: 119 (1846) = Pectocarya DC. ex Meisn.
Mattia Schult., Observ. Bot.: 30 (1809) = Cynoglossum L., Sp. Pl.: 134 (1753)
Rindera Pall., Reise Russ. Reich. 1: 486 (1771)
Ctenospermum Post & Kuntze,Lex. Gen. Phan.: 152 (1903)

Lehm., Index Seminum (HBG, Hamburgensis) 1837: 5, 7 (1837), nomen nudum = Pectocarya DC. ex Meisn.
Type Species/EspeciesTipo:
Pectocarya lateriflora (Lamarck) DC., designated by Meisner 1840; see Veno 1979

Distribution of Pectocarya:
LEFT: POWO. 2019. Plants of the World Online. Facilitated by the Royal Botanic Gardens, Kew. Published on the Internet. Accessed 31 Dec 2019.
RIGHT: GBIF. 2019. Global Biodiversity Information Facility. Accessed 31 Dec 2019.







11 species and 12 minimally ranked taxa
NA=North America; SA=South America; SA*=possibly non-native in SA (see Veno 1979, p.32)
Pectocarya anisocarpa Veno NA   Mexico, USA
Pectocarya anomala I.M.Johnston   SA Chile, Peru
Pectocarya dimorpha (I.M.Johnston) I.M.Johnston   SA Chile
Pectocarya heterocarpa (I.M.Johnston) I.M.Johnston NA   Mexico, USA
Pectocarya lateriflora (Lamarck) DC.   SA Argentina, Bolivia, Peru
Pectocarya linearis (Ruiz & Pavon) DC. NA SA Argentina, Chile, Mexico, USA
      var. ferocula I.M.Johnston NA SA Argentina, Chile, Mexico, USA
      var. linearis   SA Argentina, Chile
Pectocarya penicillata (Hooker & Arnott) A.DC. NA   Mexico, USA
Pectocarya peninsularis I.M.Johnston NA   Mexico, USA
Pectocarya platycarpa (Munz & I.M.Johnston) Munz & I.M.Johnston NA   Mexico, USA
Pectocarya recurvata I.M.Johnst. NA   Mexico, USA
Pectocarya setosa A.Gray NA   Mexico, USA
Pectocarya brachycera Veno, ined. [Veno 1979, p. 193]      
Pectocarya ferocula Veno, ined. [Veno 1979, p. 171]      

SYNONYMS (In Progress):
Pectocarya boliviana (I.M.Johnston) I.M.Johnston =Pectocarya lateriflora (Lamarck) DC. [Veno 1979, p. 185]
Pectocarya chilensis DC. =Pectocarya linearis (Ruiz & Pavon) DC. [Veno 1979, p. 180]
Pectocarya chilensis DC. var. californica Torrey [Veno 1979] =Pectocarya pusilla (A.DC.) A.Gray [Veno 1979, p. 72]
Pectocarya gracilis (Ruiz & Pavon) I.M.Johnston =Pectocarya linearis (Ruiz & Pavon) DC.
Pectocarya gracilis (Ruiz & Pavon) I.M.Johnston var. boliviana I.M.Johnston =Pectocarya lateriflora (Lamarck) DC. [Veno 1979, p. 185]
Pectocarya gracilis (Ruiz & Pavon) I.M.Johnston var. dimorpha I.M.Johnston =Pectocarya dimorpha (I.M.Johnston) I.M.Johnston [Veno 1979, p. 116]
Pectocarya gracilis (Ruiz & Pavon) I.M.Johnston var. genuina I.M.Johnston =Pectocarya linearis (Ruiz & Pavon) DC.
Pectocarya gracilis (Ruiz & Pavon) I.M.Johnston var. platycarpa Munz & I.M.Johnston =Pectocarya platycarpa (Munz & I.M.Johnston) Munz & I.M.Johnston [Veno 1979, p. 163]
Pectocarya lateriflora (Lamarck) DC. var. nuttallii =Lappula sp. Moench. [Veno 1979, p. 198]
Pectocarya linearis (Ruiz & Pavon) DC. var. ferocula =Pectocarya linearis (Ruiz & Pavon) DC. var. ferocula I.M.Johnston
Pectocarya linearis (Ruiz & Pavon) DC. var. penicillata M.E.Jones =Pectocarya penicillata (Hooker & Arnott) A.DC. [Veno 1979, p. 135]
Pectocarya linearis (Ruiz & Pavon) DC. var. platycara (Munz & I.M.Johnston) Cronquist =Pectocarya platycarpa (Munz & I.M.Johnston) Munz & I.M.Johnston [Veno 1979, p. 163]
Pectocarya miser A.Nelson =Pectocarya penicillata (Hooker & Arnott) A.DC. [Veno 1979, p. 136]
Pectocarya penicillata (Hooker & Arnott) A.DC. var. genuina I.M.Johnston =Pectocarya penicillata (Hooker & Arnott) A.DC. [Veno 1979, p. 136]
Pectocarya penicillata (Hooker & Arnott) A.DC. var. heterocarpa I.M.Johnston =Pectocarya heterocarpa (I.M.Johnston) I.M.Johnston [Veno 1979, p. 96]
Pectocarya pusilla (A.DC.) A.Gray =Gruvelia pusilla A.DC.
Pectocarya pusilla (A.DC.) A.Gray var. flagellaris Brand =Pectocarya pusilla (A.DC.) A.Gray [Veno 1979, p. 72]
Pectocarya setosa A.Gray var. aptera I.M.Johnston =Pectocarya setosa A.Gray [Veno 1979, p. 84]
Pectocarya setosa A.Gray var. genuina I.M.Johnston =Pectocarya setosa A.Gray
Pectocarya setosa A.Gray var. holoptera I.M.Johnston =Pectocarya setosa A.Gray [Veno 1979, p. 85]


