Two new combinations in Euploca Nutt. (Heliotropiaceae, Boraginales) and a conspectus of the species of the Guiana Shield area

Christian Feuillet

doi:10.3897/phytokeys.61.6260

Research Article

Two new combinations in Euploca Nutt. (Heliotropiaceae, Boraginales) and a conspectus of the species of the Guiana Shield area

Christian Feuillet^‡

‡ Smithsonian Institution, Washington, United States of America

Corresponding author: Christian Feuillet ( feuillec@si.edu )

Academic editor: Sandy Knapp

This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.

Citation: Feuillet C (2016) Two new combinations in Euploca Nutt. (Heliotropiaceae, Boraginales) and a conspectus of the species of the Guiana Shield area. PhytoKeys 61: 101-124. https://doi.org/10.3897/phytokeys.61.6260

Abstract

Heliotropium foliatum and Tournefortia humilis are transferred to Euploca Nutt. respectively as Euploca foliata comb. n. and Euploca humilis comb. n. A collection from Guyana has been recently identified as Euploca humistrata, a species previously considered a Brazilian endemic. A collection from French Guiana documents for the first time the species in that country. A key to the species of the Guiana Shield area is given. The species of Euploca from the Guiana Shield are listed with synonymy and a brief description: E. filiformis, E. humilis, E. humistrata, E. lagoensis, E. polyphylla, E. procumbens.

Keywords

Boraginales , Brazil, Euploca , Guiana Shield, Guianas, Heliotropiaceae , Venezuela

Introduction

The genus Euploca Nuttall (1836: 189) was created for the new E. convolvulacea Nutt. (1836: 190). Between 1848 and 1951, four species and two infra specific names were published that are currently considered to be synonyms of E. convolvulacea. Euploca convolvulacea was transferred to Heliotropium L. (Linnaeus 1753, 1: 130) by Gray (1857: 403). Recently Hilger and Diane (2003) reevaluated the classification of the Heliotropiaceae Schrad. (Schrader 1819: 192) on the base of molecular data of nuclear ITS1 and plastidal trnLUAA intron sequences. One of their conclusions is that Heliotropium sect. Orthostachys (R.Br.) G.Don (Brown 1810: 493; Don 1838: 361), Schleidenia Endl. (Endlicher 1839: 646), and Hilgeria Förther (1998: 132) constitute a separate genus whose earlier available name is Euploca. Their analysis indicates that Ixorhea Fenzl (1886: 287) is sister to the rest of the family consisting of two clades: one composed of Euploca and genus Myriopus Small (1933: 1131) resurrected to accommodate the species of Tournefortia sect. Cyphocyema I.M.Johnst. (Johnston 1930: 72), the other one of Heliotropium and Tournefortia L. (1753, 1: 140). Morphologically Euploca and Myriopus differ from Heliotropium and Tournefortia by the shape and structure of the fruits and their curved embryo. Euploca species are herbs or subshrubs and have usually dry fruits, in contrast Myriopus species are shrubs or passive climbers and have 4-lobed fleshy fruits. Hilger and Diane (2003: p. 47–49) made 20 new combinations in Euploca. Melo and Semir (2006; 2009) described three new species and made 12 new combinations. More new combinations were made: one by Luebert (Luebert et al. 2011), four by Degen and Melo (2012), and two by Feuillet (2012). There are probably more than 45 species of Heliotropium and Schleidenia awaiting a transfer to Euploca. Nevertheless, the nomenclature of the species of Euploca in the Guiana Shield region should be stable after the new combination E. humilis given below.

Methods

I studied the literature and type specimens. In the citation of types, “photo” refers to a print deposited in herbaria and “scan” refers to a picture posted online, available directly through herbarium sites, or sites like JSTOR or Europeana.

For the purpose of this paper, the Guiana Shield area is defined as the Venezuelan Guayana, the Guianas (Guyana, Surinam and French Guiana), and the part of Brazil north of the Amazon River and east of the Rio Negro.

Nomenclature and new combinations

Euploca Nutt., 1836

Euploca Nutt., Trans. Amer. Phil. Soc., ser. 2, 5: 189. 1836.

Heliotropium [unranked] Orthostachys R.Br., Prodr.: 493. 1810.

Type species. Heliotropium foliatum R.Br. (lectotype designated by Johnston 1928, pg. 46) [= Euploca foliata (R.Br.) Feuillet, PhytoKeys 61: 103. 2016.].

Preslaea Mart., Nov. Gen. Sp. 2: 75, t. 164. 1827; non PresliaOpiz 1824 (LAMIACEAE).

Type species. Preslaea paradoxa Mart. [= Euploca paradoxa (Mart.) J.I.M.Melo & Semir].

Orthostachys (R.Br.) Spach, Hist. Nat. Vég. 9: 32. 1838. [“1840” see Stafleu and Cowan (1985: TL2 vol. 5: 767)]

Type species. Based on Heliotropium [unranked] Orthostachys R.Br.

Heliotropium sect. Orthostachys (R.Br.) G.Don, Gen. Syst. 4: 361. 1838.

Type species. Based on Heliotropium [unranked] Orthostachys R.Br.

Schleidenia Endl., Gen. Pl. 646. 1839.

Type species. Based on Preslaea Mart.

Heliotropium subgen. Orthostachys (R.Br.) Rchb., Deut. Bot. Herb.-Buch: 112. 1841.

Type species. Based on Heliotropium [unranked] Orthostachys R.Br.

Heliotropium sect. Euploca (Nutt.) A.Gray, Proc. Amer. Acad. Arts 10: 49. 1874.

Type species. Based on Euploca Nutt.

Heliotropium sect. Orthostachys subsect. Axillaria I.M.Johnst., Contr. Gray Herb. 81: 47. 1928.

Type species. Heliotropium paradoxum (Mart.) Gürke, in Engl. and Prantl, Nat. Pflanzenfam. 4(3a): 97. 1893; non Vatke 1875. (lectotype designated by Förther 1998, pg. 135) [= Euploca paradoxa (Mart.) J.I.M.Melo & Semir].

Heliotropium sect. Orthostachys subsect. Bracteata I.M.Johnst., Contr. Gray Herb. 81: 47. 1928.

Type species. Heliotropium bracteatum R.Br., Prodr.: 493. 1810.

Heliotropium sect. Orthostachys subsect. Ebracteata I.M.Johnst., Contr. Gray Herb. 81: 47. 1928.

Type species. Heliotropium procumbens Mill., Gard. dict.. ed. 8: 10. 1768; non Kunth 1818. [= Euploca procumbens (Mill.) Diane & Hilger].

Hilgeria Förther, Sendtnera 5: 132. 1998.

Type species. Hilgeria hypogaea (Urb. & Ekman 1929: 105) Förther [= Euploca hypogaea (Urb. & Ekman) Diane & Hilger].

Type species

Euploca convolvulacea Nutt.

Description

Herbs or subshrubs. Leaves alternate, simple, entire. Trichomes on a pedestal of distinctly enlarged foliar epidermis cells. Inflorescences of single, scorpioid cymes, or flowers solitary. Flowers perfect; calyx 5–toothed or 5–lobed, persistent; corolla white, small, 5–merous, lobe’s margin involute; stamens 5, included; nectar disc at the base of the ovary; ovary functionally 4–locular, locules 1-seeded, style terminal, undivided, or absent, the stigma cryptically or clearly apically 2- or 4-fid. Fruit a schizocarp, lobed or unlobed, dry, 2–4 bony 1-seeded nutlets, embryos curved.

Distribution

North America to Argentina, Africa, Arabian Peninsula to Indochina, Australia.

Euploca foliata (R.Br.) Feuillet, comb. nov.

urn:lsid:ipni.org:names:77153393-1

Heliotropium foliatum R.Br., Prodr.: 493. 1810.

Type. Australia. Northern Territory: Carpentaria mainland opposite Grote Eyland, 4 Jan 1803, R. Brown 2932 (lectotype, designated by Craven 1996, pg. 630: BM n.v. [scan!]; isotypes: K!, MEL n.v., NSW n.v.).

Type

Based on Heliotropium foliatum R.Br.

Notes

When they resurrected Euploca, Hilger and Diane (2003: 42) said that “summarizing all urges a formal taxonomic recombination of EUPLOCA, including all species of Heliotropium section Orthostachys, Schleidenia, and Hilgeria into the genus Euploca, the oldest available generic name in this group.” Heliotropium foliatum is the type of H. sect. Orthostachys (Johnston 1928: 46). Although it is recognized as a good species (example: Förther 1998: 195), Heliotropium foliatum was not transferred to Euploca until now. Robert Brown did not cite a type collection, only mentioning “v. v.” (=seen alive).

Euploca humilis (L.) Feuillet, comb. nov.

urn:lsid:ipni.org:names:77153394-1

Tournefortia humilis L., Sp. Pl.: 141. 1753.

