Asteraceae, Apiaceae and Related Groups

1.1. The Place of Asteraceae and Apiaceae in the APG IV System

Modern systematics of flowering plants relies on the phylogenetic APG IV system (2016), in which groups are distinguished primarily by descent rather than by external similarity. According to this system, both Asteraceae and Apiaceae belong to the large clade Asterids — one of the main branches of eudicots (APG IV, 2016; cited in Singh, 2019). Within the asterids, both groups fall into the informal clade Campanulids, but they belong to different orders: Asteraceae to the order Asterales, and Apiaceae to the order Apiales (APG IV, 2016; cited in Singh, 2019). This is a fundamentally important point: the external similarity of umbel-like inflorescences and capitulum-like inflorescences does not mean close kinship. Before us are two different evolutionary lineages that independently achieved high economic importance.

The order Asterales includes, besides Asteraceae, such families as Campanulaceae — the bellflowers, including lobelias (Lammers, 2007), Goodeniaceae — the goodenia family (Carolin, 2007), Menyanthaceae — the buckbean family, Calyceraceae — the calycera family, and a number of others. The order Apiales includes Apiaceae — the umbellifers, Araliaceae — the aralia family, Pittosporaceae — the pittosporum family, Myodocarpaceae, Pennantiaceae, and some other groups (APG IV, 2016; cited in Singh, 2019). The same large branch Campanulids also includes the order Dipsacales, in its modern broad sense including Caprifoliaceae s.l., into which in APG III–IV were also included Dipsacaceae — the teasel family — and Valerianaceae — the valerian family (Mayer, 2016; Hofmann & Bittrich, 2016; Weberling & Bittrich, 2016).

Thus, already at the entrance to the course we see two different “poles” of diversity. Asteraceae is the largest family of flowering plants, a kind of evolutionary giant. Apiaceae is a family of considerably smaller volume, but with an exceptionally high concentration of food, spice-aromatic, medicinal, and poisonous plants. Their comparison allows us to show that economic importance is not always proportional to the number of species, and that systematics helps one see, behind external diversity, deep differences in structure, biochemistry, and evolutionary history.

1.2. Asteraceae — the Largest Family of Flowering Plants

Asteraceae, or Compositae — the composites — is the largest family of flowering plants. According to modern estimates, it includes more than 1600 genera and about 23,000 species (Anderberg et al., 2007). This is a cosmopolitan group, absent only from Antarctica. Its diversity is especially great in herbaceous communities, savannas, and mountain regions, whereas in humid tropical lowland forests the composites are comparatively few (Anderberg et al., 2007). Such wide distribution is connected with a unique combination of morphological, biochemical, and reproductive adaptations.

The main diagnostic feature of Asteraceae is the capitulum inflorescence. It may contain from 1 to 500 or more small flowers seated on a common enlarged receptacle. On the outside, the capitulum is surrounded by an involucre of bracts — phyllaries. The receptacle often bears membranous scales — paleae, or receptacular bracts. The individual flowers in the capitulum are usually small, with an inferior ovary, and the calyx is transformed into a pappus, consisting of bristles, scales, or teeth. The corolla is sympetalous: tubular, ligulate, bilabiate, or pseudoligulate. The stamens usually unite by their anthers into a tube, and the stigma brings the pollen out — this is the so-called secondary pollen presentation, characteristic of composites (Anderberg et al., 2007). The fruit is a cypsela, often with a pappus, which ensures anemochory, epizoochory, or other methods of dispersal.

Within Asteraceae, several large subfamilies are distinguished: Barnadesioideae, Mutisioideae, Carduoideae, Cichorioideae, and Asteroideae (Anderberg et al., 2007). Barnadesioideae is a small South American group occupying a basal position; the remaining subfamilies include the main diversity of the family. Cichorioideae contain, in particular, the tribe Cichorieae — the chicory tribe, to which lettuce, chicory, and dandelion belong. Asteroideae is the largest subfamily, including sunflower, chamomile, yarrow, wormwood, marigolds, calendula, and many others.

The economic importance of Asteraceae is exceptionally great and diverse. Among the most important cultivated plants one may name Helianthus annuus — the common sunflower, one of the world’s main oilseed species, and H. tuberosus — Jerusalem artichoke, or earth apple. Among vegetables is Lactuca sativa — garden lettuce. Cynara cardunculus gives artichoke and cardoon. Carthamus tinctorius — safflower, a source of edible and technical oil, as well as a dye. Cichorium intybus — chicory, used as a vegetable, coffee substitute, and fodder plant. Tanacetum cinerariifolium — pyrethrum, a source of natural insecticides — pyrethrins. Artemisia dracunculus — tarragon, a spice-aromatic plant. Many species of Artemisia — wormwoods — are medicinal, essential-oil, fodder, and weed plants. Calendula, Matricaria, Achillea, Taraxacum are widely known as medicinal and ornamental plants (Anderberg et al., 2007). Composites also provide fodder grasses, weed species, and ornamental crops — from asters and chrysanthemums to dahlias and marigolds.

Thus, Asteraceae is an example of an evolutionary “explosion”: an enormous number of species, a wide spectrum of life forms, diverse inflorescences and fruits, and high biochemical specialization. In the agrosphere they are represented by oilseed, vegetable, medicinal, essential-oil, ornamental, fodder, and weed plants.

1.3. Apiaceae — a Small but Strategically Important Family

Apiaceae, or Umbelliferae — the umbellifers, or celery family — is a family including, according to modern data, 466 genera and about 3800 species (Reduron, 2021). In number of species it is considerably inferior to Asteraceae, but in economic concentration of valuable plants it occupies one of the first places among flowering plants. Umbellifers are distributed almost worldwide, but the greatest diversity is concentrated in temperate regions of Eurasia and North America. In humid tropics they are rare and confined mainly to high mountains. Mediterranean and arid conditions favor high species diversity (Reduron, 2021).

Modern ideas about the origin of Apiaceae connect their appearance with Australasia — the region including Australia, Tasmania, New Zealand, New Guinea, New Caledonia, and a number of island groups. The origin of the family is dated to the Late Cretaceous — Early Eocene, about 87 million years ago. The divergence of the main subfamilies in the Southern Hemisphere occurred in the interval 45.9–71.2 million years ago: Apiodeae and Saniculoideae — in South Africa, Azorelloideae — in South America, Mackinlayoideae — in Australasia (Reduron, 2021). In the APG IV system, Apiaceae belong to the order Apiales (APG IV, 2016; cited in Singh, 2019).