1. Calyx radially symmetric, lobes > nutlets; nutlet body ± circular to obovate or ± rhomboid angular; lower cauline leaves opposite, fused at base (Sect. Gruvelia)
    2. Nutlets homomorphic, equally divergent, nutlet body ± rhomboid angular; calyx lobe tip with slender, uncinate hairs; stem not branched or branched near base,
        sparsely strigose; pedicel in fruit 1–2.3 mm ...... Gruvelia pusilla
2' Nutlets heteromorphic, divergent in pairs, nutlet body ± circular to obovate; calyx lobe tip with pungent, straight bristles; stem branched above, ± hispid;
        pedicel in fruit ± 0.5 mm ...... P. setosa
1' Calyx not radially symmetric, ± bilateral or irregular, lobes ≤ nutlets; nutlet body linear or oblanceolate to oblong or ovate-elliptic; lower cauline leaves generally
    alternate, not fused at base (Sect. Pectocarya)
    3. Nutlets of cauline fruits homomorphic, all nutlets similar in size and/or degree of ornamentation
s         4. Nutlets conspicuously winged, the wing margin often bearing bristles or teeth, wing ≥ teeth or bristles
            5. Nutlet wing ± lobed on both sides proximally, with uncinate hairs or bristles distally, not toothed or lacerate ...... P. penicillata
5’ Nutlet wing not appearing lobed, hairs and bristles absent, toothed the length of the nutlet, teeth stout ...... P. platycarpa (2)
        4' Nutlets not conspicuously winged, nutlet body appearing to bear bristles or teeth ± directly, wing wanting or << teeth or bristles
            6. Stems ascending to erect; fruit not flat; nutlets strongly recurved to coiled ...... P. recurvata
            6' Stems prostrate to decumbent; fruit ± flat; nutlets straight or slightly recurved at tip ...... P. linearis var. ferocula (2)
    3' Nutlets of cauline fruits heteromorphic, the lower nutlet or both lower and upper nutlets toward inflorescence axis differing from nutlets away from inflorescence
        axis in size and/or degree of ornamentation
           7. Cauline and basal fruits  ± similar in shape and margin ornamentation (fruits monomorphic)
               8. Nutlets away from inflorescence axis winged, wing margins ± undulate, not or barely toothed, not lacerate ...... P. anisocarpa
8' Nutlets away from inflorescence axis winged or not, if winged, wing margin not entire to undulate, toothed to lacerate along ± length of nutlet
                   9. Stems ascending to erect; nutlet wing broad, tooth width at base ± = length, nutlet body ± oblong ...... P. platycarpa s(2)
                   9' Stems prostrate to decumbent; nutlet wing, if present, narrow, tooth width at base generally << length, nutlet body ± linear ...... P. linearis var. ferocula (2)
          7' Cauline and basal fruits strongly dissimilar (fruits dimorphic); basal fruit nutlets ± homomorphic, and having nutlets ± without marginal ornamentation,
              cauline fruit nutlets heteromorphic and with obvious nutlet ornamentation
              10. Nutlets of cauline fruits in 2 planes; pair away from inflorescence axis incurved, nutlet wing lacerate-toothed; pair toward inflorescence axis
                     spreading to deflexed; pedicel in fruit partially adnate to lower nutlet toward inflorescence axis; corolla limb diameter 0.5-1 (1.5) mm ...... P. heterocarpa
10' Nutlets of cauline fruits all in 1 plane; pair away from inflorescence axis flat, nutlet wing generally entire with uncinate bristles; pair toward
                     inflorescence axis flat; pedicel in fruit free from nutlets; corolla limb diameter 1.5-3.5 mm ...... P. peninsularis

