Paper Boraginaceae

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Computer-generated keys to the flora of Egypt. 6.

The Boraginaceae
A. El-Gazzar*1, A. El-Ghamery2, A. El-Saied2, A.H. Khattab3, A.A. El-Kady2
1
Botany Department, Faculty of Science, Suez Canal University (El-Arish Branch), N. Sinai, Egypt
2
Department of Botany and Microbiology, Faculty of Science, Al-Azhar University, Cairo, Egypt
3
The Herbarium, Botany Department, Faculty of Science, Cairo University, Cairo
*Corresponding author: elgazzar_adel@hotmail.com
Abstract:
Manually-constructed keys to many groups of the Egyptian flora are in urgent need of improvement
and updating. To construct a conventional substitute of the key to representatives of the Boraginaceae, a data
matrix was compiled to accommodate 54 characters recorded comparatively for the 49 species belonging to
14 genera which represent this family in the flora of Egypt. The 54 characters were accurately and lucidly
defined to cover as much of the easily observable aspects of vegetative and floral variation in the plants as
possible. The data matrix was analyzed using the key-generating package of programs DELTA. The analysis
produced a conventional key with a detailed description of every species in terms of the 54 characters. The
key is decidedly a marked improvement over its predecessors in that it is strictly comparative and leads
directly to the full scientific name of any taxon, instead of having to use a key to the genera followed by a
second key to the infra-generic taxa.
Key words: Boraginaceae, conventional key, DELTA, Egypt, flora, identification
INTRODUCTION
The Boraginaceae Jussieu comprises 110 genera and 1595 species (APG III, 2014)[Angiosperm
Phylogeny Group], but the numbers reach 130-135 genera with 2400-2600 species in Bergianska website
(2014). The plants are perennial, biennial, or annual herbs, less often lianas, shrubs, or trees, usually bristly
or pubescent and scabrid. Leaves are simple, alternate, rarely opposite, with serrate or entire margin.
Inflorescences often scorpioid cymes, sometimes solitary; bracts present or absent. Flowers are bisexual,
actinomorphic or, rarely, slightly zygomorphic. Calyx is usually 5-parted or lobed, mostly persistent. Corolla
tubular, funnel-shaped, campanulate or rotate; tube with 5 appendages, rarely more, mostly trapeziform,
rarely absent, a ring of hairs present sometimes; limb usually 5-parted; lobes overlapping, rarely twisted in
bud. Stamens 5, inserted on corolla tube, included or rarely exserted; anthers introrse, 2-loculed, usually
dorsifixed at base, less usually medifixed, dehiscence longitudinal. Nectaries present on disc below ovary or
at base of corolla tube. Ovary is superior, 2-carpels; locules 2 and each with 2 ovules, or divided by
secondary septae into 4 locules each with 1 ovule; ovules nearly anatropous, semi- anatropous, or atropous.
Style gynobasic or, rarely, terminal, branched or not. Fruit 1–4-seeded drupe or 4 nutlets (mericarps); nutlets
mostly dry, often ornamented with wings, prickles and/or glochids (stiff bristles with barbed or anchor-like
tips). Seeds vertical or oblique, with basal attachment; endosperm oily or absent; embryo straight, less often
curved; cotyledons flat, fleshy; coat membranous (Barroso 1986; Al-Shehbaz, 1991; Watson and Dallwitz
1992 onwards; APG III, 2014).
The map shown in the APG III (2014) account of the Boraginaceae indicates that members of this
family are widely distributed in the world. They cover the whole of N. America, Europe, Asia (except most
of India and the Indonesian Archipelago), Australia, New Zealand, North Africa, East Africa, Madagascar,
the Cape Province, and western South America. Detailed accounts of the geographical distribution of the
Boraginaceae are given by Al-Shehbaz (1991) and Thorne (1992). According to Retief and Vanwyk, (1997)
members of this family grow mainly in dry, cliffy and sunny habitats.
Anatomical studies on members of the Boraginaceae are numerous and only a representative selection
of the more recent studies is presented here. Selvi and Bigazzi (2001) studied leaf surface and anatomy in
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tribe Boragineae. Foliar anatomy of Heliotropium was also the subject of study by Abbasi et al. (2011),
Ahmed and Kordofani (2012), and Alwahibi and Bukhari (2013). Similarly, foliar anatomy was studied by
Akçin and Baki (2007) in three Symphytum species, by Akçin et al. (2012) in four Cynoglossum species, and
by Güven et al. (2013) in six Onosma species. Stomatal profile in the foliar epidermis was studied by Dasti et
al. (2003) in 31 species belonging to 15 genera, while foliar trichomes were extensively surveyed by Al-
Nowaihi et al. (1987), Selvi and Bigazzi (2001), Diane et al. (2003), Taia (2006), Ventrella and Marinho
(2008), Perveen (2009), and Mehrabian et al. (2014). Ovchinnikova (2009) used details of nutlet surface
sculpture to determine the position of tribe Eritrichieae in the Boraginaceae, while Keshavarzi et al. (2013)
used 23 characters of stem and fruit anatomy to evaluate the relationships of four Anchusa species. Nodal
anatomy is rarely studied in angiosperms and it seems that the only study concerning members of the
Boraginaceae is that of Trivedi et al. (1976) on certain representatives of the family in India. According to
the account of Boraginaceae in the Bergianska website (2014), the nodes are usually unilacunar with one or
three leaf traces, or trilacunar with three leaf traces. Using scanning electron microscopy, Rabaey et al.
(2010) were able to elucidate the phylogenetic significance of the distribution of bordered pits in the
secondary xylem of 105 species representing the major groups of the Boraginaceae.
Pollen morphology of the Boraginaceae was studied extensively. Thus, Clarke (1977) maintained that
the Boraginaceae is one of the more eurypalynous families. Diez and Valdes (1991) studied the pollen
morphology of 33 species of Boraginaceae from the Iberian Peninsula belonging to the tribes Cynoglosseae
and Eritrichieae and they confirm the eurypalinous character of this family. Scheel et al. (1996) studied the
pollen morphology of 30 taxa and classified them into nine pollen types based on aperture characteristics and
surface ornamentation. Other palynological studies of different members of the Boraginaceae from various
parts of the world include those by Nowicke and Miller (1990), El Ghazaly (1995), Qureshi (1997), Bigazzi
and Selvi (1998), Liu Jiaxi et al. (2001a and b), Khatamsaz (2001), Hargrove and Simpson (2003), Bigazzi et
al. (2006), Melo et al. (2006), Binzet et al. (2010), Binzet (2011), Falatoury et al. (2011), Mehrabian et al.
(2012), Coutinho et al. (2012) and Fukuda and Ikeda (2012).
Members of the Boraginaceae are of little economic importance. Only a few species are used in
traditional medicine for treating wounds, fever, chest pain, and skin diseases (Neuwinger, 2000). Some
species are grown as ornamentals, sources of timber or dye producers. The family is generally regarded as
closely related to the Hydrophyllaceae (with similar coiled inflorescence), and the Lamiaceae (with
gynobasic style); Watson and Dallwitz (1992 onwards), APG III (2014).
The present study has been undertaken to benefit from the inherent facilities of the program package
DELTA in producing much improved keys for the identification of the genera and species representing the
Boraginaceae in the flora of Egypt. Reference to previous applications of this package to other groups in the
Egyptian flora can be found in El-Gazzar et al. (2013).
MATERIAL AND METHODS

The Boraginaceae is represented in the flora of Egypt by 19 genera and 58 species (Boulos 2002) but it
was possible to collect herbarium specimens of only 49 species representing 14 genera of this family from
the two major herbaria in Egypt, the herbarium of Botany Department, Faculty of science, Cairo
University (CAI) and the herbarium of Flora and Phytotaxonomy Researches Unit at the Agricultural
Museum, Ministry of Agriculture, Dokki, Giza (CAIM); acronyms are according to the Index
Herbariorum (Holmgren et al.1990). The number of specimens representing each taxon ranged between one
and eight. The identity of available specimens was verified by re-identifying them with the aid of the local
flora (Boulos, 2002) and the floras of neighboring countries (e.g. Andrews 1956; Feinbrun-Dothan, 1977 and
1978). Nomenclature was updated from the two websites (http://www.theplantlist.org/), and
(http://www.tropicos.org), where full lists of synonyms and author citations can be found. Full names with
author citations of taxa and collection data of most of the specimens are given in Appendix 1.
As many aspects of variation in vegetative and floral morphology as can be found in the available
specimens were recorded comparatively in a data matrix. The wide range of anatomical, cytological,
palynological and chemical variation in members of the this family was deliberately avoided so that only the
easily observable features were recorded. For uniformity in the usage of the descriptive terminology to define
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the characters and their states, reference was made to the standard comprehensive dictionary compiled by
Stearn (1966).
The data matrix was subjected to analysis under the program suit DELTA which is a multi-purpose
format for generating conventional (i.e. printable) and interactive (i.e. online) identification keys (Dallwitz et
al. 1993 onwards, Dallwitz and Paine 2005, Dallwitz 2010; http://delta-intkey.com; www.sourceforge.net).
Being essentially a format for translating taxonomic data, DELTA produces descriptions of taxa in natural
language (detailed descriptions) and in serial numbers of characters and character-states (item descriptions).
RESULTS
The Characters
Definition of the 54 characters and their character-states which were recorded comparatively for the 49
species belonging to 14 genera representing the Boraginaceae in the flora of Egypt are listed in Table 1. The
numbers of qualitative, multistate and quantitative characters in that list are 49, 2 and 3, respectively.
Table 1. List of the 54 characters and character-states recorded comparatively for 14 genera and 49 species
representing the Boraginaceae in the flora of Egypt and used to construct a conventional key to them.
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#1. Plant/1. herb/2. woody shrub/
#2. Stem/1. glabrous/2. hairy/
#3. Stem/1. erect/2. prostrate/
#4. Stem branching/1. from base/2. above the base/
#5. Internode length/cm/ [Quantitative character].
#6. Basal leaves/1. alternate/2. opposite/
#7. Leaves/1. pedicelled/2. sessile to sub-sessile/
#8. Leaves blade/1. simple/2. pinnatipartite/
#9. Leaves/1. lanceolate/2. linear/3. oblong/4. ovate/ [Multistate character].
#10. Leaf surface/1. wrinkled/2. smooth/
#11. Base of leaf blade/1. symmetric/2. asymmetric/
#12. Leaf veins/1. prominent/2. not prominent/
#13. Leaf margin/1. entire/2. serrate/3. undulate/ [Multistate character].
#14. Leaf apex/1. acute/2. obtuse/
#15. Simple hairs on leaves/1. present/2. absent/
#16. Glandular hairs on leaves/1. present/2. absent/
#17. Bulbs hairs on leaves/1. present/2. absent/
#18. Hispid hairs on leaves/1. present/2. absent/
#19. Woolly hairs on leaves/1. present/2. absent/
#20. Bracteoles/1. present/2. absent/
#21. Bracteoles/1. enclosing calyx/2. not enclosing calyx/
#22. Inflorescence/1. raceme/2. circinnate/
#23. Inflorescence/1. leafy/2. leafless/
#24. Number of flowers/bract/1. less than 8/2. more than 10/
#25. Flower/1. pedicelled/2. sessile to sub-sessile/
#26. Pedicle/1. much longer than calyx/2. as long as calyx or shorter/
#27. Sepal fusion/1. more than half the length/2. Less than half the length/
#28. Apex of calyx lobes/1. acute/2. filiform/
#29. Simple hairs on sepals/1. present/2. absent/
#30. Glandular hairs on sepals/1. present/2. absent/
#31. Hispid hairs on sepals/1. present/2. absent/
#32. Woolly hair on sepals/1. present/2. absent/
#33. Sepal length/cm/ [Quantitative character].
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#34. Petal color/1. blue, purple or pink/2. Yellow or white/