Within the family, four subfamilies are distinguished: Apiodeae, Saniculoideae, Azorelloideae, and Mackinlayoideae (Reduron, 2021). All the main cultivated vegetable and spice-aromatic umbellifers belong to Apiodeae. Saniculoideae include only one widely used aromatic plant — Eryngium foetidum, or sawtooth coriander. Azorelloideae and Mackinlayoideae contain no commonly cultivated vegetables or spices (Reduron, 2021). Thus, the economic core of the family is concentrated in one large subfamily.

Morphologically, Apiaceae are easily recognized by the compound umbel — an inflorescence in which the main axis bears rays ending in simple umbels. Usually there is an involucre — of bracts at the base of the common umbel — and an involucel — at the base of the partial umbels. Leaves are often repeatedly dissected, with sheathing petioles. Plants are almost always aromatic thanks to a network of secretory canals containing essential oils, resins, coumarins, and other compounds (Reduron, 2021). The fruit is a schizocarp, at maturity splitting into two mericarps suspended on a carpophore. On the surface of the mericarps, ribs are clearly visible, and in the grooves between them are vittae, or oil canals. These features — the number of ribs, fruit shape, presence and arrangement of vittae — are of key importance for identifying genera and species (Reduron, 2021).

The economic importance of Apiaceae is enormous. Among root vegetables, one should first name Daucus carota — carrot, Apium graveolens — celery, including root celery, and Pastinaca sativa — parsnip. Less often used are the roots of parsley (Petroselinum crispum subsp. tuberosum), alexanders, or smyrnium — Smyrnium olusatrum, great earthnut — Bunium bulbocastanum, earth chestnut — Conopodium majus, skirret — Sium sisarum, turnip-rooted chervil — Chaerophyllum bulbosum. In North America, the underground parts of Canadian honewort — Cryptotaenia canadensis, Indian parsley — Lomatium spp., and yampah — Perideridia gairdneri are sometimes eaten. In South America, the roots of arracacha — Arracacia xanthorrhiza — are widely prepared. In Africa, a few root umbellifers from the genera Annesorhiza and Chamarea are known (Reduron, 2021).

Leafy parts are also widely used: petioles and leaves of celery and fennel — Foeniculum vulgare, as well as numerous spice-aromatic plants: alexanders, angelica — Angelica spp., chervil — Anthriscus cerefolium, coriander — Coriandrum sativum, dill — Anethum graveolens, parsley — Petroselinum crispum, sweet cicely — Myrrhis odorata (Reduron, 2021).

Many umbellifers produce edible fruits, which in everyday speech are called “seeds”: ajwain — Trachyspermum ammi, anise — Pimpinella anisum, caraway — Carum carvi, celery, coriander, cumin — Cuminum cyminum, dill, fennel, zira — Elwendia persica (Reduron, 2021). Some species are used to flavor drinks: anise — in anisette, ouzo, and raki; caraway — in kümmel and aquavit; angelica — in Chartreuse and vermouth (Reduron, 2021).

In medicine, umbellifers are known as sources of antispasmodic, carminative, cosmetic, diuretic, laxative, sedative, stimulating, and stomachic remedies. Among pharmacologically important species are bishop’s weed (Ammi majus), Asiatic pennywort (Centella asiatica), species of Bupleurum (Bupleurum spp.), Phloidocarpus sibiricus, Visnaga daucoides. Some yield resins and gums: Dorema ammoniacum, species of Ferula, including F. assa-foetida, F. gummosa, F. tingitana (Reduron, 2021).

At the same time, among Apiaceae there are dangerous poisonous plants: hemlock water dropwort — Oenanthe crocata, poison hemlock — Conium maculatum, water hemlock — Cicuta spp. Some species cause dermatitis when wet skin contacts bright sun — for example, hogweeds (Heracleum) and parsnip (Pastinaca) (Reduron, 2021). Finally, many umbellifers are grown as ornamentals: angelica, astrantia, bupleurum, sea holly, hogweed (Reduron, 2021).

Thus, Apiaceae is a family with a relatively moderate number of species, but with an extremely high concentration of plants that determine the food, medicinal, and cultural traditions of many peoples. If Asteraceae is a “giant in number of species,” then Apiaceae is a “giant in economic density.”

1.4. Two Poles — Two Evolutionary Paths

Comparison of Asteraceae and Apiaceae shows two different evolutionary paths. Asteraceae bet on the pseudanthium — the capitulum, which functions as one large flower — on an enormous number of small flowers, on secondary pollen presentation, on a diversity of cypsela fruits with pappi, and on broad biochemical radiation. This allowed them to become the largest family of flowering plants and to occupy almost all terrestrial habitats except Antarctica (Anderberg et al., 2007).

Apiaceae took the path of the compound umbel, aromatic secretory canals, the schizocarp with vittae, and a comparatively narrow but very effective food and medicinal specialization. They are not so rich in species, but it is among them that carrot, celery, parsnip, parsley, coriander, dill, fennel, caraway, anise, and many other crops are concentrated (Reduron, 2021). Moreover, this family also contains dangerous poisonous plants, which makes accurate identification especially important.

For a specialist in the agrosphere, knowledge of the systematics of these families is necessary for several reasons. First, it allows one to correctly identify species, genus, and family, which is critical for phytosanitary control, quarantine, and plant protection. Second, understanding phylogenetic relationships helps predict potential pests and diseases, as well as select related species for breeding. Third, the morphology of inflorescences and fruits — the capitulum and cypsela in Asteraceae, the compound umbel and schizocarp in Apiaceae — underlies identification manuals and diagnostic keys.

Thus, Asteraceae and Apiaceae are two examples of successful but fundamentally different evolutionary strategies. Asteraceae is the largest family of flowering plants, setting the “quantitative” pole of diversity. Apiaceae is a family of moderate volume, but with exceptional economic value, setting the “qualitative” pole of concentration of useful plants. Both groups belong to the clade Asterids, but belong to different orders — Asterales and Apiales (APG IV, 2016; cited in Singh, 2019). Their further study in the module will be built on a sequential analysis of morphology, phylogeny, key genera, and practical importance.