 Pectocarya in South America
(From Horn 2000)

1. Pflanze meistens unverzweigt, selten wenig verzweigt, Stengelblätter kreuzgegenständig, Rand der Klausen nur als kleine Wulst mit kräftigen
    Krallenhaaren ausgebildet (Gruvelia) ...... P. pusilla = Gruvelia pusilla
1' Pflanze stark verzweigt, Rand der Klausen breit, tief gezähnt, mit kräftigen Krallenhaaren (Pectocarya) ...... 2
   2. Blüten groß, im Durchmesser größer als 2,5 mm ...... 3
       3. Klausen im Umriß schmal länglich ...... P. linearis
       3' Klausen im Umriß breit eiförmig bis trullat ...... P. lateriflora
   2' Blüten sehr klein, im Durchmesser kleiner als 2,0 mm. ...... 4
       4. Pflanze an der Basis mit kleistogamen Blüten; je zwei Klausen einer chasmogamen Blüte gleichgestaltet (dimorph), Rand der Klausen zur Hälfte gezähnt,
           Zähne kurz kegelförmig, wie aufgeblasen ...... P. dimorpha
       4' Pflanze nur mit chasmogamen Blüten, alle vier Klausen einer Blüte gleichgestaltet, Rand der Klausen fast bis zur Basis gezähnt, Zähne nicht wie aufgeblasen ...... 5
          5. Klausen dicht besetzt mit langen Hakenhaaren, Klausenkörper daher kaum zu erkennen, Zähne lang, schmal kegelförmig ...... P. anomala
          5' Klausen spärlich besetzt mit kurzen Hakenhaaren, Klausenkörper daher gut zu erkennen, Zähne lang, breit dreieckig ......
P. boliviana = P. lateriflora

Translation to English:
1. Plant mostly unbranched, rarely slightly branched, calyx lobes opposite, margin of fruit only as a small bulge having strong uncinate
    hairs formed (Gruvelia) ...... P. pusilla = Gruvelia pusilla
1' Plant with many branches, margin of the fruit broad, deeply toothed, with strong uncinate hairs (Pectocarya) ...... 2
   2. Flowers large, larger than 2.5 mm in diameter ...... 3
       3. Fruit narrowly elongate in outline ...... P. linearis
       3' Fruit broadly ovate to trullate in outline ...... P. lateriflora
   2' Flowers very small, less than 2.0 mm in diameter ...... 4
       4. Plant at the base with cleistogamic flowers; two fruits each of a chasmogamous flower shaped alike (dimorphic), margin of the fruits half-serrated,
              teeth short conical, as if inflated . ...... P. dimorpha
       4' Plant only with chasmogamous flowers, all four fruitsof a flower shaped alike, margin of the fruits toothed almost to the base, teeth not as if inflated ...... 5
          5. Fruits densely covered with long hooked hairs, therefore the fruit body hardly recognizable, teeth long, narrowly conical ...... P. anomala
          5' Fruits sparsely occupied with short hooked hairs, therefore the fruit body easily recognizable, teeth long, broad triangular ......
P. boliviana = P. lateriflora

Pectocarya in South America
(From Johnston 1927. Note: P. gracilis is now treated as a synonym of P. linearis and P. boliviana as a synonym of P. lateriflora)