#35. Petal outer surface/1. hairy/2. glabrous/
#36. Petal length/cm/ [Quantitative character].
#37. Petal apex/1. acute/2. obtuse/
#38. Petal fusion/1. more than half the length/2. Less than half the length/
#39. Petal lobes/1. equal/2. unequal/
#40. Corolla throat/1. with scales (5 fornices; Fig. 1) /2. without scales/
#41. No. of stamens/1. 2/2. 5/
#42. Anthers/1. exerted/2. included/
#43. Staminal filaments/1. hairy/2. glabrous/3. reduced/
#44. Appendix on anther/1. present/2. absent/ [Fig. 4]
#45. Anthers/1. sagittate/2. not sagittate/ [Fig. 4]
#46. Style/1. hairy/2. glabrous/
#47. Style/1. terminal (Fig. 3)/2. gynobasic (Fig. 2)/
#48. Style/1. inserted/2. exerted/
#49. Style/1. Bifid (Fig. 2)/2. Undivided (Figs. 1 & 3)/
#50. Stigma/1. Conical (Fig. 3)/2. capitate-globose (Figs.1 & 2)/
#51. Stigma/1. as long as style or shorter/2. much longer than style/
#52. Ovary/1. hairy/2. glabrous/
#53. Nectar disc/1. present/2. absent/ (Fig. 3)
#54. Gynophore/1. present/2. absent/ (Figs.2 & 3).
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Figs. 1 – 4. Variation in floral morphology of Boraginaceae.
Figure 1. Anchusa undulata ssp. hybrida.

Simple style with (black) capitate stigma and 5 (brown) appendages at corolla throat and opposite to corolla
blue lobes.
Figure 2. Arnebia hispidissima.

Ovary (black) deeply divided; style gynobasic, bifid with two capitate stigmas.
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Figure 3. Heliotropium strigosum.

Ovary syncarpous, on nectar disc; style (black) simple, with one black conical (arrow-head) stigma.
Figure 4. Trichodesma africanum.

Five dark brown anthers with cordate base and long tapering apices.
The Key
Application of the program suite DELTA to the data recorded for the 14 genera and 49 species of
Boraginaceae led to the construction of the following conventional key.
Characters: 54 indata, 51 included, 34 in key.
Items: 49 indata, 49 included, 49 in key.
Parameters: Rbase =1.40 Abase = 2.00 Reuse = 1.01 Varywt = 0.80
Characters included: 1–4 6–32 34–35 37–54
Character reliabilities: 1–54,5
1. Leaves lanceolate.................................................................................................................................... 2
Leaves linear......................................................................................................................................... 13
Leaves oblong........................................................................................................................................27
Leaves ovate......................................................................................................................................... 33
2(1). Staminal filaments hairy......................................................................................................................... 3

Staminal filaments glabrous.................................................................................................................... 4
Staminal filaments reduced..................................................................................................................... 7
3(2). Leaf apex acute; hispid hairs on leaves present; woolly

hairs on leaves absent; bracteoles present................................................................ Nonea vivianii DC.
Leaf apex obtuse; hispid hairs on leaves absent; woolly hairs on leaves
present; bracteoles absent.................................................. Paracaryum intermedium (Fresen.) Lipsky
4(2). Leaf margin entire...................................................................................................................................5

Leaf margin serrate........................................................................................... Anchusa milleri Spreng.
Leaf margin undulate............................................................................................................................. 6
5(4). Leaf apex acute; stem branching from base; glandular hairs on leaves present; inflorescence
circinnate......................................................................... Echium sabulicolum Pomel var sabulicolum
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Leaf apex obtuse; stem branching above the base; glandular hairs on
leaves absent; inflorescence raceme........................................... Anchusa humilis (Desf.) I.M. Johnst.
6(4). Stem erect; simple hairs on leaves absent; glandular hairs on leaves absent; bulbs
hairs on leaves Present........................... Anchusa undulata L. ssp hybrida(Ten.) Beg. In Beg & Furi
Stem prostrate; simple hairs on leaves present; glandular hairs on
leaves present; bulbs hairs on leaves absent.................................................... Anchusa hispida Forssk.
7(2). Leaf margin entire................................................................................................................................... 8

Leaf margin serrate............................................................................................................................... 12
Leaf margin undulate..................... Heliotropium bacciferum Forssk ssp bacciferum var bacciferum
8(7). Basal leaves alternate; leaf surface smooth; simple hairs on leaves present;
bracteoles not enclosing calyx................................................................................................................ 9
Basal leaves opposite; leaf surface wrinkled; simple hairs on leaves absent;
bracteoles enclosing calyx.................................................................................................................... 10
9(8). Leaf apex acute; plant woody shrub; leaves pedicelled;

leaf veins prominent.......................................................... Heliotropium zeylanicum (Burm. F.) Lam.
Leaf apex obtuse; plant herb; leaves sessile to sub-sessile; leaf
veins not prominent............................................................. Lappula sinaica (DC.) Asch. & Schweinf.
10(8). Glandular hairs on leaves present; bulbs hairs on leaves absent; number
of flowers/bract more than 10................................ Trichodesma africanum (L.) R.Br. var africanum
Glandular hairs on leaves absent; bulbs hairs on leaves present; number
of flowers/bract less than 8....................................................................................................................11
11(10). Style hairy; style terminal; ovary hairy; nectar disc

present.............................. Trichodesma africanum (L.) R.Br. Var heterotrichum Bornm. & Kneuk.
Style glabrous; style gynobasic; ovary glabrous; nectar disc
Absent....................................... Trichodesma africanum (L.) R.Br. Var abyssinicum Brand in Engl.
12(7). Leaf apex acute; leaves sessile to sub-sessile; simple hairs on leaves present;
bulbs hairs on leaves absent...................................................................... Anchusa aegyptiaca (L.) DC.
Leaf apex obtuse; leaves pedicelled; simple hairs on leaves absent; bulbs
hairs on leaves present...........................Heliotropium pterocarpum (DC.) Hochst. & Steud. ex Bunge
13(1). Staminal filaments hairy....................................................................................................................... 14

Staminal filaments glabrous.................................................................................................................. 15
Staminal filaments reduced................................................................................................................... 24
14(13). Style hairy; stem branching from base; leaves sessile to sub- sessile; hispid
hairs on leaves present........................................................... Moltkiopsis ciliata (Forssk.) I.M. Johnst.
Style glabrous; stem branching above the base; leaves pedicelled; hispid
hairs on leaves absent................. Heliotropium strigosum Willd. var brevifolium (Wal.) C.B. Clarke
15(13). Hispid hairs on leaves present; hispid hairs on sepals present.............................................................16

Hispid hairs on leaves absent; hispid hairs on sepals absent................................................................ 20
16(15). Leaf apex acute.................................................................................................................................... 17

Leaf apex obtuse................................................................................................................................... 18
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17(16). Style hairy; leaf veins prominent; apex of calyx lobes filiform; simple hairs
on sepals present.................................... Arnebia decumbens (Vent.) Coss. & Kralik var decumbens
Style glabrous; leaf veins not prominent; apex of calyx lobes acute; simple
hairs on sepals absent...................................................................... Arnebia hispidissima (Lehm.) DC.
18(16). Style hairy; leaf veins prominent; bulbs hairs on leaves

present; petal fusion more than half the length .................................................................................. 19
Style glabrous; leaf veins not prominent; bulbs hairs on leaves absent; petal
fusion less than half thelength....................................... Buglossoides incrassata (Guss.) I.M. Johnst.
19(18). Inflorescence raceme; number of flowers/bract more than 10; glandular

hairs on sepals present; petal color yellow or white....................................... Arnebia linearifolia DC.
Inflorescence circinnate; number of flowers/bract less than 8; glandular
hairs on sepals absent; petal color blue, purple or pink................................... Echium rauwolfii Delile
20(15). Leaf apex

acute.......................................................................................................................................................21
Leaf apex obtuse................................................................................................................................... 22
21(20). Stem erect; style glabrous; plant woody shrub;

leaf veins not prominent................................................................... Alkanna strigosa Boiss. & Hohen.
Stem prostrate; style hairy; plant herb; leaf veins
Prominent.......................................................Echium angustifolium Mill. Ssp sericeum (Vahl) Klotz
22(20). Style hairy; stem branching from base; bulbs hairs on leaves present;
bracteoles enclosing calyx.................................................................................................................... 23
Style glabrous; stem branching above the base; bulbs hairs on leaves absent;
bracteoles not enclosing calyx ............................................ Buglossoides tenuiflora (L.f.) I.M.Johnst.
23(22). Plant herb; leaf veins prominent; simple hairs on leaves absent;
inflorescence circinnate............................................... Echium angustifolium Mill. Ssp angustifolium
Plant woody shrub; leaf veins not prominent; simple hairs on leaves
present; inflorescence raceme.................................................................. Echiochilon fruticosum Desf.
24(13). Style hairy; simple hairs on leaves absent........................................................................................... 25

Style glabrous; simple hairs on leaves present...................................................................................... 26
25(24). Plant herb; stem branching above the base; leaf veins prominent; inflorescence
leafless.........................Arnebia decumbens (Vent.) Coss. & Kralik var macrocalyx Coss. & Kralik
Plant woody shrub; stem branching from base; leaf veins not prominent;
inflorescence leafy..........................Lappula spinocarpos (Forssk.) Asch. ex Kuntze var spinocarpos
26(24). Stem erect; plant woody shrub; stem branching from base; glandular hairs on leaves
absent........................................Lappula spinocarpos (Forssk.) Asch. ex Kuntze var inermis Botsch.
Stem prostrate; plant herb; stem branching above the base;
glandular hairs on leaves present.................................................................. Alkanna tinctoria Tausch.
27(1). Staminal filaments hairy........................................................................................................................28

Staminal filaments glabrous .................................................................................................................29
Staminal filaments reduced .................................................................................................................32
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28(27). Leaf margin entire; leaf apex obtuse; leaves sessile to sub-sessile; bulbs
hairs on leaves absent........................................................................................ Echium horridum Batt.
Leaf margin serrate; leaf apex acute; leaves pedicelled; bulbs hairs on
leaves present........................................................ Heliotropium ramosissimum (Lehm.) Sieb. ex DC.
29(27). Stem erect; leaf veins not prominent; glandular hairs on leaves present;
bulbs hairs on leaves absent.................................................................................................................. 30
Stem prostrate; leaf veins prominent; glandular hairs on leaves absent; bulbs
hairs on leaves present...........................................................................................................................31
30(29). Basal leaves alternate; leaf apex acute; style glabrous;

stem branching above the base.........................................................................Asperugo procumbens L.
Basal leaves opposite; leaf apex obtuse; style hairy;
stem branching from base............................................................................... Arnebia tinctoria Forssk.
31(29). Leaf margin entire; leaf apex obtuse; style hairy;

stem branching from base................................................................................. Echium rubrum Forssk.
Leaf margin serrate; leaf apex acute; style glabrous;
stem branching above the base........................................................................ Coldenia procumbens L.
32(27). Plant herb; stem hairy; leaves sessile to sub-sessile;

leaf surface smooth...................................................................... Paracaryum rugulosum (DC.) Boiss.
Plant woody shrub; stem glabrous; leaves pedicelled;
leaf surface wrinkled.............................................................................. Heliotropium curassavicum L.
33(1). Staminal filaments hairy.............................................. Heliotropium hirsutissimum Grauer in Weber

Staminal filaments glabrous.................................................................................................................. 34
Staminal filaments reduced................................................................................................................... 38
34(33). Leaf margin entire; flower pedicelled; style terminal.......................................................................... 35

Leaf margin undulate; flower sessile to sub-sessile; style gynobasic .................................................. 37
35(34). Leaf apex acute; simple hairs on leaves present; bulbs hairs on leaves absent ...................................36
Leaf apex obtuse; simple hairs on leaves absent; bulbs hairs on
leaves present.............................................................. Heliotropium lasiocarpum Fisch. & C.A. Mey.
36(35). Plant herb; stem branching above the base; leaf veins prominent; woolly
hairs on leaves present................................................................................... Heliotropium supinum L.
Plant woody shrub; stem branching from base; leaf veins not prominent;
woolly hairs on leaves absent............................................................ Heliotropium ovalifolium Forssk.
37(34). Leaf apex acute; style glabrous; leaf surface smooth; glandular
hairs on leaves present....................................................................... Heliotropium arabinense Fresen.
Leaf apex obtuse; style hairy; leaf surface wrinkled; glandular
hairs on leaves absent....................................................................... Heliotropium aegyptiacum Lehm.
38(33). Leaf apex acute; bracteoles present; bracteoles enclosing

calyx; petal outer surface glabrous....................................................................................................... 39
Leaf apex obtuse; bracteoles absent; bracteoles not enclosing calyx; petal
outer surface hairy................................................................................................................................ 40
39(38). Basal leaves alternate; leaf margin undulate; style glabrous;

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plant herb..................................................................................................Alkanna orientalis (L.) Boiss.