2.1. Phylogenetic Position and Monophyly

Asteraceae, or Compositae — the composites — occupy the order Asterales in the APG IV system (2016). This order belongs to the clade Campanulids, which in turn belongs to the large clade Asterids (APG IV, 2016; cited in Singh, 2019). Thus, the closest relatives of composites among economically important groups are the bellflowers — Campanulaceae (Lammers, 2007), goodenia — Goodeniaceae (Carolin, 2007), buckbeans — Menyanthaceae, as well as calyceras — Calyceraceae and a number of other families (Anderberg et al., 2007). All of them together with Asteraceae form the Asterales clade, well supported by molecular and morphological data.

The monophyly of Asteraceae is beyond doubt. It is confirmed both by molecular-phylogenetic analyses and by a unique combination of morphological characters. The most important synapomorphies of the family are the capitulum inflorescence, syngenesious anthers, an inferior ovary with a single basal ovule, a calyx transformed into a pappus, and secondary pollen presentation, in which pollen is shed into the tube of united anthers and brought out by the style (Anderberg et al., 2007). Not one of these characters individually is absolutely unique, but their combination reliably diagnoses the family.

According to modern estimates, Asteraceae include more than 1600 genera and about 23,000 species (Anderberg et al., 2007). This is the largest family of flowering plants, cosmopolitan, absent only from Antarctica. The greatest diversity is concentrated in herbaceous communities, savannas, and mountain regions; in humid tropical lowland forests composites are comparatively few (Anderberg et al., 2007).

2.2. General Morphological Characterization

Asteraceae are predominantly herbaceous plants, but among them there are subshrubs, shrubs, lianas, and even trees up to 30 m tall and more. Underground organs are diverse: taproots, rhizomes, tubers, caudices. Stems are usually erect, but may be prostrate, climbing, sometimes succulent. Leaves are almost always alternate, less often opposite or whorled, without stipules, simple or variously dissected. Many composites are characterized by secretory canals (resin ducts) and laticifers containing latex; laticifers are especially abundant in Cichorioideae (Anderberg et al., 2007).

Anatomically, composites are rather uniform. They are characterized by anisocytic or anomocytic stomata, simple or glandular hairs, and calcium oxalate druses are common. In the stem, medullary bundles often develop, and secondary growth may be anomalous, especially in lianas. The wood of composites is usually without tracheids, with libriform fibers; vessels with simple perforations (Anderberg et al., 2007).

2.3. The Capitulum Inflorescence — the Key Character of the Family

The main diagnostic feature of Asteraceae is the capitulum inflorescence. It is a terminal or axillary inflorescence with an enlarged receptacle, on which small sessile flowers are densely, often spirally, arranged. On the outside, the capitulum is surrounded by an involucre, consisting of one or several rows of phyllaries — bracts that may be herbaceous, leathery, membranous, sometimes brightly colored and serving to attract pollinators (Anderberg et al., 2007). The receptacle may be naked, pitted, hairy, or bear paleae — membranous bracts that sometimes completely cover the cypselae.

The number of flowers in a capitulum varies from 1 to 500 and more. The capitulum functions as a single “flower” — a classic example of a pseudanthium, or false flower. Such a structure ensures high pollination efficiency and economy of material in forming reproductive organs. It was the capitulum that allowed composites to occupy enormous territories and become one of the most successful families of flowering plants.

2.4. Flowers, Androecium, Gynoecium, and Secondary Pollen Presentation

The flowers of Asteraceae are always small, usually 5-merous, with an inferior ovary. The calyx is reduced and transformed into a pappus, consisting of bristles, scales, or teeth; sometimes the pappus is absent. The corolla is sympetalous, tubular, ligulate, bilabiate, or pseudoligulate. In a capitulum there may be flowers of different types: tubular (actinomorphic), ligulate (zygomorphic, with a 5-toothed ligule), bilabiate (with two lips), pseudoligulate (with a 3-toothed ligule), and filiform (sterile or female) (Anderberg et al., 2007).

The androecium consists of 5 stamens attached to the corolla. The anthers unite into a tube surrounding the style. Stamens are introrse, opening by longitudinal slits. The gynoecium is of two carpels, the ovary inferior, unilocular, with one basal ovule. The style is one, the stigma bilobed. Pollen is shed into the anther tube while still in the bud, and then brought out by the growing style — this is secondary pollen presentation. In most composites the flowers are protandrous: first functioning as male, then as female. This promotes cross-pollination (Anderberg et al., 2007).

2.5. Fruits, Seeds, and Dispersal

The fruit of Asteraceae is a cypsela, dry, one-seeded, indehiscent. The cypsela often bears a pappus, which may function as a parachute for anemochory, as a hook for epizoochory, or as an elaiosome for myrmecochory. In some composites the fruits have wings, spines, hooks, or fleshy coverings. Seeds usually lack endosperm or have a thin layer of it; the embryo is straight and large (Anderberg et al., 2007).

Dispersal of fruits is diverse: anemochory (dandelion, groundsel), epizoochory (beggarticks, burdock), myrmecochory (some species with elaiosomes), endozoochory (fleshy fruits in some tropical composites). In a number of species heterocarpy is observed — the formation of fruits of different types within one capitulum or one plant, which broadens the possibilities for dispersal and germination (Anderberg et al., 2007).

2.6. Karyology, Biochemistry, and Reproductive Systems

Chromosome numbers of Asteraceae vary from n = 2 to n = 120, but x = 9 is most common, and it is probably ancestral for the family. Polyploidy, aneuploidy, and dysploidy are widespread and play an important role in the evolution of the group. Many composites are self-sterile, which promotes cross-pollination; apomixis has been noted in some (Anderberg et al., 2007).

Biochemically, Asteraceae are extremely rich. They are characterized by inulin as a storage carbohydrate, acetylenic compounds, sesquiterpene lactones, flavonoids, essential oils, and other metabolites. It is these compounds that determine the medicinal, insecticidal, aromatic, and toxic properties of many representatives. For example, pyrethrins of Tanacetum cinerariifolium are natural insecticides; sesquiterpene lactones of many species are toxic to livestock; chicory inulin is used as a prebiotic (Anderberg et al., 2007).

2.7. Internal Phylogeny and Subfamilies

Modern classification of Asteraceae distinguishes several subfamilies reflecting the main phylogenetic lines. The following are most recognized: Barnadesioideae, Mutisioideae, Carduoideae, Cichorioideae, and Asteroideae (Anderberg et al., 2007). Sometimes Gymnarrhenoideae, Corymbioideae, and others are also distinguished, but in this review we will limit ourselves to the main ones.