Type. “Tournefortia foliis lanceolatis” pl. by Plumier, in Burman, Pl. Amer., 224, t. 227, f. 2. 1760 (lectotype designated by Miller in Cafferty and Jarvis 2004, pg. 804). Epitype: FRANCE. Martinique: Case Pilote, Feb 1868, L. Hahn 416 (designated by Miller in Cafferty and Jarvis 2004, pg. 804: BM n.v.).

Type

Based on Tournefortia humilis L.

Notes

When Melo and Semir (2009: 289) transferred Heliotropium ternatum Vahl (1794, 3: 21) to Euploca, they did not cite synonyms and they overlooked the case of Tournefortia humilis L. There had been a doubt about the exact identity of the Linnean species (Johnston 1949: 136; Förther 1998: 190), but the typification of Tournefortia humilis L. by Miller (in Cafferty and Jarvis 2004: 804) placed clearly that species in the synonymy of Heliotropium ternatum Vahl (non H. humile Lam.). The epithet of T. humilis could not be transferred to Heliotropium, but there is no previous Euploca humilis and the priority rule dictates the adoption of the new combination (McNeill et al. 2012: Art. 11.4).

Because the name T. humilis L. or/and H. humile (L.) R.Br. ex Roem. & Schult. (1819, 4: 37; non Lam., 1791: 393) have been used in synonymy (Johnston 1928: 69; 1949: 135; and especially the recent revision of Heliotropium Förther 1998: 190, 234; and the typification Miller, in Cafferty and Jarvis 2004: 804); because it is a Linnean name; and because the authors of the new combination Euploca ternata (Melo and Semir 2009: 289) should have been aware of that name, there are little chances to make a successful proposal to conserve Euploca ternata.

Conspectus of Euploca for the Guiana Shield region

The last treatment of Heliotropium (including Euploca) for the Guianas (Johnston 1935) included four of the six species treated below: H. filiforme=E. filiformis, H. lagoense=E. lagoensis, H. procumbens=E. procumbens, H. ternatum=E. humilis. Since then E. humistratahas been identified from a seasonally flooded savanna in Guyana, the distribution area of E. lagoensis includes French Guiana. Besides, E. polyphylla not present in the Guianas occurs in the Venezuelan Guayana (State of Bolívar). The species present in the Guiana Shield region are usually found on sand near water or in seasonally dry localities.

Key to the species of Euploca in the Guiana Shield

1	Flowers solitary; pedicels 1–3 mm long; corolla white and yellow	2
–	Inflorescences 1–18 cm long; flowers sessile or sub-sessile, or pedicels obvious; corolla white or yellow	3
2	Stem glabrous or puberulent; leaf blades glabrous or puberulent; corolla white, yellow at throat, lobes elliptic	*E. lagoensis*
–	Stem villose; leaf blades with long, stiff trichomes; corolla tube yellow, lobes white, ovate	*E. humistrata*
3	Petiole 4–15 mm long; raceme mostly geminate or ternate, ebracteate; stigma sessile, base as wide as the apex of the ovary, forming together a blunt cone	*E. procumbens*
–	Petiole 0.3–2 mm long; raceme single or paired, bracteate; stigma morphologically clearly separate from the ovary, looking together like a mushroom or like a head with a hat or with a clear style	4
4	Leaf blade Inflorescences always single; bract filiform to narrowly elliptic; calyx length greater than or less than half the corolla tube; corolla white; style 0.3–0.5 mm long	5
–	Inflorescences usually single and paired on the same plant; bracts ovate; calyx nearly as long as the corolla tube; corolla yellow; style 0.5–1 mm long	*E. polyphylla*
5	Stem relatively coarse and stiff; leaves drying rather light colored; inflorescences up to 20 cm long, with 1–2 cm long peduncle; calyx shorter than half the corolla tube; corolla 4–6 mm long, salverform; anthers apically joined; stigma on a short style	*E. humilis*
–	Stem wiry, very slender; leaves usually drying dark colored; inflorescences epedunculate, up to 10 cm long; calyx longer than the corolla tube; corolla 2–2.5 mm long, tube urceolate; anthers not joined at apex; stigma sessile	*E. filiformis*

Euploca filiformis (Lehm.) J.I.M.Melo & Semir, 2009

Euploca filiformis (Lehm.) J.I.M.Melo & Semir, Kew Bull. 64: 289. 2009.

Heliotropium filiforme Lehm., Gött. Gel. Anz. 3(152): 1515. 1817.

Type. Venezuela. Near Orinoco, Bonpland and Humboldt 1202 [an error, most likely for 1203] (lectotype, designated by Förther 1998, pg. 195: B-W 3246/1 n.v. [scan!]; isotypes MEL 233288 n.v., P! [2 sheets; scans!], P-Bonpl! [scan!]). [same type collection as next, but different holotype]

Heliotropium filiforme Kunth, Nov. Gen. Sp. (quarto ed.) 3: 86, t. 204. 1818, nom. illeg. non H. filiforme Lehm. 1817.

Type. Venezuela. Near Orinoco, Bonpland and Humboldt 1203 (holotype P-Bonpl! [scan!]; isotypes B-Willd 3246/1 n.v. [scan!], MEL 233288 n.v., P! [2 sheets; scans!]. [same type collection as previous, but different holotype]

Heliotropium tenue Roem. & Schult., Syst. Veg. 4: 737. 1819.

Type. Brazil. Pará, Sieber for Hoffmannsegg s.n. (holotype B-WILLD-3248!; isotypes HAL n.v., MEL 233189 n.v., P!).

Preslaea stenostachya A.St.-Hil., Voy. Distr. Diam. 2: 434. 1833.

Type. Brazil. “Ad ripas fluminis Parahyba”, St.-Hilaire B228 (lectotype, designated by Förther 1998, pg. 235: P! [scan!]; isotypes K!, M n.v., MPU!, P! [scan!]).

Heliotropium littorale Mart. ex Colla, Herb. Pedem. 4: 226. 1835.

Type. Brazil. “In arenosis maritimis”, Martius s.n. (holotype TO n.v.; isotype BM n.v.).

Note. placed in the synonymy fide Förther 1998 who saw the isotype at BM.

Heliotropium pusillum Colla, Herb. Pedem. 4: 227. 1835.

Type. Brazil. Martius s.n. (holotype TO n.v.).

Heliotropium helophilum Mart., Flora 21(2), Beibl. 4: 85. 1838.

Type. Brazil. “Propre Cujaba”, 1837, Herb. Fl. Bras. 165, n°267 (lectotype, designated by Förther 1998, pg. 199: BR n.v. [scan!]; isotypes E n.v. [scan!], FI-W!, G-DC!, GH n.v., L! [2 sheets], LE n.v. [2 sheets], M n.v., P! [2 sheets; scans!], W n.v.).

Schleidenia stenostachya (A.St.-Hil.) DC., Prodr. 9: 558. 1845.

Type. Based on Preslaea stenostachya A.St.-Hil.

Schleidenia filiformis (Lehm.) Fresen., Fl. Bras. 8(1): 40. 1857.

Type. Based on Heliotropium filiforme Lehm.

Heliotropium stenostachyum (A.St.-Hil.) Gürke, in Engler and Prantl, Nat. Pflanzenfam. 4(3a): 97. 1893.

Type. Based on Preslaea stenostachya A.St.-Hil.

Type

Based on Heliotropium filiforme Lehm.

Description

Annual herb or subshrub, up to 0.25 m tall. Stems slender, erect or decumbent, up to 0.4 m long, sparingly strigose. Leaves: petiole 0.1–0.2 cm long, slender; lamina linear or elliptic or lanceolate or oblanceolate, 1–2.5 × 0.15–0.35 cm, apex acute, base acute, margin entire, both surfaces sericeous to strigose. Inflorescences epedunculate, slender scorpioid cymes, up to 15 cm long, bearing scattered bracts, filiform or subulate, 0.1–0.2 cm long, glabrescent. Flower sub–sessile: calyx longer than the corolla tube, lobes, 1.5–2 mm long at anthesis, glabrescent; corolla white, tubular, outside strigose, inside glabrescent, tube narrowed in the middle, lobes ovate with broadly open sinuses, about 1 mm long; stamens sessile, anther with an apical appendage, weakly coherent; ovary subglobose or globose, strigose; style lacking or nearly so; stigma sessile or sub–sessile, fertile base much wider than the apex of the ovary. Fruit on 0.5–1 mm long pedicel, depressed–globose; nutlets almost 1.5 mm long.

Distribution

Venezuela (Amazonas, Bolívar), Guyana, Surinam, French Guiana, Brazil (Amapá, Amazonas, Pará, Roraima); also in Belize, Mexico, southeastern USA and from Trinidad to eastern Bolivia and Paraguay.