Basal leaves opposite; leaf marginentire; style hairy;
plant woody shrub..........................................................................Trichodesma ehrenbergii Schweinf.
40(38). Bulbs hairs on leaves present; woolly hairs on leaves absent; number
of flowers/bract more than 10; glandular hairs on
sepals present......................................................... Heliotropium digynum (Forssk.) Asch. Ex C. Chr.
Bulbs hairs on leaves absent; woolly hairs on leaves present; number
of flowers/bract less than 8; glandular hairs on sepals
absent....................Heliotropium bacciferum Forssk. ssp. bacciferum var. erosum Hadidy in Boulos
3.2.2.3 Detailed descriptions
Alkanna tinctoria Tausch.

Plant herb. Stem hairy. Stem prostrate. Stem branching above the base. Internode length 0.7–1.4 cm.
Basal leaves alternate. Leaves sessile to sub-sessile. Leaves blade simple. Leaves linear. Leaf surface smooth.
Base of leaf blade symmetric. Leaf veins not prominent. Leaf margin entire. Leaf apex acute. Simple hairs on
leaves present. Glandular hairs on leaves present. Bulbs hairs on leaves absent. Hispid hairs on leaves absent.
Woolly hairs on leaves absent. Bracteoles present. Bracteoles enclosing calyx. Inflorescence raceme.
Inflorescence leafless. Number of flowers/bract less than 8. Flower pedicelled. Pedicle as long as calyx or
shorter. Sepal fusion Less than half the length. Apex of calyx lobes acute. Simple hairs on sepals present.
Glandular hairs on sepals present. Hispid hairs on sepals absent. Woolly hair on sepals absent. Sepal length
0.3–0.6 cm. Petal color blue, purple or pink. Petal outer surface glabrous. Petal length 0.6–0.9 cm. Petal apex
obtuse. Petal fusion Less than half the length. Petal lobes equal. Corolla throat covered with scales. No. of
stamens 5. Anthers included. Staminal filaments reduced. Appendix on anther absent. Anthers not sagittate.
Style glabrous. Style gynobasic. Style inserted. Style undivided. Stigma capitate-globose. Stigma as long as
style or shorter. Ovary glabrous. Nectar disc absent. Gynophore absent.
Alkanna orientalis (L.) Boiss.

Plant herb. Stem hairy. Stem erect. Stem branching above the base. Internode length 0.7–1.5 cm. Basal
leaves alternate. Leaves sessile to sub-sessile. Leaves blade simple. Leaves ovate. Leaf surface smooth. Base
of leaf blade symmetric. Leaf veins not prominent. Leaf margin undulate. Leaf apex acute. Simple hairs on
leaves present. Glandular hairs on leaves present. Bulbs hairs on leaves absent. Hispid hairs on leaves absent.
Inflorescence leafless. Number of flowers/bract more than 10. Flower pedicelled. Pedicle as long as calyx or
Glandular hairs on sepals present. Hispid hairs on sepals absent. Woolly hair on sepals absent. Sepal length
0.5–1.5 cm. Petal color Yellow or white. Petal outer surface glabrous. Petal length 1.1–1.6 cm. Petal apex
obtuse. Petal fusion more than half the length. Petal lobes equal. Corolla throat covered with scales. No. of
Alkanna strigosa Boiss. & Hohen.

Plant woody shrub. Stem hairy. Stem erect. Stem branching from base. Internode length 0.1–0.5 cm.
leaves present. Glandular hairs on leaves absent. Bulbs hairs on leaves absent. Hispid hairs on leaves absent.
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Inflorescence leafy. Number of flowers/bract less than 8. Flower pedicelled. Pedicle as long as calyx or
shorter. Sepal fusion more than half the length. Apex of calyx lobes acute. Simple hairs on sepals present.
Glandular hairs on sepals absent. Hispid hairs on sepals absent. Woolly hair on sepals absent. Sepal length
0.5–1 cm. Petal color blue, purple or pink. Petal outer surface glabrous. Petal length 1–1.5 cm. Petal apex
obtuse. Petal fusion more than half the length. Petal lobes equal. Corolla throat covered with scales. No. of
stamens 5. Anthers included. Staminal filaments glabrous. Appendix on anther absent. Anthers not sagittate.
Anchusa aegyptiaca (L.) DC.

Plant herb. Stem hairy. Stem erect. Stem branching above the base. Internode length 0.5–4 cm. Basal
leaves alternate. Leaves sessile to sub-sessile. Leaves blade simple. Leaves lanceolate. Leaf surface smooth.
Base of leaf blade symmetric. Leaf veins not prominent. Leaf margin serrate. Leaf apex acute. Simple hairs
on leaves present. Glandular hairs on leaves absent. Bulbs hairs on leaves absent. Hispid hairs on leaves
present. Woolly hairs on leaves absent. Bracteoles present. Bracteoles enclosing calyx. Inflorescence raceme.
Glandular hairs on sepals absent. Hispid hairs on sepals present. Woolly hair on sepals absent. Sepal length
obtuse. Petal fusion Less than half the length. Petal lobes equal. Corolla throat without scales. No. of stamens
5. Anthers included. Staminal filaments reduced. Appendix on anther absent. Anthers not sagittate. Style
glabrous. Style gynobasic. Style inserted. Style undivided. Stigma capitate-globose. Stigma as long as style
or shorter. Ovary glabrous. Nectar disc absent. Gynophore absent.
Anchusa hispida Forssk.

Plant herb. Stem hairy. Stem prostrate. Stem branching above the base. Internode length 1–2.5 cm. Basal
Base of leaf blade symmetric. Leaf veins not prominent. Leaf margin undulate. Leaf apex obtuse. Simple
hairs on leaves present. Glandular hairs on leaves present. Bulbs hairs on leaves absent. Hispid hairs on
leaves present. Woolly hairs on leaves absent. Bracteoles present. Bracteoles enclosing calyx. Inflorescence
raceme. Inflorescence leafless. Number of flowers/bract more than 10. Flower pedicelled. Pedicle as long as
calyx or shorter. Sepal fusion Less than half the length. Apex of calyx lobes acute. Simple hairs on sepals
present. Glandular hairs on sepals absent. Hispid hairs on sepals present. Woolly hair on sepals absent. Sepal
length 0.1–0.4 cm. Petal color blue, purple or pink. Petal outer surface glabrous. Petal length 0.1–0.3 cm.
Petal apex obtuse. Petal fusion Less than half the length. Petal lobes unequal. Corolla throat without scales.
No. of stamens 5. Anthers included. Staminal filaments glabrous. Appendix on anther absent. Anthers not
sagittate. Style glabrous. Style gynobasic. Style inserted. Style undivided. Stigma capitate-globose. Stigma as
long as style or shorter. Ovary glabrous. Nectar disc absent. Gynophore absent.
Anchusa humilis (Desf.) I.M. Johnst.

Plant herb. Stem hairy. Stem prostrate. Stem branching above the base. Internode length 2–5 cm. Basal
Base of leaf blade symmetric. Leaf veins not prominent. Leaf margin entire. Leaf apex obtuse. Simple hairs
on leaves absent. Glandular hairs on leaves absent. Bulbs hairs on leaves absent. Hispid hairs on leaves
present. Woolly hairs on leaves absent. Bracteoles present. Bracteoles enclosing calyx. Inflorescence raceme.
Inflorescence leafless. Number of flowers/bract more than 10. Flower sessile to sub-sessile. Pedicle as long
as calyx or shorter. Sepal fusion Less than half the length. Apex of calyx lobes acute. Simple hairs on sepals
11
Petal apex obtuse. Petal fusion Less than half the length. Petal lobes equal. Corolla throat without scales. No.
of stamens 5. Anthers included. Staminal filaments glabrous. Appendix on anther absent. Anthers not
sagittate. Style glabrous. Style gynobasic. Style inserted. Style undivided. Stigma capitate-globose. Stigma as
long as style or shorter. Ovary glabrous. Nectar disc absent. Gynophore absent.
Anchusa milleri Spreng.

Plant herb. Stem hairy. Stem prostrate. Stem branching above the base. Internode length 0.1–0.5 cm.
Basal leaves alternate. Leaves sessile to sub-sessile. Leaves blade simple. Leaves lanceolate. Leaf surface
smooth. Base of leaf blade symmetric. Leaf veins not prominent. Leaf margin serrate. Leaf apex obtuse.
Simple hairs on leaves absent. Glandular hairs on leaves absent. Bulbs hairs on leaves absent. Hispid hairs on
leaves present. Woolly hairs on leaves absent. Bracteoles present. Bracteoles enclosing calyx. Inflorescence
raceme. Inflorescence leafless. Number of flowers/bract less than 8. Flower pedicelled. Pedicle as long as
absent. Glandular hairs on sepals absent. Hispid hairs on sepals present. Woolly hair on sepals absent. Sepal
length 0.3–0.6 cm. Petal color Yellow or white. Petal outer surface glabrous. Petal length 0.8–1.1 cm. Petal
apex obtuse. Petal fusion more than half the length. Petal lobes unequal. Corolla throat without scales. No. of
Anchusa undulata L. ssp. hybrida (Ten.) Beg. in Beg & Furi

Base of leaf blade symmetric. Leaf veins not prominent. Leaf margin undulate. Leaf apex obtuse. Simple
hairs on leaves absent. Glandular hairs on leaves absent. Bulbs hairs on leaves present. Hispid hairs on leaves
absent. Woolly hairs on leaves absent. Bracteoles present. Bracteoles not enclosing calyx. Inflorescence
raceme. Inflorescence leafless. Number of flowers/bract more than 10. Flower pedicelled. Pedicle as long as
calyx or shorter. Sepal fusion more than half the length. Apex of calyx lobes acute. Simple hairs on sepals
absent. Glandular hairs on sepals present. Hispid hairs on sepals present. Woolly hair on sepals absent. Sepal
Petal apex obtuse. Petal fusion more than half the length. Petal lobes equal. Corolla throat covered with
scales. No. of stamens 5. Anthers included. Staminal filaments glabrous. Appendix on anther absent. Anthers
not sagittate. Style hairy. Style gynobasic. Style inserted. Style undivided. Stigma capitate-globose. Stigma
as long as style or shorter. Ovary glabrous. Nectar disc absent. Gynophore absent.
Arnebia decumbens (Vent.) Coss. & Kralik var. decumbens