Barnadesioideae is a small South American group (9 genera, 91 species) occupying a basal position in the family. It is characterized by unique three-celled hairs, spiny pollen, and the absence of an inversion in chloroplast DNA that is present in all other composites. This is an important phylogenetic marker (Anderberg et al., 2007).

Mutisioideae is a predominantly South American group (about 60 genera, 720 species), characterized by bilabiate corollas, long tailed anthers, and often large inflorescences. It includes the genera Mutisia, Gerbera, Nassauvia, and others (Anderberg et al., 2007).

Carduoideae is a large group (at least 93 genera, 2600 species), distributed mainly in the Old World. It is characterized by spiny leaves and involucres, tubular flowers, and often compound inflorescences. The most important tribe is Cynareae, or Cardueae, including artichoke, cardoon, safflower, thistles, knapweeds, and burdocks (Anderberg et al., 2007).

Cichorioideae is a large group (241 genera, about 2900 species), characterized by ligulate flowers, laticifers with latex, and often the absence of tubular flowers. The most important tribe is Cichorieae, including lettuce, chicory, dandelion, black salsify, and salsify (Anderberg et al., 2007).

Asteroideae is the largest subfamily (1210 genera, about 17,000 species), including most composites. It is characterized by tubular flowers in the center and ligulate or pseudoligulate flowers on the periphery, as well as a double tectum of pollen grains. It includes sunflower, chamomile, yarrow, wormwood, calendula, marigolds, asters, chrysanthemums, and many others (Anderberg et al., 2007).

2.8. Most Important Genera and Species

Among the enormous diversity of Asteraceae, the following genera and species are of particular economic importance.

Helianthus — sunflower. H. annuus — common sunflower, one of the world’s main oilseed crops, a source of edible and technical oil, as well as valuable fodder. H. tuberosus — Jerusalem artichoke, or earth apple, a root-tuber vegetable and fodder plant (Anderberg et al., 2007).

Lactuca — lettuce. L. sativa — garden lettuce, one of the most widespread vegetable crops, including many varieties: leaf, head, romaine, asparagus, and others (Anderberg et al., 2007).

Cynara — artichoke. C. cardunculus — artichoke and cardoon; in the first, the unopened inflorescences (capitula) are eaten, in the second, the leaf petioles (Anderberg et al., 2007).

Carthamus — safflower. C. tinctorius — safflower, a source of edible and technical oil, as well as a dye (Anderberg et al., 2007).

Tanacetum — pyrethrum. T. cinerariifolium — Dalmatian pyrethrum, a source of natural insecticides — pyrethrins (Anderberg et al., 2007).

Artemisia — wormwood. Includes many species, among which are medicinal, essential-oil, fodder, and weed plants. A. dracunculus — tarragon, a spice-aromatic plant (Anderberg et al., 2007).

Calendula — calendula. C. officinalis — pot marigold, a widely used medicinal and ornamental plant (Anderberg et al., 2007).

Matricaria — chamomile. M. chamomilla — German chamomile, a source of medicinal essential oil and anti-inflammatory remedies (Anderberg et al., 2007).

Achillea — yarrow. A. millefolium — common yarrow, a medicinal and ornamental plant (Anderberg et al., 2007).

Cichorium — chicory. C. intybus — common chicory, a vegetable, fodder, and medicinal plant; the roots are used as a coffee substitute and a source of inulin (Anderberg et al., 2007).

Taraxacum — dandelion. T. officinale — common dandelion, a food, medicinal, and weed plant; widely known for its cypselae with a pappus (Anderberg et al., 2007).

In addition, important are Smallanthus sonchifolius — yacón, a root-tuber plant of the Andes; Stevia rebaudiana — stevia, a source of natural sweeteners; Echinacea purpurea — purple coneflower, an immunostimulant medicinal plant; Guizotia abyssinica — niger seed, an oilseed crop of Africa; Scorzonera hispanica — black salsify, a root vegetable; Tragopogon porrifolius — salsify, or oyster plant; Arctium lappa — greater burdock, a medicinal and food plant (Anderberg et al., 2007).

2.9. Economic Importance

The economic importance of Asteraceae is exceptionally great and diverse. In agriculture they are represented by several groups of crops.

Oilseeds. Sunflower (Helianthus annuus) is the most important oilseed crop of the temperate zone. Safflower (Carthamus tinctorius) yields edible and technical oil. Niger seed (Guizotia abyssinica) is an oilseed crop of Africa and India (Anderberg et al., 2007).

Vegetables. Lettuce (Lactuca sativa), artichoke and cardoon (Cynara cardunculus), chicory (Cichorium intybus), endive (Cichorium endivia), black salsify (Scorzonera hispanica), salsify (Tragopogon porrifolius), Jerusalem artichoke (Helianthus tuberosus), yacón (Smallanthus sonchifolius), Indian lettuce (Pterocypsela indica), para cress (Acmella oleracea), wasabi-like plants, crown daisy (Glebionis coronaria), butterbur (Petasites japonicus) (Anderberg et al., 2007).

Medicinal. Calendula (Calendula officinalis), chamomile (Matricaria chamomilla), yarrow (Achillea millefolium), wormwood (Artemisia* spp.), echinacea (Echinacea purpurea), dandelion (Taraxacum officinale), beggarticks (Bidens spp.), burdock (Arctium lappa), milk thistle (Silybum marianum), stevia (Stevia rebaudiana**) (Anderberg et al., 2007).

Essential-oil and spice-aromatic. Tarragon (Artemisia dracunculus), wormwood (Artemisia* spp.), tansy (Tanacetum spp.), chamomile (Matricaria** spp.) (Anderberg et al., 2007).

Insecticidal. Pyrethrum (Tanacetum cinerariifolium) — a source of pyrethrins, natural insecticides (Anderberg et al., 2007).

Ornamental. Asters (Aster spp., Symphyotrichum spp., Callistephus spp.), chrysanthemums (Chrysanthemum spp.), dahlias (Dahlia spp.), marigolds (Tagetes spp.), zinnias (Zinnia spp.), gerberas (Gerbera spp.), gazanias (Gazania spp.), daisies (Bellis spp.), calendula (Calendula spp.), yarrow (Achillea spp.), echinacea (Echinacea spp.), black-eyed Susan (Rudbeckia spp.), goldenrod (Solidago spp.), gaillardia (Gaillardia spp.), cosmos (Cosmos spp.), ornamental sunflower (*Helianthus spp.), and many others (Anderberg et al., 2007).