Selected specimens studied

Guyana: Cuyuni River, Matope Falls, 4 June 1952, Forest Department 6940 (Fanshawe 3376) (K, NY); Upper Takutu-Upper Essequibo, Dadanawa, 24 Oct 1957, Cook 22 (K, NY). Surinam: near Coppename River, Central Suriname Nature Reserve, 4°25'N, 56°31'W, 50–75 m, 22 Feb 2004, Clarke 11037 (US); near Ulemari River, 13 km upstream from confluence with Litani River, 3°13'N, 54°15'W, 150 m, 2 Apr 1998, Hammel 21275 (MO, US). French Guiana: Itany River, Saut upstream from Touinké, 26 Nov 1977, Cremers 5119 (CAY); Oyapock River, Sauts Fourmi, 18 Nov 1984, Granville 6916 (CAY, P).

Euploca humilis (L.) Feuillet, comb. nov.

urn:lsid:ipni.org:names:77153394-1

Tournefortia humilis L., Sp. Pl. 1: 141. 1753.

Type. “Tournefortia foliis lanceolatis” in Plumier in Burman, Pl. Amer., 224, t. 227, f. 2, 1760 [http://biodiversitylibrary.org/page/779658] (lectotype designated by Miller in Cafferty and Jarvis 2004: 804). Epitype: France. Martinique: Case Pilote, Feb 1868, L. Hahn 416 (designated, by Miller in Cafferty and Jarvis 2004, 804: BM n.v.)).

Heliotropium ternatum Vahl, Symb. Bot. 3:21. 1794.

Type. [Jamaica], “in India occidentali” (holotype C n.v. [scan!; microfiche 37: I, 3; photo GH!]).

Heliotropium hirtum Lehm., Neue Schriften Naturf. Ges. Halle 3(2): 10. 1817.

Type. Venezuela. Cumana, Humboldt and Bonpland s.n. (lectotype, designated by Förther 1998, pg. 199: B-WILLD 3247!; isotypes MEL 233292 n.v., P! [scan!], P-Bonpl! [scan!]) [same type collection as H. hispidum Kunth (see below), but a different holotype].

Heliotropium demissum Roem. & Schult., Syst. Veg. ed 15bis 4: 37. 1819.

Type. nom. illeg. renaming of Tournefortia humilis L.

Heliotropium humile (L.) R.Br. ex Roem. & Schult., Syst. Veg. ed 15bis 4: 37. 1819; non Lam. 1791.

Type. nom. illeg. in syn. of previous.

Heliotropium hispidum Kunth, Nov. Gen. Sp. (quarto ed.) 3: 87. 1818; illeg. not Lehm. 1817.

Type. Venezuela. near Cumana, Sep, Humboldt and Bonpland s.n. (holotype P-Bonpl! [scan!]; isotypes B-WILLD 3247!, MEL 233292 n.v., P! [scan!]) [same type collection as H. hirtum Lehm. (see above), but a different holotype].

Pioctonon ternatum (Vahl) Raf., Sylva Tellur.: 88. 1838.

Type. Based on Heliotropium ternatum Vahl

Pioctonon antillanum Raf., Sylva Tellur. 88. 1838.

Type. nom. illeg. renaming of Tournefortia humilis L.

Tournefortia incana (G.Mey.) G.Don, Gen. Hist. 4: 368. 1838; non Lam. 1791.

Type. Based on Messerschmidia incana G.Mey.; nom. inval.

Tournefortia meyeri DC., Prodr. 9: 530. 1845.

Type. Renaming of Messerschmidia incana G. Mey.; nom. inval.

Heliotropium fruticosum var. ternatum (Vahl) DC., Prodr. 9: 542. 1845.

Type. Based on Heliotropium ternatum Vahl

Heliotropium oaxacanum DC., Prodr. 9: 543. 1845.

Type. Mexico. Oaxaca, Aug 1834, Andrieux 205 (holotype G-DC !; isotypes FI-W n.v. [fragment GH n.v.], P!, W n.v.).

Heliophytum passerinoides Klotzsch, in Rich. Schomburgk, Reis. Br.-Guiana 3: 1152. 1849 [“1848”]; nom. nud.

Type. Based on Guyana. Rupununi: May 1842, Schomburgk 573 (B†).

Schleidenia hispida (Kunth) Fresen., Fl. Bras. 8(1): 37. 1857.

Type. Based on Heliotropium hispidum Kunth

Schleidenia fumana Fresen., in Mart.: Fl. Bras. 8, 1: 40. 1857.

Syntype. Brazil. Minas Gerais, Vauthier s.n. (G n.v.). Type : Brazil. “ad Formigas et Ribeirao Catinga”, Pohl 565 (= 3081 bzw. distr. nr. 1588) (lectotype, designated by Förther 1998, pg. 235: W n.v., photo MSB-55474; isotypes B† [photo F-17322, GH!, NY!, US!], F-874797 n.v. [scan!], GH, K!, M n.v., [as s.n.] W n.v.).

Heliotropium fumana (Fresen.) Gürke, in Engler and Prantl: Nat. Pflanzenf. 4, Abt. 3a: 97. 1893.

Type. Based on Schleidenia fumana Fresen.

Heliotropium mexicanum Greenm., Proc. Amer. Acad. Arts 33(25): 484. 1898; illegitimate: not Sessé & Moç. 1888.

Syntypes: Mexico. Dugès s.n.; Pelmer 31, 98; L.C. Smith 209, 391; Alvarez 750; Pringle 6726. Type. Mexico. Iron Mtn, near Durando, June 1897, Palmer 141 (lectotype, designated by Frohlich 1981, pg. 100: GH n.v.; isotypes BM n.v., F n.v., M n.v., MICH n.v. [scan!], MO!, NY!, S n.v., US!).

Heliotropium strictissimum sensu N.E. Br. 1901; not A. DC. 1845.

Heliotropium ternatum var. fumana (Fresen.) I.M.Johnst., J. Arnold Arb. 16:62. 1935.

Type. Based on Schleidenia fumana Fresen.

Heliotropium greenmanii Wiggins, Contr. Dudley Herb. 4:22. 1950.

Type. Based on Heliotropium mexicanum Greenm.

Euploca ternata (Vahl) J.I.M.Melo & Semir, Kew Bull. 64: 289. 2009.

Type. Based on Heliotropium ternatum Vahl

Type

Based on Tournefortia humilis L.

Description

Suffrutescent herbs up to 30 cm tall; stems stiff, erect or decumbent, 10–50 cm long, with abundant, loosely appressed trichomes. Leaves: petiole subnul–2 mm long, thin; lamina lanceolate to linear, revolute, 0.5–3 × 0.1–0.8 cm, apex acute, base acute, margin +/- revolute, adaxially sericeous, abaxially tomentose, trichomes base very large and looking calcified. Inflorescences terminal or axillary, up to 20 cm long, stiff scorpioid cymes, single, peduncle 1–2 cm long, with scattered linear or narrowly elliptic bracts, 2–3 mm long, sericeous. Flower subsessile: pedicel 0.5–1 mm long, calyx 2–3 mm long, deeply lobed, lobes unequal, shorter than 1/2 the length of the corolla tube, becoming twice as large at maturity, sericeous in and out; corolla white, strigose outside, tube narrow, 2–2.7 mm long, throat yellow, lobes spreading, 1.3–2 mm long, ovate, sinuses rounded and plicate; stamens subsessile, anthers ovate with short, obtuse, hairy, apical long appendages which are apically joined; ovary globose, 0.3–0.4 mm diam., glabrous; style short but obvious, about 0.5 mm long; stigma about as large as the ovary, split at apex. Fruit depressed globose, 4–lobed, 1.5–1.8 mm diam., strigose or hirsute; nutlets 1–1.5 mm long.

Distribution

Venezuela (Bolívar) and Guyana; also Central America, Mexico, and the West Indies Southward to southern Brazil.

Selected specimens studied

Guyana: Potaro-Siparuni, Iwokrama, Kurupukari Falls, 4°40'N, 58°40'W, 100 m, 9 Dec 1994, Mutchnick 595 (BRG, CAY, L, MO, NY, US); Upper Takutu-Upper Essequibo, near Sand Creek, Baboon Hill, 3°00'N, 59°31'W, 120–150 m, 21 June 1989, Gillespie 1801 (BRG, CAY, L, MO, US); near Sand Creek, Sep 1948, Wilson-Browne 128 (NY).

Euploca humistrata (Cham.) J.I.M.Melo & Semir, 2009

Fig. 1

Euploca humistrata (Cham.) J.I.M.Melo & Semir, Kew Bull. 64(2): 288. 2009

Heliotropium humistratum Cham., Linnaea 4: 462. 1829.

Type [fide Förther 1998: 200]: Brazil. Minas Gerais: Faz de Piedade, 1818, Sellow s.n. (holotype LE n.v.; isotypes B† [fragment GH n.v.; photo F-17325, GH, NY, US], G-DC!, HAL n.v. [scan!], K!).

Schleidenia humistrata (Cham.) Fresen., Fl. Bras. 8(1): 34. 1857.

Type: Based on Heliotropium humistratum Cham.

Type

Based on Heliotropium humistratum Cham.