Plant herb. Stem hairy. Stem erect. Stem branching above the base. Internode length 1–2 cm. Basal leaves
alternate. Leaves sessile to sub-sessile. Leaves blade simple. Leaves linear. Leaf surface smooth. Base of leaf
blade symmetric. Leaf veins prominent. Leaf margin entire. Leaf apex acute. Simple hairs on leaves absent.
Glandular hairs on leaves absent. Bulbs hairs on leaves present. Hispid hairs on leaves present. Woolly hairs
on leaves absent. Bracteoles present. Bracteoles enclosing calyx. Inflorescence raceme. Inflorescence
leafless. Number of flowers/bract more than 10. Flower pedicelled. Pedicle as long as calyx or shorter. Sepal
fusion Less than half the length. Apex of calyx lobes filiform. Simple hairs on sepals present. Glandular hairs
on sepals absent. Hispid hairs on sepals present. Woolly hair on sepals absent. Sepal length 0.5–1.2 cm. Petal
color Yellow or white. Petal outer surface hairy. Petal length 0.5–1.4 cm. Petal apex obtuse. Petal fusion
more than half the length. Petal lobes equal. Corolla throat without scales. No. of stamens 5. Anthers
included. Staminal filaments glabrous. Appendix on anther absent. Anthers not sagittate. Style hairy. Style
gynobasic. Style inserted. Style undivided. Stigma capitate-globose. Stigma as long as style or shorter. Ovary
glabrous. Nectar disc absent. Gynophore absent.
12
Arnebia decumbens (Vent.) Coss. & Kralik var. macrocalyx Coss. & Kralik
Plant herb. Stem hairy. Stem erect. Stem branching above the base. Internode length 1–3.5 cm. Basal
leaves alternate. Leaves sessile to sub-sessile. Leaves blade simple. Leaves linear. Leaf surface smooth. Base
of leaf blade symmetric. Leaf veins prominent. Leaf margin entire. Leaf apex acute. Simple hairs on leaves
absent. Glandular hairs on leaves present. Bulbs hairs on leaves absent. Hispid hairs on leaves present.
shorter. Sepal fusion Less than half the length. Apex of calyx lobes acute. Simple hairs on sepals absent.
0.5–1.3 cm. Petal color Yellow or white. Petal outer surface hairy. Petal length 0.5–1.5 cm. Petal apex
obtuse. Petal fusion more than half the length. Petal lobes equal. Corolla throat without scales. No. of
Style hairy. Style gynobasic. Style inserted. Style undivided. Stigma capitate-globose. Stigma as long as style
or shorter. Ovary hairy. Nectar disc absent. Gynophore absent.
Arnebia hispidissima (Lehm.) DC.

of leaf blade symmetric. Leaf veins not prominent. Leaf margin entire. Leaf apex acute. Simple hairs on
leaves absent. Glandular hairs on leaves absent. Bulbs hairs on leaves present. Hispid hairs on leaves present.
0.4–0.7 cm. Petal color Yellow or white. Petal outer surface glabrous. Petal length 0.5–1 cm. Petal apex
Style glabrous. Style gynobasic. Style inserted. Style bifid. Stigma capitate-globose. Stigma as long as style
Arnebia linearifolia DC.

Plant herb. Stem hairy. Stem erect. Stem branching above the base. Internode length 1–3.5 cm. Basal
of leaf blade symmetric. Leaf veins prominent. Leaf margin entire. Leaf apex obtuse. Simple hairs on leaves
absent. Glandular hairs on leaves absent. Bulbs hairs on leaves present. Hispid hairs on leaves present.
Glandular hairs on sepals present. Hispid hairs on sepals present. Woolly hair on sepals absent. Sepal length
Style hairy. Style gynobasic. Style inserted. Style bifid. Stigma capitate-globose. Stigma as long as style or
shorter. Ovary glabrous. Nectar disc absent. Gynophore absent.
Arnebia tinctoria Forssk.

Plant herb. Stem hairy. Stem erect. Stem branching from base. Internode length 2–3.5 cm. Basal leaves
opposite. Leaves pedicelled. Leaves blade simple. Leaves oblong. Leaf surface smooth. Base of leaf blade
symmetric. Leaf veins not prominent. Leaf margin entire. Leaf apex obtuse. Simple hairs on leaves absent.
13
Glandular hairs on leaves present. Bulbs hairs on leaves absent. Hispid hairs on leaves absent. Woolly hairs
on leaves absent. Bracteoles present. Bracteoles enclosing calyx. Inflorescence raceme. Inflorescence
leafless. Number of flowers/bract less than 8. Flower pedicelled. Pedicle as long as calyx or shorter. Sepal
fusion more than half the length. Apex of calyx lobes acute. Simple hairs on sepals absent. Glandular hairs on
sepals present. Hispid hairs on sepals present. Woolly hair on sepals absent. Sepal length 0.5–1 cm. Petal
color blue, purple or pink. Petal outer surface glabrous. Petal length 0.5–1.2 cm. Petal apex obtuse. Petal
fusion more than half the length. Petal lobes equal. Corolla throat without scales. No. of stamens 5. Anthers
included. Staminal filaments glabrous. Appendix on anther absent. Anthers not sagittate. Style hairy. Style
gynobasic. Style inserted. Style bifid. Stigma capitate-globose. Stigma as long as style or shorter. Ovary
Asperugo procumbens L.
alternate. Leaves sessile to sub-sessile. Leaves blade simple. Leaves oblong. Leaf surface smooth. Base of
leaf blade symmetric. Leaf veins not prominent. Leaf margin entire. Leaf apex acute. Simple hairs on leaves
absent. Glandular hairs on leaves present. Bulbs hairs on leaves absent. Hispid hairs on leaves present.
Woolly hairs on leaves absent. Bracteoles present. Bracteoles not enclosing calyx. Inflorescence raceme.
Inflorescence leafless. Number of flowers/bract less than 8. Flower pedicelled. Pedicle much longer than
calyx. Sepal fusion Less than half the length. Apex of calyx lobes acute. Simple hairs on sepals absent.
Buglossoides incrassata (Guss.) I.M. Johnst.

of leaf blade symmetric. Leaf veins not prominent. Leaf margin entire. Leaf apex obtuse. Simple hairs on
leaves absent. Glandular hairs on leaves absent. Bulbs hairs on leaves absent. Hispid hairs on leaves present.
0.2–0.5 cm. Petal color blue, purple or pink. Petal outer surface hairy. Petal length 0.3–0.5 cm. Petal apex
obtuse. Petal fusion Less than half the length. Petal lobes equal. Corolla throat without scales. No. of stamens
5. Anthers included. Staminal filaments glabrous. Appendix on anther absent. Anthers not sagittate. Style
Buglossoides tenuiflora (L.f.) I.M. Johnst.

present. Glandular hairs on leaves absent. Bulbs hairs on leaves absent. Hispid hairs on leaves absent. Woolly
hairs on leaves absent. Bracteoles present. Bracteoles not enclosing calyx. Inflorescence raceme.
14
style or shorter. Ovary hairy. Nectar disc absent. Gynophore absent.
Coldenia procumbens L.
Plant herb. Stem hairy. Stem prostrate. Stem branching above the base. Internode length 0.3–1 cm. Basal
leaves alternate. Leaves pedicelled. Leaves blade pinnatipartite. Leaves oblong. Leaf surface wrinkled. Base
of leaf blade asymmetric. Leaf veins prominent. Leaf margin serrate. Leaf apex acute. Simple hairs on leaves
absent. Glandular hairs on leaves absent. Bulbs hairs on leaves present. Hispid hairs on leaves absent. Woolly
hairs on leaves absent. Bracteoles present. Bracteoles enclosing calyx. Inflorescence raceme. Inflorescence
leafy. Number of flowers/bract less than 8. Flower pedicelled. Pedicle as long as calyx or shorter. Sepal
fusion Less than half the length. Apex of calyx lobes acute. Simple hairs on sepals present. Glandular hairs
on sepals absent. Hispid hairs on sepals absent. Woolly hair on sepals absent. Sepal length 0.1–1.1 cm. Petal
color Yellow or white. Petal outer surface glabrous. Petal length 0.1–0.2 cm. Petal apex acute. Petal fusion
Less than half the length. Petal lobes unequal. Corolla throat without scales. No. of stamens 5. Anthers
included. Staminal filaments glabrous. Appendix on anther absent. Anthers not sagittate. Style glabrous.
Style gynobasic. Style inserted. Style bifid. Stigma capitate-globose. Stigma as long as style or shorter.
Ovary glabrous. Nectar disc absent. Gynophore absent.
Echiochilon fruticosum Desf.

Base of leaf blade symmetric. Leaf veins not prominent. Leaf margin entire. Leaf apex obtuse. Simple hairs
on leaves present. Glandular hairs on leaves absent. Bulbs hairs on leaves present. Hispid hairs on leaves
absent. Woolly hairs on leaves absent. Bracteoles present. Bracteoles enclosing calyx. Inflorescence raceme.
Inflorescence leafy. Number of flowers/bract less than 8. Flower sessile to sub-sessile. Pedicle as long as
present. Glandular hairs on sepals absent. Hispid hairs on sepals absent. Woolly hair on sepals absent. Sepal
length 0.1–0.3 cm. Petal color blue, purple or pink. Petal outer surface hairy. Petal length 0.4–1 cm. Petal
apex obtuse. Petal fusion more than half the length. Petal lobes unequal. Corolla throat without scales. No. of
Style hairy. Style gynobasic. Style inserted. Style undivided. Stigma capitate-globose. Stigma as long as style
Echium angustifolium Mill. ssp. angustifolium

Plant herb. Stem hairy. Stem erect. Stem branching from base. Internode length 1.5–4.5 cm. Basal leaves
alternate. Leaves sessile to sub-sessile. Leaves blade simple. Leaves linear. Leaf surface smooth. Base of leaf
blade symmetric. Leaf veins prominent. Leaf margin entire. Leaf apex obtuse. Simple hairs on leaves absent.
Glandular hairs on leaves absent. Bulbs hairs on leaves present. Hispid hairs on leaves absent. Woolly hairs
on leaves absent. Bracteoles present. Bracteoles enclosing calyx. Inflorescence circinnate. Inflorescence
on sepals absent. Hispid hairs on sepals absent. Woolly hair on sepals absent. Sepal length 0.4–1 cm. Petal
color blue, purple or pink. Petal outer surface hairy. Petal length 1–3.2 cm. Petal apex obtuse. Petal fusion
more than half the length. Petal lobes unequal. Corolla throat without scales. No. of stamens 5. Anthers
exerted. Staminal filaments glabrous. Appendix on anther absent. Anthers not sagittate. Style hairy. Style
15
gynobasic. Style exerted. Style bifid. Stigma capitate-globose. Stigma as long as style or shorter. Ovary
glabrous. Nectar disc present. Gynophore absent.
Echium angustifolium Mill. ssp. sericeum (Vahl) Klotz