Fodder. Sunflower (oilcake, silage), Jerusalem artichoke (green mass, tubers), fodder varieties of lettuce, chicory, dandelion, yarrow (Anderberg et al., 2007).

Weeds. Many composites are noxious weeds: thistle (Cirsium spp.), sow thistle (Sonchus spp.), plumeless thistle (Carduus spp.), knapweed (Centaurea spp.), Russian knapweed (Acroptilon spp.), cotton thistle (Onopordum spp.), rush skeletonweed (Chondrilla spp.), ragweed (Ambrosia spp.), cocklebur (Xanthium spp.), groundsel (Senecio spp.), dandelion (Taraxacum* spp.), common sow thistle (Sonchus oleraceus), shepherd’s purse? No, shepherd’s purse is Brassicaceae. Also Parthenium hysterophorus causes contact dermatitis, and pollen of Ambrosia** causes hay fever (Anderberg et al., 2007).

Toxic. Many species of Senecio are hepatotoxic to livestock; Xanthium is poisonous; some Centaurea, Cirsium may accumulate nitrates (Anderberg et al., 2007).

2.10. Conclusions

Asteraceae is the largest family of flowering plants, distinguished by the unique capitulum inflorescence, secondary pollen presentation, an inferior ovary with one ovule, and a cypsela fruit, often with a pappus. The family includes five main subfamilies: Barnadesioideae, Mutisioideae, Carduoideae, Cichorioideae, and Asteroideae, of which the last is the largest and economically most important. Among Asteraceae are the most important oilseed, vegetable, medicinal, essential-oil, insecticidal, ornamental, fodder, and weed plants. Sunflower, lettuce, artichoke, chicory, chamomile, calendula, yarrow, wormwood, and pyrethrum are only a small part of the enormous economic potential of this family. Knowledge of the systematics of Asteraceae is necessary for species identification, breeding, plant protection, and rational use of plant resources.

3.1. Phylogenetic Position and Monophyly

Apiaceae, or Umbelliferae — the umbellifers, or celery family — occupy the order Apiales in the APG IV system (2016). This order, like Asterales, belongs to the clade Campanulids, which belongs to the large clade Asterids (APG IV, 2016; cited in Singh, 2019). Thus, Apiaceae and Asteraceae are not closely related groups, but representatives of different evolutionary lines within one large asterid branch. Their external similarity in inflorescence structure (umbel and capitulum) is not the result of close kinship, but represents an example of convergent evolution — the independent origin of similar solutions in different lineages.

The order Apiales includes, besides Apiaceae, such families as Araliaceae — the aralia family (ginseng, aralia, ivy), Pittosporaceae — the pittosporum family, Myodocarpaceae, Pennantiaceae, and some other groups (APG IV, 2016; cited in Singh, 2019). The closest relatives of Apiaceae among economically important groups are Araliaceae, which is confirmed by both molecular and morphological data. The monophyly of Apiaceae is beyond doubt and is confirmed by a unique combination of characters: compound umbel, schizocarp, secretory canals with essential oils, and the characteristic structure of fruits with ribs and vittae (Reduron, 2021).

Modern ideas about the origin of Apiaceae connect their appearance with Australasia — the region including Australia, Tasmania, New Zealand, New Guinea, New Caledonia, and a number of island groups. The origin of the family is dated to the Late Cretaceous — Early Eocene, about 87 million years ago (Nicolas and Plunkett, 2014; cited in Reduron, 2021). The divergence of the main subfamilies in the Southern Hemisphere occurred in the interval 45.9–71.2 million years ago: Apiodeae and Saniculoideae — in South Africa, Azorelloideae — in South America, Mackinlayoideae — in Australasia (Calvino et al., 2016; cited in Reduron, 2021). Thus, Apiaceae is an ancient group whose evolutionary history is closely connected with Gondwana and the subsequent breakup of continents.

According to modern estimates, Apiaceae include 466 genera and about 3800 species (Plunkett et al., 2018; cited in Reduron, 2021). In number of species they are considerably inferior to Asteraceae, but in economic concentration of valuable plants they occupy one of the first places among flowering plants. Umbellifers are distributed almost worldwide, but the greatest diversity is concentrated in temperate regions of Eurasia and North America. In humid tropics they are rare and confined mainly to high mountains. Mediterranean and arid conditions favor high species diversity (Reduron, 2021).

3.2. General Morphological Characterization

Apiaceae are predominantly herbaceous plants, but among them there are subshrubs, shrubs, and even small trees. Some species are aquatic and marsh plants. Stems are usually erect, hollow or filled with pith, often ribbed, furrowed. Underground organs are diverse: taproots, rhizomes, tubers, bulbs. Leaves are almost always alternate, rarely opposite or whorled, usually repeatedly dissected, with sheathing petioles. Sheaths may be inflated, as in Ferula. Leaf blades are often pinnately or palmately dissected, less often entire (Bupleurum). Many umbellifers are characterized by heterophylly — different leaf shapes on one plant (Reduron, 2021).

Anatomically, Apiaceae are characterized by the presence of secretory canals (resin ducts) located in roots, stems, leaves, and fruits. These canals contain essential oils, resins, coumarins, polyacetylenes, and other compounds that determine the aroma, taste, and biological activity of the plants. The family is also characterized by laticifers, especially in aboveground organs. Stomata are anomocytic. Hairs are simple, multicellular, sometimes glandular. Calcium oxalate druses and starch grains are often found in tissues (Reduron, 2021).

3.3. The Umbel Inflorescence — the Key Character of the Family

The main diagnostic feature of Apiaceae is the compound umbel. It is an inflorescence in which the main axis (peduncle) bears rays — first-order branches ending in simple umbels (umbellules). Simple umbels consist of pedicels bearing individual flowers. Thus, the compound umbel is an inflorescence of inflorescences, which ensures compact arrangement of a large number of small flowers and high pollination efficiency (Reduron, 2021).

At the base of the common umbel there is often an involucre of bracts. At the base of the partial umbels there is an involucel of bracteoles. The presence, number, and shape of involucre and involucel bracts are important diagnostic characters. In some genera (for example, Coriandrum) the involucre and involucel may be absent or reduced. In Daucus the involucre consists of repeatedly dissected bracts, which is a characteristic feature. In Astrantia and some other genera, the involucre bracts are large, brightly colored, and serve to attract pollinators (Reduron, 2021).