1. Nutlets equally divergent, much compressed, obviously angular and rhomboidal in outline (sect. Gruvelia) ...... P. pusilla = Gruvelia pusilla
1' Nutlets divergent in pairs, thickish, not obviously angular nor rhomboidal in outline (sect. Pectocarya)
    2. Back and sides of nutlets armed with coarse subterete subulate appendages tipped with uncinate hairs ...... P. anomala
    2' Back of nutlets smooth or at most papillate, margin with a lacerately or pectinately cut wing-margin bearing uncinate hairs
        3. Fruit dimorphic, that produced at base of plant with reflexed subpersistent nutlets obviously different from the divaricate nutlets produced by
            the normal fruit of the middle and upper stems ...... P. dimorpha
        3' Fruit of one sort, that produced at the base of the plant similar to that on the upper stem
            4. Nutlets linear, 2.5-3 mm long, sides parallel ...... P. gracilis = P. linearis
            4' Nutlets obovate or obovate-oblong
                5. Nutlet body 2-2.5 mm long, with medial dorsal ridge ...... P. lateriflora
                5' Nutlet body 1.5-2 mm long, lacking medial dorsal ridge ...... P. boliviana = P. lateriflora

Pectocarya in North & South America
(From Veno 1979.)

1. Calyx appendaged with 5 to 10 subterete, subulate horns armed with subretrorse uncinate prickles; upper two calyx lobes elongate, indurate, and galeate, partially enclosing a nutlet; mature nutlets two. (Section Harpagonella) ...... P. palmeri = Harpagonella palmeri
1. Calyx not as above; mature nutlets four ...... 2.
    2. Calyx radially symmetrical; lobes equally divergent, sub equal in length, greatly exceeding the nutlets; basal and basal-cauline leaves opposite, decussate, and connate at the base; upper cauline leaves alternate; nutlet margins a narrow rim or spreading wing, entire or subentire or rarely cleft. (Section Gruvelia) ...... 3.
        3. Calyx lobes apically uncinate bristled; nutlets equally divergent, broadly obtrullate to subrhomboidal, homomorphic (rarely heteromorphic); margins narrow rims (rarely absent), entire; stems slender, strigose ...... P. pusilla = Gruvelia pusilla
        3. Calyx lobes setose-hispid; nutlets geminate, broadly obovatc, heteromorphic (rarely all four nutlets unmargined); margins broad wings, entire to subentire (rarely cleft); stems stiff, setose-hispidulous ...... P. setosa
    2. Calyx bilaterally symmetrical; upper lobes two, exceeding the lower three lobes, subequal or ~ the lower lobes irregularly reduced and displaced by fusion of the pedicel to the lower adaxial nutlet, only slightly exceeding or equal to or shorter than the nutlets; only basalmost leaves opposite, remaining cauline leaves alternate; nutlets wing margined or appendaged with subulate horns; margins involute, erect, ascending, or spreading; margin form subentire, dentate, or lacerate. (Section Pectocarya) ...... 4.
        4. Nutlets wing margined ...... 5.
            5. Fruits dimorphic, dissimilar basal and cauline fruits; nutlets distinctly heteromorphic, opposing pairs of nutlets dissimilar ...... 6.
                6. Fruiting pedicel partially fused to the lower adaxial nutlet; calyx obliquely distorted, lower adaxial and medial lobes greatly reduced, and alternately displaced along the pedicel; nutlet pairs usually somewhat biplanar, abaxial nutlets ascendingly curved, adaxial nutlets deflexed and somewhat descendingly curved; nutlet outline oblong to oblanceolate ...... P. heterocarpa
                6. Fruiting pedicels free; calyx +/- regular, three lower lobes subequal; nutlets +/- planar to slightly deflexed; nutlet outline elliptic to orbicular obovate or oblong to broadly obovate ...... 7.
                    7. Nutlet outline elliptic to orbicular obovate; margins of lower or, most commonly, both adaxial nutlets distinctly reduced to absent, to 0.4 mm wide, entire to subentire; margins of abaxial nutlets well developed, ascending-spreading, cleft (to parted) dentate, 0.25 - 0.75 mm wide; one to four nutlets of basal fruits unmargined, often yellowish; North America ...... P. peninsularis
                    7. Nutlet outline oblong to broadly obovate; margin of lower or, occasionally, both adaxial nutlets reduced, to 1.0 mm wide, parted to divided dentate with teeth often intermittent; margins of abaxial nutlets well developed, involute to erect to ascending-spreading, irregularly cleft to divided into broad dentate segments, 0.35 - 1.2 mm wide; all nutlets of the basal fruits narrowly margined, parted to divided dentate with teeth occasionally intermittent; South America ...... P. dimorpha