Plant herb. Stem hairy. Stem prostrate. Stem branching from base. Internode length 0.2–2.1 cm. Basal
of leaf blade symmetric. Leaf veins prominent. Leaf margin entire. Leaf apex acute. Simple hairs on leaves
absent. Glandular hairs on leaves present. Bulbs hairs on leaves absent. Hispid hairs on leaves absent. Woolly
hairs on leaves absent. Bracteoles present. Bracteoles enclosing calyx. Inflorescence circinnate. Inflorescence
on sepals absent. Hispid hairs on sepals absent. Woolly hair on sepals absent. Sepal length 0.4–0.8 cm. Petal
color blue, purple or pink. Petal outer surface hairy. Petal length 1–2.5 cm. Petal apex obtuse. Petal fusion
more than half the length. Petal lobes unequal. Corolla throat without scales. No. of stamens 5. Anthers
exerted. Staminal filaments glabrous. Appendix on anther absent. Anthers sagittate. Style hairy. Style
gynobasic. Style exerted. Style bifid. Stigma capitate-globose. Stigma as long as style or shorter. Ovary
Echium horridum Batt.

leaf blade symmetric. Leaf veins not prominent. Leaf margin entire. Leaf apex obtuse. Simple hairs on leaves
present. Glandular hairs on leaves absent. Bulbs hairs on leaves absent. Hispid hairs on leaves present.
Woolly hairs on leaves absent. Bracteoles present. Bracteoles enclosing calyx. Inflorescence circinnate.
Inflorescence leafy. Number of flowers/bract more than 10. Flower pedicelled. Pedicle as long as calyx or
0.5–1.4 cm. Petal color blue, purple or pink. Petal outer surface hairy. Petal length 1–2.2 cm. Petal apex
acute. Petal fusion more than half the length. Petal lobes equal. Corolla throat without scales. No. of stamens
5. Anthers included. Staminal filaments hairy. Appendix on anther absent. Anthers not sagittate. Style hairy.
Style gynobasic. Style inserted. Style bifid. Stigma capitate-globose. Stigma as long as style or shorter.
Ovary glabrous. Nectar disc present. Gynophore absent.
Echium rauwolfii Delile

Plant herb. Stem hairy. Stem erect. Stem branching above the base. Internode length 0.4–2 cm. Basal
absent. Glandular hairs on leaves absent. Bulbs hairs on leaves present. Hispid hairs on leaves present.
stamens 5. Anthers exerted. Staminal filaments glabrous. Appendix on anther absent. Anthers not sagittate.
Style hairy. Style gynobasic. Style exerted. Style bifid. Stigma capitate-globose. Stigma as long as style or
16
Echium rubrum Forssk.

Plant herb. Stem hairy. Stem prostrate. Stem branching from base. Internode length 0.6–3.3 cm. Basal
leaves alternate. Leaves sessile to sub-sessile. Leaves blade simple. Leaves oblong. Leaf surface smooth.
Base of leaf blade symmetric. Leaf veins prominent. Leaf margin entire. Leaf apex obtuse. Simple hairs on
leaves absent. Glandular hairs on leaves absent. Bulbs hairs on leaves present. Hispid hairs on leaves absent.
as calyx or shorter. Sepal fusion more than half the length. Apex of calyx lobes acute. Simple hairs on sepals
length 0.5–1.1 cm. Petal color blue, purple or pink. Petal outer surface hairy. Petal length 1–2 cm. Petal apex
obtuse. Petal fusion more than half the length. Petal lobes unequal. Corolla throat without scales. No. of
shorter. Ovary hairy. Nectar disc present. Gynophore absent.
Echium sabulicolum Pomel var. sabulicolum

Plant herb. Stem hairy. Stem prostrate. Stem branching from base. Internode length 1–2.1 cm. Basal
leaves absent. Glandular hairs on leaves present. Bulbs hairs on leaves absent. Hispid hairs on leaves present.
Inflorescence leafless. Number of flowers/bract less than 8. Flower sessile to sub-sessile. Pedicle as long as
length 0.4–1.1 cm. Petal color blue, purple or pink. Petal outer surface hairy. Petal length 1–2.1 cm. Petal
apex obtuse. Petal fusion more than half the length. Petal lobes equal. Corolla throat without scales. No. of
shorter. Ovary glabrous. Nectar disc present. Gynophore absent.
Heliotropium aegyptiacum Lehm.

Plant woody shrub. Stem hairy. Stem erect. Stem branching above the base. Internode length 0.9–2.5 cm.
Basal leaves alternate. Leaves pedicelled. Leaves blade simple. Leaves ovate. Leaf surface wrinkled. Base of
leaf blade symmetric. Leaf veins prominent. Leaf margin undulate. Leaf apex obtuse. Simple hairs on leaves
hairs on leaves absent. Bracteoles absent. Bracteoles not enclosing calyx. Inflorescence circinnate.
present. Glandular hairs on sepals absent. Hispid hairs on sepals absent. Woolly hair on sepals present. Sepal
length 0.2–0.4 cm. Petal color Yellow or white. Petal outer surface hairy. Petal length 0.3–0.5 cm. Petal apex
stamens 5. Anthers included. Staminal filaments glabrous. Appendix on anther absent. Anthers sagittate.
Style hairy. Style gynobasic. Style inserted. Style undivided. Stigma conical. Stigma as long as style or
shorter. Ovary hairy. Nectar disc absent. Gynophore absent.
Heliotropium arabinense Fresen.

Plant woody shrub. Stem hairy. Stem erect. Stem branching above the base. Internode length 0.5–1 cm.
Basal leaves alternate. Leaves pedicelled. Leaves blade simple. Leaves ovate. Leaf surface smooth. Base of
leaf blade symmetric. Leaf veins prominent. Leaf margin undulate. Leaf apex acute. Simple hairs on leaves
17
absent. Glandular hairs on leaves present. Bulbs hairs on leaves absent. Hispid hairs on leaves absent. Woolly
Inflorescence leafless. Number of flowers/bract less than 8. Flower sessile to sub-sessile. Pedicle as long as
absent. Glandular hairs on sepals present. Hispid hairs on sepals absent. Woolly hair on sepals absent. Sepal
Style glabrous. Style gynobasic. Style inserted. Style undivided. Stigma conical. Stigma much longer than
style. Ovary hairy. Nectar disc present. Gynophore absent.
Heliotropium bacciferum Forssk. Ssp. bacciferum var. bacciferum

Basal leaves alternate. Leaves pedicelled. Leaves blade simple. Leaves lanceolate. Leaf surface smooth. Base
of leaf blade symmetric. Leaf veins prominent. Leaf margin undulate. Leaf apex obtuse. Simple hairs on
leaves absent. Glandular hairs on leaves absent. Bulbs hairs on leaves present. Hispid hairs on leaves absent.
Woolly hairs on leaves absent. Bracteoles absent. Bracteoles not enclosing calyx. Inflorescence circinnate.
Glandular hairs on sepals absent. Hispid hairs on sepals absent. Woolly hair on sepals present. Sepal length
Style hairy. Style terminal. Style inserted. Style undivided. Stigma conical. Stigma much longer than style.
Heliotropium bacciferum Forssk. Ssp. bacciferum var. erosum Hadidy in Boulos

hairs on leaves present. Bracteoles absent. Bracteoles not enclosing calyx. Inflorescence circinnate.
0.2–1 cm. Petal color Yellow or white. Petal outer surface hairy. Petal length 0.4–1.2 cm. Petal apex obtuse.
Petal fusion more than half the length. Petal lobes equal. Corolla throat without scales. No. of stamens 5.
Anthers included. Staminal filaments reduced. Appendix on anther absent. Anthers not sagittate. Style hairy.
Style terminal. Style inserted. Style undivided. Stigma conical. Stigma much longer than style. Ovary
Heliotropium digynum (Forssk.) Asch. ex C. Chr.

present. Glandular hairs on leaves absent. Bulbs hairs on leaves present. Hispid hairs on leaves absent.
18
Glandular hairs on sepals present. Hispid hairs on sepals absent. Woolly hair on sepals present. Sepal length
0.1–0.5 cm. Petal color Yellow or white. Petal outer surface hairy. Petal length 0.3–0.6 cm. Petal apex acute.
Petal fusion more than half the length. Petal lobes equal. Corolla throat without scales. No. of stamens 5.
Anthers included. Staminal filaments reduced. Appendix on anther absent. Anthers not sagittate. Style hairy.
Style terminal. Style inserted. Style undivided. Stigma conical. Stigma as long as style or shorter. Ovary
hairy. Nectar disc present. Gynophore absent.
Heliotropium hirsutissimum Grauer in Weber

alternate. Leaves pedicelled. Leaves blade simple. Leaves ovate. Leaf surface smooth. Base of leaf blade
symmetric. Leaf veins prominent. Leaf margin entire. Leaf apex acute. Simple hairs on leaves present.
on leaves absent. Bracteoles absent. Bracteoles not enclosing calyx. Inflorescence circinnate. Inflorescence
leafless. Number of flowers/bract more than 10. Flower sessile to sub-sessile. Pedicle as long as calyx or
stamens 5. Anthers included. Staminal filaments hairy. Appendix on anther absent. Anthers sagittate. Style
hairy. Style terminal. Style inserted. Style undivided. Stigma conical. Stigma much longer than style. Ovary
hairy. Nectar disc absent. Gynophore absent.
Heliotropium lasiocarpum Fisch. & C.A. Mey.

Plant woody shrub. Stem hairy. Stem erect. Stem branching above the base. Internode length 1–7 cm.
leaf blade symmetric. Leaf veins prominent. Leaf margin entire. Leaf apex obtuse. Simple hairs on leaves
Ovary hairy. Nectar disc absent. Gynophore absent.
Heliotropium ovalifolium Forssk.

Plant woody shrub. Stem hairy. Stem erect. Stem branching from base. Internode length 0.5–3 cm. Basal
leaves alternate. Leaves pedicelled. Leaves blade simple. Leaves ovate. Leaf surface smooth. Base of leaf
blade symmetric. Leaf veins not prominent. Leaf margin entire. Leaf apex acute. Simple hairs on leaves
19
Heliotropium pterocarpum (DC.) Hochst. & Steud. ex Bunge

alternate. Leaves pedicelled. Leaves blade simple. Leaves lanceolate. Leaf surface smooth. Base of leaf blade
symmetric. Leaf veins not prominent. Leaf margin serrate. Leaf apex obtuse. Simple hairs on leaves absent.
leafless. Number of flowers/bract less than 8. Flower sessile to sub-sessile. Pedicle as long as calyx or
stamens 5. Anthers included. Staminal filaments reduced. Appendix on anther absent. Anthers sagittate. Style
glabrous. Style terminal. Style inserted. Style undivided. Stigma conical. Stigma as long as style or shorter.
Heliotropium ramosissimum (Lehm.) Sieb. ex DC.