Distribution

Prostrate herbs. Stems villose, trichomes hyaline, somewhat ferruginous around the nodes. Leaves: petiole short; lamina lanceolate, 0.5–0.8 × 0.15–0.2 cm, apex acute, base attenuate, margin entire, both surfaces villose. Inflorescence: solitary, supra-axillary flowers, on 1–1.5 mm long pedicel. Flowers: calyx as long as the corolla tube, lobes, 2.2–2.7 mm long at anthesis, puberulent on both sides; corolla tube yellow with white lobes, outside strigose, inside pubescent, tube more or less cylindric, lobes ovate with narrow sinuses, about 1 mm long; stamens sessile, anther with a small appendage; ovary ca. 0.3 mm diam., globose, glabrous; style lacking; stigma sessile, widely conical. Fruit on 1.2–2 mm long pedicel, subglobose, 1.5 mm diam., glabrous, rostrate; nutlets about 1.2 mm long.

Distribution

Guyana; previously only known from a few localities outside the Guiana Shield, Brazil (Ceará, Goiás, Minas Gerais), Venezuela (Apure). In our area it is known only by GUYANA, Upper Takutu–Upper Essequibo, W of the Kanuku Mtns, savanna near Mountain Point, 27 Feb–5 Mar 1985, fl, C. Feuillet 1632 (CAY).

Notes

Melo and Semir (2009: 288) cited the type as “Oliveira 239 (holotype BHCB!; isotype BHMH!)”. This is obviously a misunderstanding. Melo and Semir (2010: 117) cited the holotype, Sellow s.n., as being at GH. In fact at GH is a fragment of the isotype, now destroyed, that was at B. Förther (1998: 200) cites the holotype at LE.

Figure 1.

Euploca humistrata. Guyana, Upper Takutu-Upper Essequibo, Rupununi savanna, savanna near Mountain Point, 27 Feb–5 Mar 1985, Feuillet 1632 (CAY). Picture by Ingrid Lin at US.

Euploca lagoensis (Warm.) Diane & Hilger, in Hilger and Diane, 2003

Fig. 2

Euploca lagoensis (Warm.) Diane & Hilger, in Hilger and Diane: Bot. Jahrb. Syst. 125 (1): 48. 2003.

Schleidenia lagoensis Warm., Kjoeb. Vidensk. Meddel 1867: 15. 1868.

Type: Brazil. Minas Gerais: Lagoa Santa ad ripam lacus, 5 Nov 1864, Warming s.n. (holotype C n.v. [scan!; photos F-21971, M, MO]; isotypes GH n.v. [fragment], P! [scan!]).

Schleidenia subracemosa Warm., Kjoeb. Vidensk. Meddel 1867: 15. 1868.

Type: Brazil. Minas Gerais: Lagoa Santa, 28 Jan 1866, Warming s.n. (holotype C [photo F-21972, GH, US]; isotype P! [scan!]).

Heliotropium lagoense (Warm.) Gürke, in Engler and Prantl, Nat. Pflanzemf. 4(3a): 97. 1893.

Type: Based on Schleidenia lagoensis Warm.

Heliotropium trinitense Urb., Symb. Ant. 7(3): 350. 1912.

Type: Trinidad. in Piarco Savannah, 10 May 1895, Lunt 6030 (holotype B †; lectotype, designated by Förther 1998, pg. 225: S n.v. [scan!]; isotypes [fragments of holotype GH n.v. and lectotype GH n.v.]).

Type

Based on Schleidenia lagoensis Warm.

Description

Annual herbs, up to 15 cm tall, stems decumbent or prostrate, up to 30 cm long, glabrous or with scattered slender appressed trichomes. Leaves: petiole about 1 mm long, glabrous; lamina oblanceolate, 0.5–1.5 × 0.1–0.3 cm, apex acute to acuminate, base attenuate, margin entire, ciliate, adaxially glabrous, abaxially glabrous to puberulent. Flowers single, extra–axillary on leafy stems: pedicel up to 3 mm long, calyx of 5 lanceolate or cuneate, unequal lobes, 1.5–2.5 mm long at anthesis, puberulent in and out; corolla white, yellow at throat, tube swollen at base, 3–4 mm long, puberulent in and out, lobes ovate, 1–1.5 mm long, with sinus rounded, occasionally plaited with a minute lobule; stamens sessile, inserted about 0.5 mm from base, anthers joined, oblong, with an hairy apical appendage nearly as large as the anther; ovary obpyriform, about 0.5 mm diam.; style lacking or confused with the apex of the ovary; stigma sessile or subsessile, much larger at base; fruit rostrate, glabrous or nearly so, about 2 mm diam., obpyriform; nutlets ca. 1.5–2 mm long.

Distribution

Venezuela (Bolívar), Surinam, French Guiana, and Brazil (Amazonas); also Greater Antilles and from Mexico to Brazil and eastern Bolivia. Euploca lagoensis is new to French Guiana. I have seen only few specimens from the Guiana Shield.

Selected specimens studied

Venezuela, Bolívar, near San Carlos, Laguna de la Culebra, 6 Apr 1925, fl, Pittier 11703 (P). Surinam, In 1885, W.F.R. Suringar s.n. (L); Mennega 907 (L). French Guiana, Savane Maillard, 6 km SE de Tonate-Macouria, 4°58'N, 52°26'W, 25 Nov 1999, Raynal-Roques and Jérémie 24694 (CAY, K, P, NY).

Note

Melo and Semir (2010) cited the holotype as being at M. I do not think there is a specimen at M, only a photograph. Furthermore, if Warming worked in Munich, it was about 7 years after he described Schleidenia lagoensis. In the 1867 he was working in Copenhagen, I do not see any reason to reject Förther’s (1998) choice of the specimen at C as the holotype.

Figure 2.

Euploca lagoensis. French Guiana, SE of Tonate-Macouria, Savane Maillard, 25 Nov 1999, Raynal-Roques and Jérémie 24694 (CAY). (Picture by Piero Delprete at CAY).

Euploca polyphylla (Lehm.) J.I.M.Melo & Semir, 2009

Euploca polyphylla (Lehm.) J.I.M.Melo & Semir, Kew Bull. 64(2): 289. 2009.

Heliotropium polyphyllum Lehm., Neue Schriften Naturf. Ges. Halle 3(2): 9. 1817.

Type (fide Förther 1998: 214). Venezuela. near Orinoco, Humboldt s.n. (holotype MEL 233310 n.v.; isotypes B-Willd 3252 n.v., P! [scan!], S 11-21939 n.v. [scan!]).

Heliotropium foliosum Roem. & Schult., Syst. Veg. 4: 737. 1819.

Type. Venezuela. Orinoco, Apr 1800, Humboldt and Bonpland 808 (holotype B-WILLD 3252-01 n.v.).

Preslaea linifolia A.St.-Hil., Voy. Distr. Diam. 2: 433. 1833.

Type. Brazil. Cabo Frio, St.-Hilaire 433 (lectotype, designated by Förther 1998, pg. 234: P!; isotypes B †, M n.v., MPU!, P! [2 sheets]).

Heliotropium bahiense DC., Prodr. 9: 544. 1845.

Syntype: Brazil. Blanchet 92 (G n.v., NY!, P!). Type. Brazil. Bahia, 1836, Salzmann 368 (lectotype, designated by Förther 1998, pg. 181: G-DC n.v.; isotypes B †, F n.v., H n.v., HAL n.v., K!, M n.v., MPU!, P! [4 sheets]).

Heliotropium polyphyllum var. blanchetii DC., Prodr. 9: 544. 1845.

Type. Brazil. Bahia, Serra Jacobina, 1837, Blanchet 2651 (lectotype, designated by Förther 1998, pg. 214: G-DC n.v.; isotypes B† [photo F-17340, GH, NY, US], BM n.v. [fragment GH n.v.], BR n.v., F-520830 n.v. [photo F-57735], FI-W!, HAL n.v., K! [scan !], LE n.v., M n.v., NY!, P! [2 sheets], TUB n.v., US-292391!, W-118162 n.v.).

Schleidenia linifolia (A.St.-Hil.) DC., Prodr. 9: 558. 1845.

Type. Based on Preslaea linifolia A.St.-Hil.

Schleidenia pubescens Fresen., Fl. Bras. 8(1): 35. 1857.

Type. Brazil. Rio de Janeiro, Gaudichaud [524] (lectotype, designated by Förther 1998, pg. 236: P!; isotype B†).

Schleidenia polyphylla (Lehm.) Fresen., Fl. Bras. 8(1): 36. 1857.

Type. Based on Heliotropium polyphyllum Lehm.

Schleidenia polyphylla var. blanchetii (DC.) Fresen., Fl. Bras. 8(1): 36. 1857.

Type. Based on Heliotropium polyphyllum var. blanchetii DC.

Schleidenia bahiensis (DC.) Fresen., Fl. Bras. 8(1): 43. 1857.

Type. Based on Heliotropium bahiense DC.

Heliotropium polyphyllum var. laevenworthii A.Gray, Syn. Fl. N. Amer. 2(1): 185. 1878.