            5. Fruits monomorphic, basal and cauline fruits similar; nutlets homomorphic, opposing pairs of nutlets similar, or heteromorphic ...... 8.
                8. Nutlet margins bilobed, bidentate, subentire, or shallowly dentate; nutlet outline oblong, oblanceolate, or obscurely panduriform ...... 9.
                    9. Nutlets distinctly heteromorphic; margins of lower or, most commonly, both adaxial nutlets reduced, entire (-subentire); margins of abaxial nutlets well developed, ascending to erect, subentire (lobed to cleft) or rarely bidentate; all nutlet margins of basalmost fruits slightly reduced ...... P. anisocarpa
                    9. Nutlets homomorphic; margins of all nutlets well developed or adaxial margins only slightly reduced, involute to erect (-ascending), commonly bidentate into broad segments or occasionally subentire; nutlet margins of basalmost fruits well developed ...... P. penicillata
                8. Nutlet margins pectinate-dentate or lacerate-dentate; nutlet outline linear, linear-oblanceolate to oblanceolate or obovate to broadly obovate ...... 10.
                    10. Nutlet margin pectinate-dentate, erect to ascending; nutlet outline linear to oblanceolate; nutlets homomorphic to heteromorphic, nutlet surface smooth. ...... 11.
                          11. Nutlets strongly recurved to scorpiod (-coiled), narrow, linear, ~ homomorphic, length/width= 6/1 (- 5/1); margin erect; margin teeth narrow, two or more times longer than wide ...... P. recurvata
                          11. Nutlets only slightly to moderately recurved, linearoblanceolate to oblanceolate, homomorphic to heteromorphic, length/width = 5/1 to 4/1 (-3/1); margin erect to ascending; margin teeth narrow and less than two time longer than wide, or teeth wide and two or more times longer than wide ...... 12.
                                12. Nutlets ~ planar or with apices somewhat recurved, linear-oblanceolate, homomorphic to slightly heteromorphic, length/width = 5/1 (-4/1); margin erect to strictly ascending, (cleft-) divided dentate; margin teeth less than two times longer than wide; North America, coastal California and Baja California Norte, Mexico; South America, Argentina (~rare Chile) ...... P. ferocula = P. linearis var. ferocula
                                12. Nutlets moderately recurved, oblanceolate, heteromorphic with the lower or both adaxial nutlet margins and/or nutlet size reduced, length/width = 4/1 (-3/1); margin sub- involute to ascending; margin teeth wide, two or more times longer than wide ...... 13.
                                      13. Plants erect to ascending (- decumbent); nutlet margins strictly to widely ascending, broadly and regularly cleft to parted dentate; North America. southeastern California, Arizona, and Sonora, Mexico ...... P. platycarpa
                                      13. Plants prostrate to decumbent; nutlet margins subinvolute to erect to ascending. often mediallaterally constricted, irregularly parted to divided dentate; South America, Chile, rare Argentina ...... P. linearis
                    10. Nutlet margin lacerate-dentate, spreading; nutlet outline broad obovate to subdeltoid; nutlets heteromorphic, lower adaxial nutlet reduced ~n size, more densely pubescent and deeply rugose than the three remaining nutlets; nutlet surface dorsal-medially ridged or short papillate, smooth to rugose, pubescent to glabrous ...... P. lateriflora.
        4. Nutlets appendaged with subterete, subulate horns ...... 14.
            14. Nutlet horns lateral and dorsal-medial; nutlet surface deeply rugose to convoluted ...... P. anomala
            14. Nutlet horns lateral only; nutlets dorsal-medially naked or intermittently short papillate; nutlet surface smooth to obscurely or moderately rugose ...... P. brachycera, ined.



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      with a revised dichotomous key and photographic atlas for the genus in the region. Aliso 31: 1-13.
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      Fakultät für Biologie der Ludwig – Maximilians – Universität München. 
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     University of California, Los Angeles.
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      y Uruguay) 2 Dicotyledoneae: Acanthaceae-Fabaceae (Abarema-Schizolobium), ed.:

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