Flowers in the umbel are usually small, 5-merous, actinomorphic or slightly zygomorphic. The calyx is reduced to five teeth or completely absent. The corolla consists of five free petals, usually white, yellow, pink, or purple. Petals often have tips curved inward. The androecium consists of five stamens alternating with the petals. The gynoecium is of two carpels, the ovary inferior, bilocular, with one ovule in each locule. There are two styles, enlarged at the base into a stylopodium — a nectary. Flowers are usually protandrous, which promotes cross-pollination. Pollinators are diverse insects: bees, flies, beetles, butterflies (Reduron, 2021).

3.4. Fruits, Seeds, and Their Taxonomic Significance

The fruit of Apiaceae is a schizocarp, at maturity splitting into two mericarps suspended on a carpophore — a thin axis, forked or entire. Each mericarp contains one seed. On the surface of the mericarps, ribs are clearly visible: five primary ribs — three dorsal and two marginal. Ribs may be filiform, prominent, keeled, corky, toothed, winged, spiny, or barely noticeable. In the grooves between the ribs — valleculae — are located vittae — oil canals containing essential oils. Vittae may be solitary or numerous, located in the valleculae or on the commissural side. The number, shape, and arrangement of vittae are the most important diagnostic characters for identifying genera and species (Reduron, 2021).

The shape of the fruit, degree of compression, development of ribs, presence of wings or spines, and number of vittae — all these characters are widely used in the systematics of Apiaceae. Fruits may be almost spherical (Coriandrum), bilobed (Bifora), beaked (Anthriscus, Scandix), dorsally compressed (Pastinaca, Peucedanum), or laterally compressed (Torilis, Visnaga, Smyrnium). Seeds usually have endosperm, the embryo is small and straight (Reduron, 2021).

Dispersal of fruits is diverse. Anemochory is characteristic of plants with winged or compressed fruits (Laserpitium, Thapsia, Prangos). Epizoochory — for plants with spiny fruits (Daucus, Torilis). In some species the entire compound umbel functions as a dispersal unit (Falcaria, Petagnaea, Trinia). Vegetative reproduction also occurs: by rhizomes, stolons, tubercles (Reduron, 2021).

3.5. Karyology and Reproductive Systems

Chromosome numbers of Apiaceae vary from n = 3 (Sium suave) to n = 77 (Lomatium columbianum, 2n = 154). The most common is n = 11, giving the most frequent diploid level 2n = 22. Polyploidy occurs, but is comparatively rare among vegetable umbellifers — only in Daucus montanus (2n = 66) and some North American species of Lomatium and Perideridia. Aneuploidy and dysploidy have been noted in the genera Bunium and Bupleurum. Genome size (C-value) varies from 0.63 pg (Oenanthe fistulosa) to 5.48 pg (Daucus montanus). B-chromosomes have been found in 24 genera and 40 species (Pimenov et al., 2003; cited in Reduron, 2021).

Reproductive systems of Apiaceae are diverse. Most species are cross-pollinated, but self-pollinated and apomictic forms occur. Flowers are usually protandrous. In some species andromonoecy is observed — the presence of male and bisexual flowers on one plant. Less often gynodioecy (female and bisexual flowers on different plants) or dioecy (male and female flowers on different plants) occurs. Many umbellifers are characterized by asynchrony of flowering: first the central umbel opens, then the lateral ones; moreover, in the first umbels bisexual flowers predominate, while in later ones the proportion of male flowers increases. This is of practical importance for seed production: for seed collection, umbels of the first two flowering phases should be used (Reduron, 2021).

3.6. Internal Phylogeny and Subfamilies

Modern classification of Apiaceae distinguishes four subfamilies: Apiodeae, Saniculoideae, Azorelloideae, and Mackinlayoideae (Reduron, 2021). Sometimes Platysace and other groups are also distinguished, but in this review we will limit ourselves to the main ones.

Apiodeae is the largest subfamily, including the overwhelming majority of genera and species. It is characterized by vittae in the valleculae and on the commissural side, as well as rib oil canals, usually very small or absent. All the main cultivated vegetable and spice-aromatic umbellifers belong to this subfamily: carrot, celery, parsnip, parsley, coriander, dill, fennel, caraway, anise, cumin, lovage, angelica, chervil, alexanders, skirret, turnip-rooted chervil, and many others (Reduron, 2021).

Saniculoideae is a small subfamily characterized by large rib oil canals and usually the absence of vittae in the valleculae. The fruit surface often bears scales, bristles, or spines. The only widely used aromatic plant of this subfamily is Eryngium foetidum, or sawtooth coriander, used as a spice in tropical Asia and Latin America (Reduron, 2021).

Azorelloideae is a subfamily characterized by an endocarp of fibrous sclereids, absence of vittae in the valleculae, presence of rib oil canals (but not large), and dorsally compressed fruits. It contains no commonly cultivated vegetables or spices (Reduron, 2021).

Mackinlayoideae is a subfamily with an endocarp of fibrous sclereids, absence of vittae in the valleculae, presence of rib oil canals, and laterally compressed fruits. It contains no commonly cultivated plants used as vegetables or spices (Reduron, 2021).

Thus, the economic core of the family is concentrated in the subfamily Apiodeae, which is important to take into account when studying cultivated umbellifers.

3.7. Most Important Genera and Species

Among the enormous diversity of Apiaceae, the following genera and species are of particular economic importance.

Daucus — carrot. D. carota — carrot, one of the world’s most important root vegetable crops. Wild carrot (D. carota subsp. carota) is a common weed. Cultivated carrot (D. carota subsp. sativus) includes many cultivar groups: eastern, western, varieties with orange, yellow, red, purple roots. Carrot is rich in carotenoids, sugars, vitamins, and minerals. It is used fresh, for processing, and as a fodder crop (Reduron, 2021).

Apium — celery. A. graveolens — celery, including three main varieties: leaf celery (var. secalinum), petiole celery (var. dulce), and root celery (var. rapaceum). All are widely cultivated as vegetable, spice-aromatic, and medicinal plants. Celery is rich in essential oils, flavonoids, and vitamins. It is used fresh, for processing, and as a seasoning (Reduron, 2021).