Syntypes: USA. Florida: Everglades, Leavenworth s.n.; Palmer s.n. (F). Type. USA. Florida: Everglades, 1877, Garber s.n. (lectotype, designated by Förther 1998, pg. 214: GH n.v.; isotype K n.v. [scan!]).

Heliotropium pubescens (Fresen.) Gürke, in Engler and Prantl, Nat. Pflanzenfam. 4(3a): 97. 1893.

Type. Based on Schleidenia pubescens Fresen.

Heliotropium linifolium (A.St.-Hil.) Gürke, in Engler and Prantl, Nat. Pflanzenfam. 4(3a): 97. 1893; illegitimate: not Lehm. 1818.

Type. Based on Preslaea linifolia A.St.-Hil.

Heliotropium laevenworthii (A.Gray) Small, Fl. S. E. U.S.: 1006. 1903.

Type. Based on Heliotropium polyphyllum var. laevenworthii A.Gray

Heliotropium horizontale Small, Bull. New York Bot. Gard. 3: 435. 1905.

Type. USA. Florida: 9–12 Nov 1903, Small and Carter 742 (holotype NY! [photo GH]; isotypes F n.v., GH n.v. [fragment], US!).c> (holotype NY! [photo GH]; isotypes F n.v., GH n.v. [fragment], US!).

Heliotropium polyphyllum var. horizontale (Small) R.W.Long, Rhodora 72: 33. 1970.

Type. Based on Heliotropium horizontale Small

Type

Based on Heliotropium polyphyllum Lehm.

Description

Herb or subshrub. Stems prostrate or decumbent, sericeous. Leaves alternate or subopposite: petiole 0.03–0.1 cm long, sericeous; lamina narrow lanceolate or oblanceolate, 0.6–1.3 × 0.1–0.2 cm, apex acute, base cuneate, margin entire, both surfaces sericeous. Inflorescences single or paired, cymose, 1–15 cm long, bracts ovate, 0.3–0.4 cm long, sericeous inside, glabrous outside. Flower sub–sessile: calyx slightly shorter than the corolla tube, lobes, 3–4 mm long at anthesis, glabrous inside, sericeous outside; corolla yellow or white, tubular, outside sericeous, inside sericeous at throat, tube widened in the middle, lobes ovate-deltoid, with narrow sinuses, 2.5–3 mm long; stamens subsessile, anther with a long appendage; ovary subglobose, 4-sulcate, glabrous; style 0.3–0.6 mm long; stigma conical. Fruit subglobose, entirely covered by the calyx; nutlets almost 1.5 mm diam.

Distribution

Venezuela (Bolívar), Brazil (Amapá, Amazonas, Pará); also USA (Florida), Bahamas, Venezuela, eastern Brazil.

Euploca procumbens (Mill.) Diane & Hilger, in Hilger and Diane, 2003

Euploca procumbens (Mill.) Diane & Hilger, in Hilger and Diane: Bot. Jahrb. Syst. 125 (1): 48. 2003.

Heliotropium procumbens Mill., Gard. Dict. ed. 8, Heliotropium n°10. 1768.

Type. Jamaica. herb. Miller (coll. Houston?) s.n. (lectotype, designated by Howard 1989, pg. 204: BM n.v. [photo GH!]).

Heliotropium americanum Mill., Gard. Dict. ed. 8, Heliotropium n°11. 1768.

Type. Mexico. Vera Cruz, 1731, Houston s.n. (lectotype, designated by Förther 1998, pg. 178: BM n.v.).

Heliotropium inundatum Sw., Prodr. 40. 1788.

Type (fide Förther 1998: 201). Jamaica. “insulae caribaeae”, probably a Swartz coll. (holotype S?).

Heliotropium decumbens Lehm., Neue Schr. Naturf. Ges. Halle 3(2): 16. 1817.

Type. Venezuela. Sucre: Cumana, Humboldt and Bonpland 57 (lectotype, designated by Förther 1998, pg. 190; B-WILLD 3239! [scan!]; isotypes MEL n.v., HAL n.v. [scan!], P! [2 sheets; scans!], P-Bonpl!) [same type collection as Heliotropium procumbens Kunth (below), but different holotype].

Heliotropium canescens Lehm., Pl. Asperif. Nucif. 38. 1818; illegitimate: not Moench 1794.

Type. Brazil. Sieber pro Hoffmannsegg s.n. (holotype B-WILLD 3242 n.v.; isotypes MEL 233279 n.v., HAL 71587 n.v. [scan!], P! [scan!], S n.v. [scan!]).

Heliotropium procumbens Kunth, Nov. Gen. Sp. (quarto ed.) 3: 88. 1818; nom. illeg. non Mill. 1768 + homotypic (different holo-) with H. decumbens Lehm. 1817.

Type. Venezuela. Sucre: Cumana, Humboldt and Bonpland 57 (holotype P-Bonpl!; isotypes MEL n.v., HAL n.v., P! [2 sheets], B-WILLD 3239!) [same type collection as Heliotropium decumbens Lehm. (above), but different holotype].

Heliotropium canescens Kunth, Nov. Gen. Sp. (quarto ed.) 3: 88. 1818; nom. illeg. non Moench. 1794: 415.

Type. Venezuela. near Cumana, Humboldt 58 (holotype P!; isotype B-WILLD 3240/1!).

Heliotropium cinereum Kunth, Nov. Gen. Sp. (quarto ed.) 3: 89. 1818.

Type. Venezuela. [Apure:] bank of the Apures, propre Arichuna, Mar, Bonpland and Humboldt 1202 (holotype P-Bonpl.!; isotypes B-WILLD 3240/2!, F n.v., P! [2 sheets]). [Same type collection as H. humboldtianum below, but different holotype]

Heliotropium humboldtianum Roem. & Schult., Syst. Veg. 4: 737. 1819.

Type. Venezuela. [Apure:] bank of the Apures, propre Arichuna, Mar, Bonpland and Humboldt 1202 (lectotype, designated by Förther 1998, pg. 200: B-WILLD 3240/2!; isotypes F n.v., P! [2 sheets], P-Bonpl!). [Same type collection as H. cinereum above, but different holotype]

Heliotropium brasilianum Roth, Nov. Pl. Sp.: 103. 1821.

Type. Brazil. Mertens s.n. (holotype B † [photo F-17314, GH, NY, US]).

Heliotropium riparium Mart. ex Colla, Herb. Pedem. 4: 226. 1835.

Type. Brazil. Rio Belmonte, Martius s.n. (holotype TO n.v.; isotypes BM n.v., BR n.v., K!).

Heliotropium willdenowii G. Don, Gen. Hist. 4: 359. 1838.

Type. A renaming of illegitimate Heliotropium canescens Lehm.

Heliotropium rigidulum DC., Prodr. 9: 543. 1845.

Type. Mexico. near Matamoros, July 1831, Berlandier pl. exs. 234 (holotype G-DC n.v.; isotypes BM n.v., BP n.v., F n.v. [scan!], G-DC n.v., LE n.v., P! [2 sheets, scans!], W n.v.).

Heliotropium houstonii DC., Prodr. 9: 549. 1845; nom. illeg. renaming of H. procumbens Mill.

Type. Based on Heliotropium procumbens Mill.

Schleidenia elliptica Fresen., Fl. Bras. 8(1): 42. 1857.

Type. Brazil. Minas Gerais: near Salvado, Aug 1819, Martius s.n. (lectotype, designated by Förther 1998, pg. 235: M n.v.; isotype M n.v.).

Schleidenia longepetiolata Fresen., Fl. Bras. 8(1): 42. 1857.

Type. Brazil. Goyaz: Rio Maranhao, Pohl 2352 and 2384 (sic! distributed as 1593) (lectotype, designated by Förther 1998, pg. 236: W n.v. [photo F-31909, GH, US]; isotypes F n.v. [scan!; photo F-57726], K! [scan!], W n.v.).

Schleidenia inundata (Sw.) Fresen., Fl. Bras. 8(1): 43. 1857.

Type. Based on Heliotropium inundatum Sw.

Schleidenia leptostachya Fresen., Fl. Bras. 8(1): 43. 1857.

Syntype: Brazil. Bahia: Blanchet 3610 (G n.v., GH n.v. [fragments, scan!], NY!, P! [3 sheets, scans!]). Type. Brazil. Bahia: near Joazeiro, Apr 1819, Martius s.n. (lectotype, designated by Förther 1998, pg. 235: M [photo F-20310, NY, GH]; isotypes FI-W!, GH [fragment], M [2 sheets]).

Heliotropium ellipticum (Fresen.) Gürke, in Engler and Prantl, Nat. Pflanzenfam.. 4(3a): 97. 1893.

Type. Based on Schleidenia elliptica Fresen.

Heliotropium leptostachyum (Fresen.) Gürke, in Engler and Prantl, Nat. Pflanzenfam. 4(3a): 97. 1893.

Type. Based on Schleidenia leptostachya Fresen.

Heliotropium longepetiolatum (Fresen.) Gürke, in Engler and Prantl, Nat. Pflanzenfam. 4(3a): 97. 1893.