Plant herb. Stem hairy. Stem erect. Stem branching from base. Internode length 0.5–4 cm. Basal leaves
alternate. Leaves pedicelled. Leaves blade simple. Leaves oblong. Leaf surface smooth. Base of leaf blade
symmetric. Leaf veins not prominent. Leaf margin serrate. Leaf apex acute. Simple hairs on leaves present.
leafless. Number of flowers/bract more than 10. Flower sessile to sub-sessile. Pedicle as long as calyx or
hairy. Style terminal. Style inserted. Style undivided. Stigma conical. Stigma as long as style or shorter.
Heliotropium strigosum Willd. Var. brevifolium (Wal.) C.B. Clarke

Plant woody shrub. Stem hairy. Stem erect. Stem branching above the base. Internode length 0.5–2.5 cm.
Basal leaves alternate. Leaves pedicelled. Leaves blade simple. Leaves linear. Leaf surface smooth. Base of
present. Glandular hairs on leaves absent. Bulbs hairs on leaves present. Hispid hairs on leaves absent.
Ovary glabrous. Nectar disc present. Gynophore present.
20
Heliotropium supinum L.
leaves alternate. Leaves pedicelled. Leaves blade simple. Leaves ovate. Leaf surface smooth. Base of leaf
blade symmetric. Leaf veins prominent. Leaf margin entire. Leaf apex acute. Simple hairs on leaves present.
Glandular hairs on leaves absent. Bulbs hairs on leaves absent. Hispid hairs on leaves absent. Woolly hairs
on leaves present. Bracteoles absent. Bracteoles not enclosing calyx. Inflorescence circinnate. Inflorescence
on sepals absent. Hispid hairs on sepals absent. Woolly hair on sepals present. Sepal length 0.1–0.3 cm. Petal
color Yellow or white. Petal outer surface hairy. Petal length 0.2–0.5 cm. Petal apex obtuse. Petal fusion
more than half the length. Petal lobes equal. Corolla throat without scales. No. of stamens 5. Anthers
included. Staminal filaments glabrous. Appendix on anther absent. Anthers sagittate. Style hairy. Style
terminal. Style inserted. Style undivided. Stigma conical. Stigma as long as style or shorter. Ovary hairy.
Nectar disc absent. Gynophore absent.
Heliotropium zeylanicum (Burm. f.) Lam.

Plant woody shrub. Stem hairy. Stem erect. Stem branching from base. Internode length 0.9–2 cm. Basal
leaves alternate. Leaves pedicelled. Leaves blade simple. Leaves lanceolate. Leaf surface smooth. Base of
leaf blade symmetric. Leaf veins prominent. Leaf margin entire. Leaf apex acute. Simple hairs on leaves
present. Glandular hairs on leaves present. Bulbs hairs on leaves absent. Hispid hairs on leaves absent.
as calyx or shorter. Sepal fusion more than half the length. Apex of calyx lobes acute. Simple hairs on sepals
present. Glandular hairs on sepals absent. Hispid hairs on sepals absent. Woolly hair on sepals absent. Sepal
acute. Petal fusion Less than half the length. Petal lobes equal. Corolla throat without scales. No. of stamens
5. Anthers included. Staminal filaments reduced. Appendix on anther absent. Anthers sagittate. Style hairy.
Style terminal. Style inserted. Style undivided. Stigma conical. Stigma as long as style or shorter. Ovary
Heliotropium curassavicum L.
Plant woody shrub. Stem glabrous. Stem erect. Stem branching from base. Internode length 0.5–2.5 cm.
Basal leaves alternate. Leaves pedicelled. Leaves blade simple. Leaves oblong. Leaf surface wrinkled. Base
of leaf blade symmetric. Leaf veins not prominent. Leaf margin entire. Leaf apex obtuse. Simple hairs on
leaves absent. Glandular hairs on leaves absent. Bulbs hairs on leaves absent. Hispid hairs on leaves absent.
absent. Glandular hairs on sepals absent. Hispid hairs on sepals absent. Woolly hair on sepals absent. Sepal
length 0.1–0.2 cm. Petal color Yellow or white. Petal outer surface glabrous. Petal length 0.1–0.3 cm. Petal
glabrous. Style terminal. Style inserted. Style undivided. Stigma conical. Stigma much longer than style.
Lappula sinaica (DC.) Asch. & Schweinf.

alternate. Leaves sessile to sub-sessile. Leaves blade simple. Leaves lanceolate. Leaf surface smooth. Base of
21
hairs on leaves absent. Bracteoles present. Bracteoles not enclosing calyx. Inflorescence raceme.
Lappula spinocarpos (Forssk.) Asch. ex Kuntze var. spinocarpos

Plant woody shrub. Stem hairy. Stem erect. Stem branching from base. Internode length 1–2 cm. Basal
leaves absent. Glandular hairs on leaves present. Bulbs hairs on leaves absent. Hispid hairs on leaves present.
hairy. Style terminal. Style inserted. Style undivided. Stigma capitate-globose. Stigma as long as style or
Lappula spinocarpos (Forssk.) Asch. ex Kuntze var. inermis Botsch.

Plant woody shrub. Stem hairy. Stem erect. Stem branching from base. Internode length 1–2 cm. Basal
leaves present. Glandular hairs on leaves absent. Bulbs hairs on leaves absent. Hispid hairs on leaves present.
Style glabrous. Style gynobasic. Style inserted. Style bifid. Stigma capitate-globose. Stigma as long as style
Moltkiopsis ciliata (Forssk.) I.M. Johnst.

leaves present. Glandular hairs on leaves absent. Bulbs hairs on leaves present. Hispid hairs on leaves
present. Woolly hairs on leaves absent. Bracteoles present. Bracteoles enclosing calyx. Inflorescence
circinnate. Inflorescence leafless. Number of flowers/bract less than 8. Flower pedicelled. Pedicle as long as
22
length 0.3–0.5 cm. Petal color blue, purple or pink. Petal outer surface hairy. Petal length 0.8–1.3 cm. Petal
stamens 5. Anthers exerted. Staminal filaments hairy. Appendix on anther absent. Anthers not sagittate. Style
hairy. Style gynobasic. Style exerted. Style undivided. Stigma capitate-globose. Stigma as long as style or
shorter. Ovary glabrous. Nectar disc absent. Gynophore present.
Nonea vivianii DC.

present. Glandular hairs on leaves absent. Bulbs hairs on leaves absent. Hispid hairs on leaves present.
Woolly hairs on leaves absent. Bracteoles present. Bracteoles not enclosing calyx. Inflorescence raceme.
shorter. Sepal fusion more than half the length. Apex of calyx lobes acute. Simple hairs on sepals present.
acute. Petal fusion more than half the length. Petal lobes equal. Corolla throat without scales. No. of stamens
5. Anthers included. Staminal filaments hairy. Appendix on anther absent. Anthers not sagittate. Style hairy.
Style gynobasic. Style inserted. Style undivided. Stigma capitate-globose. Stigma as long as style or shorter.
Paracaryum intermedium (Fresen.) Lipsky

hairs on leaves present. Bracteoles absent. Bracteoles not enclosing calyx. Inflorescence raceme.
stamens 5. Anthers included. Staminal filaments hairy. Appendix on anther absent. Anthers not sagittate.
Paracaryum rugulosum (DC.) Boiss.

hairs on leaves absent. Bracteoles present. Bracteoles enclosing calyx. Inflorescence raceme. Inflorescence
color blue, purple or pink. Petal outer surface glabrous. Petal length 0.4–0.6 cm. Petal apex obtuse. Petal
fusion Less than half the length. Petal lobes equal. Corolla throat covered with scales. No. of stamens 5.
Anthers included. Staminal filaments reduced. Appendix on anther absent. Anthers not sagittate. Style
23
or shorter. Ovary glabrous. Nectar disc absent. Gynophore present.
Trichodesma africanum (L.) R.Br. var. abyssinicum Brand in Engl.

Plant herb. Stem hairy. Stem erect. Stem branching from base. Internode length 5–8 cm. Basal leaves
opposite. Leaves pedicelled. Leaves blade simple. Leaves lanceolate. Leaf surface wrinkled. Base of leaf
leafless. Number of flowers/bract less than 8. Flower pedicelled. Pedicle much longer than calyx. Sepal
color blue, purple or pink. Petal outer surface glabrous. Petal length 0.5–1.3 cm. Petal apex acute. Petal
fusion Less than half the length. Petal lobes equal. Corolla throat without scales. No. of stamens 5. Anthers
included. Staminal filaments reduced. Appendix on anther present. Anthers sagittate. Style glabrous. Style
gynobasic. Style inserted. Style undivided. Stigma capitate-globose. Stigma as long as style or shorter. Ovary
Trichodesma africanum (L.) R.Br. var. africanum

leafless. Number of flowers/bract more than 10. Flower pedicelled. Pedicle much longer than calyx. Sepal
on sepals absent. Hispid hairs on sepals absent. Woolly hair on sepals present. Sepal length 0.3–1 cm. Petal
included. Staminal filaments reduced. Appendix on anther present. Anthers sagittate. Style glabrous. Style
terminal. Style inserted. Style undivided. Stigma capitate-globose. Stigma as long as style or shorter. Ovary
Trichodesma africanum (L.) R.Br. var. heterotrichum Bornm. & Kneuk

Plant herb. Stem hairy. Stem erect. Stem branching from base. Internode length 1–5 cm. Basal leaves
color blue, purple or pink. Petal outer surface glabrous. Petal length 0.5–1 cm. Petal apex acute. Petal fusion
Less than half the length. Petal lobes equal. Corolla throat without scales. No. of stamens 5. Anthers
included. Staminal filaments reduced. Appendix on anther present. Anthers sagittate. Style hairy. Style
24
Trichodesma ehrenbergii Schweinf.