Pastinaca — parsnip. P. sativa — parsnip, a root vegetable and fodder plant. Parsnip roots are rich in sugars, starch, and vitamins. It is used fresh, for processing, and as a fodder crop. Wild parsnip (P. sativa subsp. sylvestris) is a common plant of meadows and roadsides and can cause dermatitis (Reduron, 2021).

Carum — caraway. C. carvi — caraway, a spice-aromatic and medicinal plant. The fruits (seeds) of caraway contain essential oil with carvone and are used as a spice in baking, cheese-making, and in the production of drinks (kümmel, aquavit). Caraway is also used in medicine as a carminative, antispasmodic, and choleretic remedy (Reduron, 2021).

Petroselinum — parsley. P. crispum — parsley, including leaf and root varieties. Leaf parsley (var. crispum, var. neapolitanum) is used as a spice-aromatic plant. Root parsley (var. tuberosum) is a root vegetable. Parsley is rich in essential oils, flavonoids, and vitamins, especially vitamin C and carotenoids (Reduron, 2021).

Coriandrum — coriander. C. sativum — coriander, including two main varieties: coriander (var. sativum) — a spice-aromatic plant whose fruits are used as a spice (coriander), and cilantro (var. microcarpum) — a vegetable plant whose leaves are used fresh. Coriander is widely cultivated in the Mediterranean, India, China, and Latin America. The fruits contain essential oil with linalool and are used in perfumery, medicine, and cooking (Reduron, 2021).

Anethum — dill. A. graveolens — dill, a spice-aromatic and medicinal plant. Leaves and young shoots are used fresh as a seasoning. The fruits (seeds) of dill are used as a spice in canning, baking, and for essential oil production. Dill is rich in essential oils, flavonoids, and vitamins (Reduron, 2021).

Foeniculum — fennel. F. vulgare — fennel, including several varieties: common fennel (var. vulgare) — a spice-aromatic and medicinal plant, vegetable fennel (var. dulce) — a vegetable plant with thickened petioles used fresh, sweet fennel (var. dulce) — a source of essential oil. Fennel is widely used in cooking, medicine, and perfumery. The fruits contain essential oil with anethole and are used as a carminative and antispasmodic remedy (Reduron, 2021).

Conium — poison hemlock. C. maculatum — poison hemlock, a deadly poisonous plant. It contains alkaloids (coniine, conhydrine, and others) that cause respiratory paralysis. All parts of the plant are poisonous, especially the fruits and rhizomes. Poison hemlock is a common plant of waste places, roadsides, and dumps. It is often confused with parsley, celery, and dill, which leads to severe poisoning. Extreme caution is required when collecting wild umbellifers (Reduron, 2021).

In addition, important are Levisticum officinale — lovage, a spice-aromatic and medicinal plant; Angelica archangelica — garden angelica, a spice-aromatic, medicinal, and ornamental plant; Anthriscus cerefolium — chervil, a spice-aromatic plant; Myrrhis odorata — sweet cicely, a spice-aromatic plant; Smyrnium olusatrum — alexanders, a vegetable and spice-aromatic plant; Sium sisarum — skirret, or sugar root, a root vegetable; Chaerophyllum bulbosum — turnip-rooted chervil, a root vegetable; Bunium bulbocastanum — great earthnut, a root vegetable; Conopodium majus — earth chestnut, a root vegetable; Arracacia xanthorrhiza — arracacha, a root vegetable of South America; Cryptotaenia canadensis — Canadian honewort, a vegetable plant of North America; Perideridia gairdneri — yampah, a root vegetable of North America; Lomatium spp. — Indian parsley, root vegetables of North America (Reduron, 2021).

3.8. Economic Importance

The economic importance of Apiaceae is exceptionally great and diverse. In agriculture they are represented by several groups of crops.

Root vegetables. Carrot (Daucus carota), root celery (Apium graveolens* var. rapaceum), parsnip (Pastinaca sativa), root parsley (Petroselinum crispum var. tuberosum), alexanders (Smyrnium olusatrum), great earthnut (Bunium bulbocastanum), earth chestnut (Conopodium majus), skirret (Sium sisarum), turnip-rooted chervil (Chaerophyllum bulbosum), arracacha (Arracacia xanthorrhiza*) (Reduron, 2021).

Leafy vegetables and spice-aromatic plants. Leaf and petiole celery (Apium graveolens), leaf parsley (Petroselinum crispum), chervil (Anthriscus cerefolium), cilantro (Coriandrum sativum* var. microcarpum), dill (Anethum graveolens), vegetable fennel (Foeniculum vulgare var. dulce), lovage (Levisticum officinale), angelica (Angelica archangelica), sweet cicely (Myrrhis odorata), alexanders (Smyrnium olusatrum), sawtooth coriander (Eryngium foetidum*) (Reduron, 2021).

Spice-aromatic (fruits, “seeds”). Caraway (Carum carvi), coriander (Coriandrum sativum), dill (Anethum graveolens), fennel (Foeniculum vulgare), anise (Pimpinella anisum), cumin (Cuminum cyminum), ajwain (Trachyspermum ammi), zira (Elwendia persica), celery (Apium graveolens), parsley (Petroselinum crispum) (Reduron, 2021).

Medicinal. Anise (Pimpinella anisum), caraway (Carum carvi), fennel (Foeniculum vulgare), coriander (Coriandrum sativum), dill (Anethum graveolens), lovage (Levisticum officinale), angelica (Angelica archangelica), bishop’s weed (Ammi majus), Asiatic pennywort (Centella asiatica), species of Bupleurum (Bupleurum spp.), Phloidocarpus sibiricus, Visnaga daucoides, Dorema ammoniacum, species of Ferula (Reduron, 2021).

Essential-oil. Anise, caraway, fennel, coriander, dill, lovage, angelica, ammi, ferula (Reduron, 2021).

Poisonous. Poison hemlock (Conium maculatum), hemlock water dropwort (Oenanthe crocata), water hemlock (Cicuta spp.), poison hemlock (Conium maculatum), hogweeds (Heracleum spp.), wild parsnip (Pastinaca sativa* subsp. sylvestris**). Some species cause dermatitis when wet skin contacts bright sun — hogweeds, parsnip (Reduron, 2021).

Weeds. Many umbellifers are common weeds: ground elder (Aegopodium podagraria), Sosnowsky’s hogweed (Heracleum sosnowskyi), poison hemlock (Conium maculatum), water dropwort (Oenanthe spp.), water hemlock (Cicuta spp.), wild parsnip (Pastinaca sativa* subsp. sylvestris), wild carrot (Daucus carota subsp. carota), wild chervil (Anthriscus spp.), cow parsley (Anthriscus sylvestris**) (Reduron, 2021).