Type. Based on Schleidenia longepetiolata Fresen.

Heliotropium bridgesii Rusby, Mem. Torrey Bot. Club 4: 224. 1895.

Type. Bolivia. Cochabamba, 1891, Bang 950 (holotype NY!; isotypes B †, BM n.v., GH n.v., K n.v., LE n.v., M n.v., MO n.v. [scan!], NY!, US! [2 sheets], W n.v.).

Heliotropium riparium Chodat, Bull. Herb. Boiss., sér. 2, 2: 817. 1902; nom. illeg. non Mart. ex Colla 1835.

Type. Paraguay. “ad ripam lacus Ypacaray”, Hassler 3893 (holotype G!; isotype P§

Type

Based on Heliotropium procumbens Mill.

Description

Annual herbs up to 50 cm long, usually cinereous; stems erect or decumbent, sericeous, trichomes appressed. Leaves: petiole slender, 0.4–2.4 cm long; lamina elliptic, obovate or broadly oblanceolate, 1–6 × 0.3–2 cm, apex acute, mucronate, base attenuate, margin entire, both surfaces sericeous. Inflorescences terminal or axillary, peduncle 1–3 cm long, slender scorpioid cymes, mostly geminate or ternate, bractless, up to 10 cm long, sericeous. Flower subsessile: calyx deeply lobed, lobes unequal, lanceolate to linear, about 2/3 as long as corolla tube; corolla white, tube 1–1.4 mm long, throat yellow, lobes spreading ovate with rounded sinus, < 0.5 mm long; stamens subsessile, anthers ovate with a small apical appendage, free; ovary about 0.3 mm diam., style lacking; stigma sessile forming with the ovary a blunt cone. Fruit: depressed-globose, 4-lobed, strigose; nutlets up to 1 mm long.

Distribution

Venezuela (Amazonas, Bolívar, Delta Amacuro), Guyana, Brazil (Pará, Roraima); also West Indies and from Southern United States to Argentina.

Selected specimens studied

Guyana. s.l., Appun 1762 (K); 1839, Rob. Schomburgk 1024 (K); Schomburgk 1026 (K); Upper Takutu-Upper Essequibo, NW Kanuku Mtns, near Nappi Village, 13 Feb 1993, Hoffman 3743 (BRG, F, L, US).

Note

Förther (1998: 200) placed Heliotropium humboltianum Roem. & Schult. 1819 in the synonymy of H. procumbens. He designated as the lectotype Humboldt and Bonpland [1202]. Humboldt 1202 is the type of H. filiforme (see above under E. filiformis).

Names not validly published

Messerschmidia incana G. Mey. 1818, nom. inval.

Messerschmidia incana G. Mey., Prim. Fl. Esseq. 92. 1818, nom. inval.

Type

not designated; no original material found.

Note

The only species known from the Guianas that would agree with Meyer’s description is E. humilis. Förther (1998: 238) lists M. incana as a synonym of H. ternatum [=Euploca humilis] with a question mark, but Meyer placed a synonym in the protolog of his new species, Tournefortia sessilifolia Poir. (1804: 360), a species described from Argentina. Because sessilifolia was available in Messerschmidia, it should have been used as the epithet. Therefore the name by Meyer is not validly published.

Acknowledgements

I want to thank the editor Sandy Knapp for her patience with my struggle with the format. I am indebted to the curators of the herbaria cited who made it easy to get loans, or to work in the collection in their care or who provided scans of types. I want to thank Ingrid Lin (US) and Piero Delprete (CAY) who scanned the specimens shown in Fig. 1 and 2, and the reviewers. I am very grateful to the Department of Botany, Smithsonian Institution, where I was welcomed from 1988 to 2014 and the Department of Botany and Plant Pathology, Oregon State University, where I do research since the summer of 2014. This is number 215 in the Smithsonian’s Biological Diversity of the Guiana Shield publication series.

References

Brown R (1810) Prodromus Florae Novae Hollandiae et Insulae van-Diemen. Johnson, London, I–VIII, 145–590. [1–144 were not published]http://www.biodiversitylibrary.org/item/21871#page/6/mode/1up

Burmann J (1760) Plantarum Americanarum 10. Schouten, Leiden, 221–262; t. 227–262. http://bibdigital.rjb.csic.es/spa/Libro.php?Libro=6191

Cafferty S, Jarvis CE (2004) Typification of Linnaean plant names in Boraginaceae. Taxon 53(3): 799–805. doi: 10.2307/4135454

Candolle AP de (1845) Borragineae.Prodromus vol. 9. Fortin and Masson, Paris, 466–559. http://www.biodiversitylibrary.org/item/7158#page/5/mode/1up

Chamisso A de (1829) Boragineae Juss. Linnaea 4: 435–496. http://www.biodiversitylibrary.org/item/10848

Chodat R (1902) Plantae hasslerianae soit Énumération des plantes récoltées au Paraguay par le Dr Émile Hassler. Bulletin de l’herbier Boissier, sér. 2, 2(9): 811–824. http://www.biodiversitylibrary.org/item/105265#page/7/mode/1up

Colla LA (1835) Herbarium Pedemontanum, Vol. 4. Typis Regiis, Turin, 1–592. http://bibdigital.rjb.csic.es/ing/Libro.php?Libro=5587

Craven LA (1996) A taxonomic revision of Heliotropium (Boraginaceae) in Australia. Australian Systematic Botany 9(4): 521–657. doi: 10.1071/SB9960521

Degen R, Melo JIM (2012) Nuevas combinaciones en el género Euploca Nutt. (Boraginaceae) del Paraguay. Candollea 67(2): 368. doi: 10.15553/c2012v672a16

Don G (1838) Boraginae, pg. 306–374. A general history of the dichlamydeous plants vol. 4. Rivington, London, I–VIII, 1–908. http://www.biodiversitylibrary.org/item/9906#page/2/mode/1up

Endlicher SFL (1839) Genera plantarum secundum ordines naturales disposita. Beck, Vienna, I–LX, 1–1484. http://bibdigital.rjb.csic.es/ing/Libro.php?Libro=1669

Europeana [consulted 31 March 2014] http://www.europeana.eu/portal/

Fenzl E (1886) Vier neue Pflanzenarten Süd-Amerikas. Verhandlungen der Kaiserlich-Koniglichen Zoologisch-Botanischen Gesellschaft in Wien 36: 287–294. http://www.biodiversitylibrary.org/item/47596#page/413/mode/1up

Feuillet C (2012) Boraginaceae. In: Acevedo-Rodríguez P, Strong MT (Eds) Catalogue of Seed Plants of the West Indies. Smithsonian Institution Scholarly Press, Washington, 157–174. http://hdl.handle.net/10088/17551

Förther H (1998) Die infragenerische Gliederung der Gattung Heliotropium L. und ihre Stellung innerhalb der subfam. Heliotropioidea (Schrad.) Arn. (Boraginaceae). Sendtnera 5: 35–241. http://www.biodiversitylibrary.org/item/52056#page/1/mode/2up

Fresenius G (1857) Heliotropieae. Flora brasiliensis 8(1): 29–58, t. 10–13. http://florabrasiliensis.cria.org.br/opus?vol=8&part=1

Frohlich MW (1981) Heliotropium. In: Nash DL, Moreno NP (Eds) Boraginaceae.Flora of Veracruz 18: 70–104.

Gray A (1857) Diagnostic characters of new species of phanerogamous plants. Memoirs of the American Academy of Arts and Sciences, ser. 2, 6: 377–452. http://www.biodiversitylibrary.org/item/49960#page/8/mode/2up

Gray A (1874) Notes on Borraginaceae. Proceedings of the American Academy of Arts and Sciences 10: 48–62. http://www.biodiversitylibrary.org/item/22173#page/70/mode/1up

Gray A (1878) Synoptical Flora of North America 2(1). American Book, New York, I–XV, 1–506. http://www.biodiversitylibrary.org/item/42613#page/5/mode/1up

Greenman JM (1898) Diagnoses of new and critical Mexican phanerogams. Proceedings of the American Academy of Arts and Sciences, 471–189. http://www.biodiversitylibrary.org/item/22279#page/9/mode/1up

Gürke RLAM (1893) Borraginaceae. In: Engler A, Prantl K. Die natürlichen Pflanfamilien 4(3a). Engelman, Leipzig, 71–131. http://www.biodiversitylibrary.org/item/16413#page/7/mode/1up

Hilger HH, Diane N (2003) A systematic analysis of Heliotropiaceae (Boraginales) based on trnL and ITS1 sequence data. Botanische Jahrbuecher fuer Systematik Pflanzengeschichte und Pflanzengeographie 125(1): 19–51. http://www.ingentaconnect.com/content/schweiz/bj/2003/00000125/00000001/art00002

Howard RA (1989) Boraginaceae. Flora of the Lesser Antilles 6(3): 188–211.