Plant woody shrub. Stem hairy. Stem erect. Stem branching from base. Internode length 2–4.9 cm. Basal
leaves opposite. Leaves pedicelled. Leaves blade simple. Leaves ovate. Leaf surface wrinkled. Base of leaf
included. Staminal filaments reduced. Appendix on anther present. Anthers sagittate. Style hairy. Style
hairy. Nectar disc absent. Gynophore absent.
DISCUSSION
Comparison between the key resulting from the present study and previous keys to genera and species
of the Boraginaceae in Egypt reveals the numerous advantages of the former over the latters. The present key
is based on a much wider range of characters from vegetative and floral morphology. The characters were
recorded and used in key generation in a strictly comparative way, so that alternative entries of any couplet in
the key are distinguished from each other by means of contrasting character-states of the same character or
set of characters. The prelude to the key shows that of the 54 characters recorded for each of the 49 taxa, only
34 were sufficient to generate the key successfully with a surplus of 20 characters. All 54 characters are
included in the detailed description of every taxon, thus serving the all-important function of confirming the
identity of these taxa. In fact, the number of confirmatory characters is much greater than the number of
surplus characters. For example, the name of the last taxon in the key would be reached by only 10 characters
so that the confirmatory characters in the detailed description of that taxon would be 44. Other species will
be identified with much fewer characters: the name Heliotropium hirsutissimum will be reached by only two
characters because of its ovate leaves (couplet 1) and hairy staminal filaments (couplet 33).
According to Boulos (2002), the Boraginaceae is represented in the Egyptian flora by 19 genera and 58
species with a number of infra-specific taxa, but the present key comprises only 49 species representing 14
genera. No specimens of the remaining 5 genera and 9 species could be located in the two major herbaria in
Egypt CAI and CAIM. However, if at any future time specimens of any of these missing genera and species
are re-discovered in the country, the data matrix on which the present key is based can be expanded to
accommodate them together with their characters and a re-run of the DELTA package of key-generating
programs would produce the updated key and descriptions.
References
Abbasi, M.; Attar, F. and Ghahremani Nejad, F. (2011). Anatomical studies on several species of
Heliotropium L. in Iran. Notulae Scientia Biologicae, 3(4): 35-43.
Ahmed, H.O. and Kordofani, M.A. (2012). Leaf and stem anatomy of five species from the genus
Heliotropium L. (Boraginaceae) in Sudan. Journal of Chemical and Pharmaceutical Research,
4(10): 4575-4581.
Akçin, O.E. and Baki, H. (2007). Micromorphology and anatomy of three Symphytum (Boraginaceae) taxa
from Turkey. Bangaladesh Journal of Botany, 36(2): 93-103.
Akçin, O.E.; Çoşkunçelebi, K. and Şenel, G. (2012). Foliar anatomy of Cynoglossum L. (Boraginaceae) from
North Anatolia, Turkey. Bangaladesh Journal of Plant Taxonomy, 19(2): 101-108.
Al-Nowaihi, A.S.; Khalifa, S.F. and Hamed, K. (1987). A contribution to the taxonomy of Boraginaceae.
Phytologia, 62: 107-125.
25
Al-Shehbaz, I.A. (1991). The genera of Boraginaceae in the Southeastern United States. Journal of the
Arnold Arborertum, Suppl. Ser., 1: 1-169.
Alwahibi, M. and Bukhary, N. (2013). Anatomical study of four species of Heliotropium L. (Boraginaceae)
from Saudi Arabia. African Journal of Plant Science, 7(1): 35-42.
Andrews, F.W. (1956). The Flowering Plants of the Sudan, volume 3. (Compositae-Gramineae). T. Buncle &
Co., Arbroath, Scotland.
APG III (2014). Angiosperm Phylogeny Group. [http://www.mobot.org].
Barroso, G. M. (1986). Sistemática de Angiospermas do Brasil, v. 3. Universidad Federale do Viçosa.
Bergianska (2014). [http://angio.bergianska.se/asterids/Boraginales/Borag-inales.html].
Bigazzi, M. and Selvi, F. (1998). Pollen morphology in the Boragineae (Boraginaceae) in relation to the
taxonomy of the tribe. Plant Systeatics and Evolution, 213: 121-151.
Bigazzi, M.; Nardi, E. and Selvi, F. (2006). Palynological contribution to the systematics of Rindera and the
allied genera Paracaryum and Solenanthes (Boraginaceae-Cynoglosseae). Willdenowia, 36: 37-46.
Binzet, R. (2011). Pollen morphology of some Onosma species (Boraginaceae) from Turkey. Pakistan
Journal of Botany, 43(2): 731-741.
Binzet, R.; Kandemir, I. and Orcan, N. (2010). Palynological classification of Onosma L. (Boraginaceae)
species from east Mediterranean region in Turkey. Acta Botanica Croatica, 69(2): 259-274.
Boulos, L. (2002). Flora of Egypt. Volume 3. Verbenaceae-Compositae. Al Hadara Publishing, Cairo. 373pp.
Clarke, C.G.S. (1977). The Northwest European Pollen Flora: 10. Boraginaceae. Review of Palaeobotany
and Palynology, 24(2): 59-101.
Coutinho, A.P.; Castro, S.; Carbajal, R.; Ortiz, S.and Serrano, M. (2012). Pollen morphology of the genus
Omphalodes Mill. (Cynoglosseae, Boraginaceae). Grana, 51(3): 194-205.
Dallwitz, M.J. (2010). Overview of the DELTA system. [http://delta-intkey.com/www/overview.htm]
Dallwitz, M.J. and Paine, T.A. (2005). Definition of the DELTA format. [http://delta-
intkey.com/standard.htm]
Dallwitz, M.J., Paine, T.A. and Zurcher, E.J. (1993 onwards). User's guide to the DELTA system: a general
system for processing taxonomic descriptions. 4th edition. [http://delta-intkey.com].
Dasti, A.A.; Bokhari, T.Z.; Malik, S.A. and Akhtar, R. (2003). Epidermal morphology in some members of
family Boraginaceae in Baluchistan. Asian Journal of Plant Sciences, 2: 42-47.
Diane, N.; Jacob, C. and Hilger, H.H. (2003). Leaf anatomy and foliar trichomes in Heliotropiaceae and their
systematic relevance. Flora, 198(6): 468-485.
Diez, M. J. & Valdes, B. (1991). Pollen morphology of the tribes Eritrichieae and Cynoglosseae
(Boraginaceae) in the Iberian Peninsula and its taxonomic significance. Botanical Journal of the
Linnean Society, 107(1): 49-66.
El-Gazzar, A.; Abdel Ghani, M.M.; El-Husseini, N.M. and Khattab, A.A. (2013). Computer-generated keys
to the flora of Egypt. 5. The Leguminosae-Papilionoideae. Assiut University Journal of Botany,
41(2): 129-181.
El-Ghazaly, G. (1995). Pollen morphology of the family Boraginaceae in Qatar. Qatar University Science
Journal, 15(1): 65-75.
Falatoury, A.N.; Pakravan, M. and Sharifnia, F. (2011). Palynological study of Nonea (Boraginaceae-
Boragineae) in Iran. Progress in Biological Sciences, 1(2): 36-43.
Feinbrun-Dothan, N. (1977). Flora Palaestina, part 3, plates. The Israel Academy of Sciences and
Humanities, Jerusalem.
Feinbrun-Dothan, N. (1978). Flora Palaestina, part 3, text. The Israel Academy of Sciences and Humanities,
Jerusalem.
Fukuda, T. and Ikeda, H. (2012). Palynological analysis and taxonomic position of the genus Mertensia
(Boraginaceae). Botany, 90(8): 722-730.
Güven, S.; Beyazoglu, O.; Makbul, S.; Türkmen, Z. and Kandemir, A. (2013). Anatomical features of six
Onosma L. (Boraginaceae) species from Turkey. Iranian Journal of Botany, 19(1): 94-103.
Hargrove, L. and Simpson, M.G. (2003). Ultrastructure of heterocolpate pollen in Cryptantha
(Boraginaceae). International Journal of Plant Sciences, 164: 137-151.
26
Holmgren, P.K., Holmgren N.H. and Barnett L.C. (1990) Index Herbariorum. Part 1. The Herbaria of the
World. Regnum Vegetabile ,vol. 120. pp. 693.
Khatamsaz, M. (2001). Pollen morphology of Iranian Boraginaceae family and its taxonomic significance.
The Iranian Journal of Botany, 9(1): 27-40.
Keshavarzi, M.; Nasrollahi, F. and Sheidai, M. (2013). Stem and fruit anatomical study of the Anchusa
species (Boraginaceae iin Iran. Phytologia Bacanica, 19(2): 193-199.
Liu Jia-Xi; Xi Yizhen; Ning Jian-Chang; Zhang Jing-Mei; Li Yaxuan; Zhao Yun-Yun and Sun Xiao-Hong
(2001a). Pollen morphology and exine ultrastructure of genus Cordia in Boraginaceae in China.
Acta Botanica Sinica, 43(9): 893-898.
Liu Jia-Xi; Zhang Yu-Long; Ning Jian-Chang; Zhao Yun-Yun; Li Ya-Xuan and Zhang Jing-Mei (2001b).
Pollen morphology of the tribe Cynoglosseae of Boraginoideae (Boraginaceae) in China. Acta
Botanica Sinica, 39(6): 515-522.
Mehrabian, A.-R.; Shaidai, M. and Mozaffarian, V. (2014). Morphology of leaf trichomes in Onosma
(Boraginaceae) and their systematic relevance in Iran. Phytologia Balcanica, 20(1): 33-48.
Mehrabian, A.-R.; Shaidai, M.; Noormohammadi, Z.; Mozaffarian, V. and Asrei, Y. (2012). Palynological
diversity in the genus Onosma L. (Boraginaceae) of Iran. Annals of Biological Research, 3(8): 3885-
3893.
Melo, J.I.M.; Sales, M.F. and Santos, F.A.R. (2006). Morfologia polinica das especies de Heliotropium L.
(Boraginaceae) ocorrentes em Pernambuco, Brasil. Ernestia, 16(1): 69-79.
Neuwinger, H.D. (2000). African Traditional Medicine. A dictionary of Plant Uses and Applications.
Medpharm Scientific Publishers, Stuttgart.
Nowicke, J.W. and Miller, J.S. (1990). Pollen morphology of the Cordioideae (Boraginaceae): Auxemma,
Cordia and Patagonula. Plant Systematics and Evolution, 5: 103-121.
Ovchinnikova, S. (2009). On the position of the tribe Eritrichieae in the Boraginaceae system. Botanica
Serbica, 33(2):141-146.
Perveen, A. (2009). Trichomes morphology as a taxonomic marker in the subfamily Heliotropioideae
(Boraginaceae) from Pakistan and Kashmir. International Journal of Biology and Biochemistry, 6:
13-19.
Qureshi, U.S. (1997). Pollen morphology of the family Boraginaceae of Pakistan and Kashmir. Ph.D. thesis,
Sind University, Pakistan. 246 pp.
Rabaey, D.; Lens, F.; Smets, E. and Jansen, S. (2010). The phylogenetic significance of vestured pits in
Boraginaceae. Taxon, 59(2): 510-516.
Retief, E. and Vanwyk, A.E. (1997). Palynology of southern African Boraginaceae: the genera Lobostemon,
Echiostachys and Echium. Grana, 36: 271-278.
Selvi, F. and Bigazzi, M. (2001). Leaf surface and anatomy in Boraginaceae tribe Boragineae with respect to
ecology and taxonomy. Flora, 196: 269-285.
Scheel, R.; Ybert, J.-P. and Barth, O.M. (1996). Pollen morphology of the Boraginaceae from Santa Catarina
State (southern Brazil), with comments on the taxonomy of the family. Grana, 35: 138-153.
Stearn, W.T. (1966). Botanical Latin. Hafner Publishing Co., New York. 566 pp.
Taia, W.K. (2006). Family Boraginaceae. Hair variations and their significance in the systematic of the
genera. Asian Journal of Plant Sciences, 5(3): 441-454.
Thorne, R.F. (1992). Classification and geography of the flowering plants. Botanical Review, 58(3): 225-348.
Trivedi, M.L.; Khanna, V. and Shailja, J. (1976). Nodal anatomy of crtain members of Boraginaceae.
Proceedings of the Indian Academy of Science – section Botany, 84: 31-36.
Ventrella, M.C. and Marinho, C.R. (2008). Morphology and histochemistry of glandular trichomes of Cordia
verbenacea DC. (Boraginaceae) leaves. Revista Brasil Botanica, 31(3): 457-467.
Watson, L. and Dallwitz, M.J. (1992 onwards). The Families of Flowering Plants: descriptions, identification
and information retrieval. Version 19th , August 2014. [http://delta-intkey.com].
27
Appendix 1. List of taxa and collection data of the 14 genera and 49 species representing the Boraginaceae in
the flora of Egypt and used in key-generation. Genera and species are arranged alphabetically.
No. Taxa Collection data
1 Alkanna tinctoria Tausch. Gunnar Täckholm, 26/3/1927, Abukir; CAI