Humboldt FWHA von, Bonpland A, Kunth CS (1818) Nova Genera et Species Plantarum vol. 3. Librairie grecque-latine-allemande, Paris, 1–456, t. 193–300. http://bibdigital.rjb.csic.es/ing/Libro.php?Libro=5760

Johnston IM (1928) Studies in the Boraginaceae VII. The South American Species of Heliotropium. Contributions from the Gray Herbarium 81: 3–73. http://www.biodiversitylibrary.org/item/122400#page/1/mode/1up

Johnston IM (1930) Studies in the Boraginaceae VIII, I.3. Treatment of Tournefortia. Contributions from the Gray Herbarium 92: 66–89. http://www.biodiversitylibrary.org/item/123764#page/111/mode/1up

Johnston IM (1935) Studies in the Boraginaceae X. The Boraginaceae of the northeastern South America. Journal of the Arnold Arboretum 16: 1–64. http://www.biodiversitylibrary.org/item/33592#page/1/mode/1up

Johnston IM (1949) Studies in the Boraginaceae XVIII. Boraginaceae of the southern West Indies. Journal of the Arnold Arboretum 30: 111–188. http://www.biodiversitylibrary.org/part/18048#/summary

JSTOR Plant Science [consulted 31 March 2014] http://plants.jstor.org/

Kunth KS (1818) Boragineae Juss. Nova Genera et Species Plantarum (quarto ed.) 3: 65–93.

Lamarck JB (1791) Tableau Encyclopédique et Méthodique des trois Règnes de la Nature. Botanique 1. Heliotropium. Panckoucke, Paris, 392–394. doi: 10.5962/bhl.title.218

Lehmann JGC (1817a) Decas novarum e Borraginearum familia plantarum. Göttingische Gelehrte Anzeigen unter der Aufsicht der Konigl. Gesellschaft der Wissenschaften 152: 1513–1515.

Lehmann JGC (1817b) Beschreibung einiger neuer und wenig bekannter Pflanzen. Neue Schriften der naturforschenden Gesellschaft zu Halle 3(2): 1–26.

Lehmann JGC (1818) Heliotropium. Pl. Asperif. nucif. 1: 19–74. http://gallica.bnf.fr/ark:/12148/bpt6k98096m/f47.image.zoom

Linnaeus C (1753) Species Plantarum, 2 vol.Laurentius Salvius, Stockholm, 1–1231. Vol. 1 [http://www.biodiversitylibrary.org/item/13829#page/1/mode/1up] and vol. 2 [http://www.biodiversitylibrary.org/item/13830#page/2/mode/1up]

Long RW (1970) Additions and nomenclatural changes in the flora of southern Florida 11. Rhodora 72: 17–46. http://www.biodiversitylibrary.org/item/14543#page/1/mode/1up

Luebert F, Brokamp G, Wen J, Weigend M, Hilger HH (2011) Phylogenetic relationships and morphological diversity in Neotropical Heliotropium (Heliotropiaceae). Taxon 60(3): 663–680. http://docserver.ingentaconnect.com/deliver/connect/iapt/00400262/v60n3/s4.pdf?expires=1360244727&id=72763714&titleid=6069&accname=Smithsonian+Institution+Libraries&checksum=CB163B8EBE40FED399C422AF1014AE1A

Martius CFP von (1827) Nova Genera et Species Plantarum 2. Wolf, Munich, 1–148. http://www.biodiversitylibrary.org/item/9620#page/148/mode/1up

Martius CFP von (1838) Martii herbarium florae brasiliensis continuatis. Flora 21(2), beibl. 4: 49–96. http://www.biodiversitylibrary.org/item/931#page/1/mode/1up

Martius CFP von (1857) Flora brasiliensis vol. 8(1). Fleischer, Leipzig, 1–448, t. 1–68. http://florabrasiliensis.cria.org.br/opus?vol=8&part=1

McNeill J, Barrie FR, Buck WR, Demoulin V, Greuter W, Hawksworth DL, Herendeen PS, Knapp S, Marhold K, Prado J, Prud’Homme van Reine WF, Smith GF, Wiersema JH, Turland NJ (2012) International Code of Nomenclature for Algae, Fungi, and Plants (Melbourne Code). Regnum Vegetabile 154. Koeltz, Königstein, I–XXX, 1–208. http://www.iapt-taxon.org/nomen/main.php

Melo JIM, Semir J (2006) A new species of Euploca (Heliotropiaceae) from Brazil. Candollea 61(2): 453–456. http://www.ville-ge.ch/cjb/publications/cando612/C612_453-456.pdf

Melo JIM, Semir J (2009) Two new species and new combinations in Euploca (Heliotropiaceae). Kew Bulletin 64(2): 285–289. doi: 10.1007/s12225-009-9109-3

Melo JIM, Semir J (2010) Taxonomia do gênero Euploca Nutt. (Heliotropiaceae) no Brasil. Acta Botanica Brasilica 24(1): 111–132. doi: 10.1590/S0102-33062010000100012

Meyer GFW (1817) Primitiae Florae Essequiboensis. Dieterich, Göttingen, I–X, 1–316. http://www.biodiversitylibrary.org/item/104399#page/5/mode/1up

Miller P (1768) Gardeners dictionary, 8th edition. The author, London, unpaged. http://www.biodiversitylibrary.org/item/10276#page/2/mode/1up

Moench C (1794) Heliotropium. Methodus Plantas Horti Botanici et Agri Marburgensis vol. 2. Academy Library, Marburg, 415.

Nuttall T (1836) Collections towards a Flora of the Territory of Arkansas. Transcriptions of the American Philosophical Society, new series 5: 139–203. http://www.jstor.org/stable/1004943

Opiz PM (1824) Naturalientausch 8: 86.

Poiret JLM (1804) Pittone. Encyclopédie Méthodique. Botanique 5: 355–362.

Rafinesque CS (1838) Sylva telluriana. The author, Philadelphia, 1–194. http://www.biodiversitylibrary.org/item/62551#page/5/mode/1up

Reichenbach HGL (1841) Der Deutsche Botaniker. Das Herbarienbuch. Arnoldischen, Dresden, II–XCV, 1–236. http://www.biodiversitylibrary.org/item/32106#page/7/mode/1up

Roemer JJ, Schultes JA (1819) Systema vegetabilium vol 4. Cottae, Stuttgart, I–LX, 1–888. http://bibdigital.rjb.csic.es/ing/Libro.php?Libro=1701

Roth AW (1821) Novae Plantarum species praesertim Indiae orientalis. Vogler, Halberstadt, I–IV, 1–412. http://www.biodiversitylibrary.org/item/41813#page/1/mode/1up

Rusby HH (1895) An enumeration of the plants collected in Bolivia by Miguel Bang, with descriptions of new genera and species. Memoirs of the Torrey Botanical Club 4: 203–274. http://www.biodiversitylibrary.org/item/99826#page/1/mode/1up

Saint-Hilaire AFCP de (1933) Voyage dans le district des diamans, vol. 2. Gide, Paris, I–III, 1–456. https://archive.org/stream/voyagesdanslint02hilgoog#page/n6/mode/1up

Schomburgk MR (1849) Reisen in Britisch-Guiana in den Jahren 1840-1884. Weber, Leipzig, 534–1260http://www.biodiversitylibrary.org/item/109594#page/11/mode/1up

Schrader HA (1819) De asperifoliis Linnei comentatis recitata. Commentationes Societatis Regiae Scientiarum Gottingensis Recentiores 4: 175–196. https://books.google.com/books?id=8FvNAAAAMAAJ&printsec=frontcover&dq=inauthor:%22K%C3%B6nigliche+Gesellschaft+der+Wissenschaften+zu+G%C3%B6ttingen%22&hl=en&sa=X&ved=0CDgQ6AEwA2oVChMIrtWQ4fvLxwIVwy2ICh3OOwWv#v=onepage&q&f=false

Small JK (1903) Flora of southeastern U.S. The author, New York, VII–XII, 1–1370. http://www.biodiversitylibrary.org/item/56866#page/7/mode/1up

Small JK (1905) Additions to the flora of subtropical Florida. Bulletin of the New York Botanical Garden 3(3): 419–440. http://www.biodiversitylibrary.org/item/97293#page/7/mode/1up

Small JK (1933) Manual of the Southeastern Flora. University of North Carolina Press, Chapel Hill, 1–1554. http://www.biodiversitylibrary.org/item/13847#page/1/mode/1up

Spach É (1838) Histoire naturelle des végétaux. Roret, Paris, 1–586. http://gallica.bnf.fr/ark:/12148/bpt6k6149996b.r=dolichos+lablab.langFR

Stafleu FA, Cowan RS (1985) Taxonomic literature, 2nd edition, 5: Sal–Ste. Scheltema, Bohn and Holkema, Utrecht, 1–1066. http://www.sil.si.edu/digitalcollections/tl-2/browse.cfm?vol=5#page/11

Swartz O (1788) Nova Genera et Species Plantarum seu Prodromus descriptionum vegetabilium. Sweder, Stockholm, 1–152. http://www.biodiversitylibrary.org/item/23300#page/9/mode/1up

Urban I (1912) Nova genera et species, part 5. Symbolae antillanae 7(3): 151–432. http://www.biodiversitylibrary.org/item/3757#page/1/mode/1up

Urban I (1929) Plantae haitenses et domingenses novae vel rariores VI. Arkiv för Botanik 22 A(10): 1–108.

Vahl MH (1794) Symbolae botanicaepart 3. Möller, Copenhagen, 1–106, t. 51–75. http://www.biodiversitylibrary.org/item/118639#page/313/mode/1up

Vatke W (1875) Plantas in itinere africano ab J. M. Hildebrandt collectas determinare pergit W. Vatke. Oesterreichische Botanische Zeitschrift 25(1): 166–169.

Warming JEB (1868) Symbolae ad floram brasiliae centralis cognoscendam, part 1. Videnskabelige Meddelelser fra den Naturhistoriske Forening i Kjøbenhavn 1867: 15: 1–24. http://www.biodiversitylibrary.org/bibliography/7547

Wiggins IL (1950) Taxonomic notes on plants from the Sonoran desert. Contributions from the Dudley herbarium 4(2): 13–31.

List of taxa

Euploca Nutt. 1836

Euploca convolvulacea Nutt. (type of Euploca)

Euploca filiformis (Lehm.) J.I.M.Melo & Semir 2009

Euploca foliata (R. Br.) Feuillet comb. nov.

Euploca humilis (L.) Feuillet, comb. nov.

Euploca humistrata (Cham.) J.I.M.Melo & Semir 2009

Euploca hypogaea (Urb. & Ekman) Diane & Hilger

Euploca lagoensis (Warm.) Diane & Hilger 2003

Euploca paradoxa (Mart.) J.I.M.Melo & Semir

Euploca polyphylla (Lehm.) J.I.M.Melo & Semir 2009

Euploca procumbens (Mill.) Diane & Hilger 2003

Euploca ternata (Vahl) J.I.M.Melo & Semir 2009 = Euploca humilis

Heliophytum passerinoides Klotzsch 1849 = Euploca humilis

Heliotropium subgen. Orthostachys (R.Br.) Rchb. 1841 = Euploca

Heliotropium sect. Orthostachys R.Br. 1810 = Euploca

Heliotropium sect. Orthostachys subsect. Axillaria I.M.Johnst. 1928 = Euploca

Heliotropium americanum Mill. 1768 = Euploca procumbens

Heliotropium bahiense DC. 1845 = Euploca polyphylla

Heliotropium brasilianum Roth 1821 = Euploca procumbens

Heliotropium bridgesii Rusby 1895 = Euploca procumbens

Heliotropium canescens Kunth 1818 = Euploca procumbens

Heliotropium canescens Lehm. 1818 = Euploca procumbens

Heliotropium cinereum Kunth 1818 = Euploca procumbens

Heliotropium decumbens Lehm. 1817 = Euploca procumbens

Heliotropium ellipticum (Fresen.) Gürke 1893 = Euploca procumbens

Heliotropium filiforme Kunth 1818 = Euploca filiformis

Heliotropium filiforme Lehm. 1817 = Euploca filiformis

Heliotropium foliatum R.Br. 1810 (type of Heliotropium sect. Orthostachys (R.Br.) G.Don) = Euploca foliata

Heliotropium foliosum Roem. & Schult. 1819 = Euploca polyphylla

Heliotropium fruticosum var. ternatum (Vahl) DC. 1845 = Euploca humilis

Heliotropium fumana (Fresen.) Gürke 1893 = Euploca humilis

Heliotropium greenmanii Wiggins 1950. = Euploca humilis

Heliotropium helophilum Mart. 1838 = Euploca filiformis

Heliotropium hirtum Lehm. 1817 = Euploca humilis

Heliotropium hispidum Kunth 1818 = Euploca humilis

Heliotropium horizontale Small, Bull. 1905 = Euploca polyphylla

Heliotropium houstonii DC. 1845 = Euploca procumbens

Heliotropium humboldtianum Roem. & Schult. 1819 = Euploca procumbens

Heliotropium humboltianum Roem. & Schult. 1819 = Euploca procumbens

Heliotropium humistratum Cham. 1829 = Euploca humistrata

Heliotropium inundatum Sw. 1788 = Euploca procumbens

Heliotropium laevenworthii (A. Gray) Small 1903 = Euploca polyphylla

Heliotropium lagoense (Warm.) Gürke 1893 = Euploca lagoensis

Heliotropium leptostachyum (Fresen.) Gürke 1893 = Euploca procumbens

Heliotropium linifolium (A. St.-Hil.) Gürke 1893 = Euploca polyphylla

Heliotropium littorale Mart. ex Colla 1835 = Euploca filiformis (Lehm.) J.I.M.Melo & Semir

Heliotropium longepetiolatum (Fresen.) Gürke 1893 = Euploca procumbens

Heliotropium mexicanum Greenm. 1898 = Euploca humilis

Heliotropium oaxacanum DC. 1845 = Euploca humilis

Heliotropium paradoxum (Mart.) Gürke 1893 = Euploca paradoxa

Heliotropium polyphyllum Lehm. 1817 = Euploca polyphylla

Heliotropium polyphyllum var. blanchetii DC. 1845 = Euploca polyphylla

Heliotropium polyphyllum var. horizontale (Small) R.W. Long 1970 = Euploca polyphylla

Heliotropium polyphyllum var. laevenworthii A. Gray 1878 = Euploca polyphylla

Heliotropium procumbens Kunth 1818 = Euploca procumbens

Heliotropium procumbens Mill. 1768 = Euploca procumbens

Heliotropium pubescens (Fresen.) Gürke 1893 = Euploca polyphylla

Heliotropium pusillum Colla 1835 = Euploca filiformis

Heliotropium rigidulum DC. 1845. = Euploca procumbens

Heliotropium riparium Chodat 1902 = Euploca procumbens

Heliotropium riparium Mart. ex Colla 1835 = Euploca procumbens

Heliotropium sect. Euploca (Nutt.) A. Gray 1874 = Euploca

Heliotropium stenostachyum (A. St.-Hil.) Gürke 1893 = Euploca filiformis

Heliotropium strictissimum sensu N.E. Br. 1901 = Euploca humilis

Heliotropium tenue Roem. & Schult. 1819 = Euploca filiformis

Heliotropium ternatum Vahl 1794 = Euploca humilis

Heliotropium ternatum var. fumana (Fresen.) I.M. Johnst. 1935 = Euploca humilis

Heliotropium trinitense Urb. 1912 = Euploca lagoensis

Heliotropium willdenowii G. Don 1838 = Euploca procumbens

Hilgeria Förther 1989 = Euploca

Hilgeria hypogaea (Urb. & Ekman) Förther 1989 (type of Hilgeria Förther) = Euploca hypogaea

Messerschmidia incana G. Mey. 1818 = Euploca humilis

Orthostachys (R. Br.) Spach 1840 = Euploca

Pioctonon antillanum Raf. 1838 = Euploca humilis

Pioctonon ternatum (Vahl) Raf. 1838 = Euploca humilis

Preslaea Mart. 1827 = Euploca

Preslaea linifolia A. St.-Hil. 1833 = Euploca polyphylla

Preslaea paradoxa Mart. 1827 (type of Preslaea Mart.) = Euploca paradoxa

Preslaea stenostachya A. St.-Hil. 1833 = Euploca filiformis

Schleidenia Endl. 1839 = Euploca

Schleidenia bahiensis (DC.) Fresen. 1857 = Euploca polyphylla

Schleidenia elliptica Fresen. 1857 = Euploca procumbens

Schleidenia filiformis (Lehm.) Fresen. 1857 = Euploca filiformis

Schleidenia fumana Fresen. 1857 = Euploca humilis

Schleidenia hispida (Kunth) Fresen. 1857 = Euploca humilis

Schleidenia humistrata (Cham.) Fresen. 1857 = Euploca humistrata

Schleidenia inundata (Sw.) Fresen. 1857 = Euploca procumbens

Schleidenia lagoensis Warm. 1868 = Euploca lagoensis

Schleidenia leptostachya Fresen. 1857 = Euploca procumbens

Schleidenia linifolia (A. St.-Hil.) DC. 1845 = Euploca polyphylla

Schleidenia longepetiolata Fresen. 1857 = Euploca procumbens

Schleidenia polyphylla (Lehm.) Fresen. 1857 = Euploca polyphylla

Schleidenia polyphylla var. blanchetii (DC.) Fresen. 1857 = Euploca polyphylla

Schleidenia pubescens Fresen. 1857 = Euploca polyphylla

Schleidenia stenostachya (A. St.-Hil.) DC. 1845 = Euploca filiformis

Schleidenia subracemosa Warm. 1868 = Euploca lagoensis

Tournefortia humilis L. 1753 = Euploca humilis

Tournefortia incana (G. Mey.) G. Don 1838 = Euploca humilis

Tournefortia meyeri DC. 1845 = Euploca humilis