A. El Hadidy, 19/8/1982, Wadi el Arbain, S. Sinai; CAI.
2 Alkanna orientalis (L.) Boiss. Nabil El Hdidy, 10/5/1965, outside the monastery of st.
Catherine; CAI.
Zohary & Jaffe, 17/2/1931, Jerusalem, eastern slopes of Mt.
3 Alkanna strigosa Boiss. & Hohen.
Scopus; CAI
4 Anchusa aegyptiaca (L.) DC. Adel Gazzar, March 1973, Amria; CAI
Loutfy Boulos, 28/4/1955, 3km north of El-Arish; CAI.
A. Kadry, Spring 1964, Arabia : Khurais Rd., 75km from
5 Anchusa hispida Forssk.
Riyadh; CAI.
Täckholm, et al, 2/6/1964, Ras El-Hekma; CAI
6 Anchusa humilis (Desf.) I.M. Johnst. Alaa Amer, 20/3/1987, Alexandria; CAI
Anchusa milleri Spreng. M. Kassas, 26/2/1964, Wadi Amloug 50km south of Suez; CAI
7
Amal Amin, 13-14/3/1952, Mariut; CAI
Anchusa undulata L. ssp. hybrida (Ten.)
8 Mustafa Imam, 22/3/1965, Alexandria-Burg El Arab road; CAI.
Beg. In Beg &
Christina Brydolf, 12/3/1969, Cyrene, Cyrenaica (Libya); CAI.
Arnebia decumbens (Vent.) Coss. &
9 Botany Dept. Excursion, 2/5/1955, Ras El Hekma; CAI.
Kralik var. decumbens
M. N. Shourbagy, June 1956, Hamadan, west of Tehran, Iran;
Arnebia decumbens (Vent.) Coss. & CAI.
10
Kralik var. macrocalyx Coss. & Loutfy Boulos, W. Jallad and J. Lahham, 11/4/1974, 35km NE of
H-4; CAI.
11 Arnebia hispidissima (Lehm.) DC. K. H. Batanouny, 22/1/1990, Hurgada-Safaga Road; CAI
Arnebia linearifolia DC. A. El Hadidy, 3/4/1980, Cairo- Alexandria desert Road 160km
12
far from Cairo; CAI.
13 Arnebia tinctoria Forssk. Nabil El Hadidi & K. H. Batanouny, 2/3/1956, Suez Road; CAI.
14 Asperugo procumbens L. A. El Hadidy, 24/4/1983, Gebel Catherine, St. Catherine S. Sinai;
CAI.
Buglossoides incrassata (Guss.) I.M. E. K. Balls & W. B. Garlay, 2/5/1934, Asia Minor (Turkey);
15
Johnst. CAI.
Buglossoides tenuiflora (L.f.) I.M. Christina Brydolf, 22/2/1970, 20 km north Tan-Tan, Morocco;
16
Johnst. CAI.
Coldenia procumbens L. J. R. Shabetai, Z4478, 15/10/1941, around Lake Faruk, Kom
17
Oshim, Fayum; CAI.
18 Echiochilon fruticosum Desf. Vivi Taecholm, 16/2/1965, Burg el Arab – Alamein road; CAI.
Vivi Täckholm, I. Elsayed and M. El Mahdi, 7/4/1967, Iking
Echium angustifolium Mill. ssp.
19 Mariut; CAI.
angustifolium
Amal Amin 13/3/1972, Burg el Arab; CAI.
Echium angustifolium Mill. Ssp
20 Ibrahim El-Garf, 29/12/1989, Sidi Barrani; CAI.
sericeum (Vahl)
21 Echium horridum Batt. Vivi Täckholm, et al, 6/2/1961, Bir Hafafit, Red Sea Coast; CAI.
Echium rauwolfii Delile Vivi Täckholm, L. Boulos, and M. Zahran, 14-18/11/1963 and
22
11/2/1964, Aniba (near Aswan) and Kom Ombo; CAI.
Christina Brydolf, April 1968, Leptis Magna, Libya; CAI.
23 Echium rubrum Forssk Ibrahim El-Garf, April 1999, Wadi Hashem, 48km before Mersa
Matruh; CAI.
Echium sabulicolum Pomel var A. G. Fahmy 1084, 2/5/1988, Mersa Matruh- Sallum Road 60 km
24
sabulicolum east of Sallum; CAI.
R. Muschler, 4/1904, Idfu; CAI.
25 Heliotropium aegyptiacum Lehm.
Dr. Pfund, (1875-1876), Dongola, Soudan; CAI.
26 Heliotropium arabinense Fresen. Gaher, 17/3/1957, Burg el Arab; CAI
28
Heliotropium bacciferum Forssk. ssp.

27 M. Kassas, 26/2/1964, 60km south of Suez; CAI.
bacciferum var. bacciferum
Heliotropium bacciferum Forssk ssp
28 Loutfy Boulos, 27/5/1961, Grand Canaries; CAI.
bacciferum var erosum Hadidy in Boulos
29 Heliotropium digynum (Forssk.) Asch. Adel Gazzar, 5/2/1973, Nekhel, Central Sinai; CAI.
Ex C. Chr. Wafaa Amer, 28/3/2005, Bir El-Abd, N. Sinai; CAI.
Heliotropium hirsutissimum Grauer in Gwaine, 16/6/1907, Sporting Club, Alexandria; CAI.
30
Weber Vivi Täckholm, July 1962, Tulkarm, Jordan; CAI.
Imam, Ibrahim, Mahdi and Sisi, 31/10/1971, 5km south of
Rosetta; CAI.
Heliotropium lasiocarpum Fisch. & C.A.
31 Adel Gazzar, 1/6/1975, Faculty of Science Garden, Alexandria;
Mey.
CAI.
Ibrahim El-Garf, 28/4/1955, Wadi Araba; CAI.
Heliotropium ovalifolium Forssk. M. Kassas, et al. E39(a), E35, 7/12/1967, Jebel Avitola- Khor
32
Tagando, Kassala, Sudan; CAI
Heliotropium pterocarpum (DC.) Vivi Täckholm et al 1495a., 5/2/1962, Mouth of Wadi Laseitit,
33 Gebel Elba; CAI.
Hochst. & Steud. Ex Bunge
Vivi Taeckholm et al, 9/2/1962, Gebel Elba; CAI.
Heliotropium ramosissimum (Lehm.) M. Kassas et al 410, 10/12/1966, Khor Gwab, Red Sea, Sudan;
34
Sieb. Ex DC. CAI.
M. Kassas 75, 12/10/1953, Merkheat Hills, Sudan; CAI.
Heliotropium strigosum Willd. Var Vivi Täckholm et al, 1025, 28/1/1962, Gebel Elba; CAI.
35
brevifolium (Wal.) C.B. V. Täckholm et al, 928. 28/1/1962, Wadi Mawaw, Gebel Elba;
CAI.
Nabil El Hadidy, 22/7/1954, on the drain, near Tukh, Qalubeya;
36 Heliotropium supinum L.
CAI.
Heliotropium zeylanicum (Burm. F.)
37 Gunnar Täckholm, 23-27/1/1929, Gebel Elba; CAI
Lam.
Heliotropium curassavicum L. Täckholm, Nabil and Imam, 3/9/1966, Wadi Digla, near Maadi;
38
CAI
Lappula sinaica (DC.) Asch. &
39 Schweinfurth, (original burnt in Berlin); CAI.
Schweinf.
Lappula spinocarpos (Forssk.) Asch. ex
40 M. Imam, 4/4/1957, Suez desert Road; CAI.
Kuntze var spinocarpos
Lappula spinocarpos (Forssk.) Asch. ex Loutfy Boulos 8757, 7/5/1976, University Campus, Al- Jubaiha,
41
Kuntze var inermis Botsch. near Amman, Jordan; CAI.
42 Moltkiopsis ciliata (Forssk.) I.M. Johnst. Wafaa Amer, Gebel Libna, N. Sinai; CAI.
Loutfy Boulos, 24/3/1955, Khan Yunis, Palestine; CAI.
43 Nonea vivianii DC. Loutfy Boulos 2256, 13/3/1968, Al- Tamimi, East Derna, Gebel
Akhdar, Libya; CAI.
Paracaryum intermedium (Fresen.) Azza El Hadidy, 23/4/1983, S. Sinai, Wadi Tala; CAI.
44
Lipsky Adel Gazzar, Feb. 1993, El Hasana, C. Sinai; CAI.
L. Boulos, David Addiss and Mazin Qumsiyeh, 8718, 2/5/1976,
45 Paracaryum rugulosum (DC.) Boiss.
Wadi Bayir, Jordan; CAI.
Kassas et al, 287, 11/12/1965, Golol, Sudan; CAI.
Trichodesma africanum (L.) R.Br. var
46 Dale Osborn & Ibrahim Helmy, 10/4/1967, Gebel Uwainat, Wadi
abyssinicum Brand in Engl.
el Brins; CAI.
Trichodesma africanum (L.) R.Br. var Vivi Täckholm el al, 4/3/1961, Wadi Hankalia, near the gold
47
africanum mines Red Sea coast; CAI.
Trichodesma africanum (L.) R.Br. var.
48 Vivi Täckholm et al, 22/11/1960, El Maadi desert; CAI.
heterotrichum Bornm. & Kneuk.
Trichodesma ehrenbergii Schweinf. Khattab 152B, 16/1/1945, Gibal Arforat, Mecca, Saudi Arabia;
49
CAI

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1

Computer-generated keys to the flora of Egypt. 6.

*Corresponding author: elgazzar_adel@hotmail.com

MATERIAL AND METHODS

#34. Petal color/1. blue, purple or pink/2. Yellow or white/

Figure 1. Anchusa undulata ssp. hybrida.

Figure 2. Arnebia hispidissima.

Figure 3. Heliotropium strigosum.

Figure 4. Trichodesma africanum.

2(1). Staminal filaments hairy......................................................................................................................... 3

3(2). Leaf apex acute; hispid hairs on leaves present; woolly

4(2). Leaf margin entire...................................................................................................................................5

7(2). Leaf margin entire................................................................................................................................... 8

9(8). Leaf apex acute; plant woody shrub; leaves pedicelled;

11(10). Style hairy; style terminal; ovary hairy; nectar disc

13(1). Staminal filaments hairy....................................................................................................................... 14

15(13). Hispid hairs on leaves present; hispid hairs on sepals present.............................................................16

16(15). Leaf apex acute.................................................................................................................................... 17

18(16). Style hairy; leaf veins prominent; bulbs hairs on leaves

19(18). Inflorescence raceme; number of flowers/bract more than 10; glandular

20(15). Leaf apex

21(20). Stem erect; style glabrous; plant woody shrub;

24(13). Style hairy; simple hairs on leaves absent........................................................................................... 25

27(1). Staminal filaments hairy........................................................................................................................28

30(29). Basal leaves alternate; leaf apex acute; style glabrous;

31(29). Leaf margin entire; leaf apex obtuse; style hairy;

32(27). Plant herb; stem hairy; leaves sessile to sub-sessile;

33(1). Staminal filaments hairy.............................................. Heliotropium hirsutissimum Grauer in Weber

34(33). Leaf margin entire; flower pedicelled; style terminal.......................................................................... 35

38(33). Leaf apex acute; bracteoles present; bracteoles enclosing

39(38). Basal leaves alternate; leaf margin undulate; style glabrous;

plant herb..................................................................................................Alkanna orientalis (L.) Boiss.

3.2.2.3 Detailed descriptions

Alkanna tinctoria Tausch.

Alkanna orientalis (L.) Boiss.

Alkanna strigosa Boiss. & Hohen.

Anchusa aegyptiaca (L.) DC.

Anchusa hispida Forssk.

Anchusa humilis (Desf.) I.M. Johnst.

Anchusa milleri Spreng.

Anchusa undulata L. ssp. hybrida (Ten.) Beg. in Beg & Furi

Arnebia decumbens (Vent.) Coss. & Kralik var. decumbens

Arnebia hispidissima (Lehm.) DC.

Arnebia linearifolia DC.

Arnebia tinctoria Forssk.

Buglossoides incrassata (Guss.) I.M. Johnst.

Buglossoides tenuiflora (L.f.) I.M. Johnst.

Echiochilon fruticosum Desf.

Echium angustifolium Mill. ssp. angustifolium

Echium angustifolium Mill. ssp. sericeum (Vahl) Klotz

Echium horridum Batt.

Echium rauwolfii Delile

Echium rubrum Forssk.

Echium sabulicolum Pomel var. sabulicolum

Heliotropium aegyptiacum Lehm.

Heliotropium arabinense Fresen.

Heliotropium bacciferum Forssk. Ssp. bacciferum var. bacciferum

Heliotropium bacciferum Forssk. Ssp. bacciferum var. erosum Hadidy in Boulos

Heliotropium digynum (Forssk.) Asch. ex C. Chr.