Ornamental. Angelica (Angelica spp.), astrantia (Astrantia spp.), bupleurum (Bupleurum spp.), sea holly (Eryngium spp.), hogweed (Heracleum spp.), hare’s ear (Bupleurum spp.), tingin-like plants (Reduron, 2021).

Fodder. Carrot, parsnip, celery, hogweed (silage), ground elder, cow parsley (Reduron, 2021).

Thus, Apiaceae is a family with a relatively moderate number of species, but with an extremely high concentration of plants that determine the food, medicinal, and cultural traditions of many peoples. If Asteraceae is a “giant in number of species,” then Apiaceae is a “giant in economic density.”

3.9. Conclusions

Apiaceae is a large and economically important family of flowering plants, occupying the order Apiales in the APG IV system. The monophyly of the family is confirmed by a unique combination of characters: compound umbel, schizocarp, secretory canals with essential oils, and the characteristic structure of fruits with ribs and vittae. The family includes four subfamilies: Apiodeae, Saniculoideae, Azorelloideae, and Mackinlayoideae, of which the first is the largest and economically most important. Among Apiaceae are the most important root, vegetable, spice-aromatic, medicinal, essential-oil, poisonous, and weed plants. Carrot, celery, parsnip, parsley, coriander, dill, fennel, caraway, anise, and cumin are only a small part of the enormous economic potential of this family. Knowledge of the systematics of Apiaceae is necessary for species identification, breeding, plant protection, and rational use of plant resources. Of particular importance is the ability to distinguish cultivated and medicinal umbellifers from poisonous ones, which requires knowledge of diagnostic characters of vegetative organs, inflorescences, and fruits.

4.1. Araliaceae — the Aralia Family

Araliaceae — the aralia family — is a small but economically important family of the order Apiales, the closest relative of Apiaceae in the APG IV system (2016). The family includes about 50 genera and 1500 species, distributed mainly in tropical and subtropical regions, less often in temperate latitudes (APG IV, 2016; cited in Singh, 2019). Unlike Apiaceae, aralias are predominantly woody plants: trees, shrubs, lianas, sometimes epiphytes. Leaves are usually alternate, simple or compound, often large, with sheathing petioles. Inflorescences are umbels, heads, or racemes, often gathered into compound inflorescences. Flowers are small, usually 5-merous, with an inferior ovary. The fruit is a berry or drupe. Aralias are characterized by secretory canals with essential oils, resins, and triterpene saponins — it is these compounds that determine their medicinal importance.

The most important genus is Panax — ginseng. P. ginseng — Asian ginseng, a perennial herbaceous plant with a fleshy root, distributed in the forests of Korea, Manchuria, and Primorye. Ginseng root contains ginsenosides — triterpene saponins with adaptogenic, tonic, and immunomodulatory action. Ginseng is widely used in traditional Chinese medicine and modern pharmacology. P. quinquefolius — American ginseng, cultivated in North America. P. notoginseng — notoginseng, or sanqi, used in Chinese medicine as a hemostatic and analgesic remedy (Singh, 2019).

Another important genus is Aralia — aralia. A. elata — Japanese aralia, or devil’s walking stick, a large shrub or small tree with prickly trunks, distributed in the Far East. Aralia roots contain aralosides — triterpene saponins with tonic and adaptogenic action. A. racemosa — American spikenard, a perennial herbaceous plant of North America whose roots were used by Native Americans as a medicine. A. nudicaulis — wild sarsaparilla, also a North American medicinal plant (Singh, 2019).

Hedera — ivy. H. helix — common ivy, an evergreen liana widely cultivated as an ornamental plant. Ivy is used for greening walls, fences, and creating hedges. Ivy leaves contain saponins and flavonoids and are used in folk medicine. H. canariensis — Canary ivy, a larger liana with large leaves (Singh, 2019).

Other economically important genera: Eleutherococcus — eleuthero. E. senticosus — Siberian ginseng, a shrub with prickly shoots distributed in the Far East. Roots contain eleutherosides with adaptogenic action. Oplopanax — devil’s club. O. elatus — devil’s club, a shrub with prickly stems whose roots are used as a tonic. Kalopanax — castor aralia. K. septemlobus — castor aralia, or dimorphanthus, a large tree with prickly trunks distributed in the Far East, an ornamental and medicinal plant. Fatsia — fatsia. F. japonica — Japanese fatsia, an evergreen shrub with large palmate leaves, widely cultivated as an ornamental plant. Schefflera — schefflera, many species of which are grown as house and greenhouse plants. Polyscias — polyscias, ornamental house plants (Singh, 2019).

Thus, Araliaceae is a family close to Apiaceae, but predominantly woody. Its economic importance is connected mainly with medicinal (ginseng, aralia, eleuthero, devil’s club) and ornamental (ivy, fatsia, schefflera, polyscias) plants.

4.2. Pittosporaceae, Myodocarpaceae, Pennantiaceae — an Overview

In the order Apiales, besides Apiaceae and Araliaceae, several other families are distinguished that do not have such broad economic importance but are important for understanding phylogenetic relationships.

Pittosporaceae — the pittosporum family — includes about 9 genera and 200–240 species, distributed mainly in Australia, New Zealand, tropical Asia, and Africa. These are trees, shrubs, or lianas with simple alternate leaves, often leathery, and bisexual 5-merous flowers with an inferior ovary. The fruit is a capsule or berry; seeds are often covered with a sticky resinous coat — hence the family name. Among economically important representatives is Pittosporum tobira — Japanese pittosporum, widely cultivated as an ornamental shrub in the subtropics. P. undulatum — sweet pittosporum, an ornamental and medicinal plant. P. crassifolium — karo, used in New Zealand as an ornamental and windbreak plant (APG IV, 2016; cited in Singh, 2019).

Myodocarpaceae is a small family including 2 genera and about 19 species, distributed in New Caledonia, Australia, and New Guinea. These are trees and shrubs with simple leaves and small flowers. Economic importance is small; some species are used as ornamentals and timber plants (APG IV, 2016; cited in Singh, 2019).

Pennantiaceae is a small family including 1 genus and about 4 species, distributed in New Zealand, Australia, and Pacific islands. These are trees and shrubs with simple leaves and small flowers. Economic importance is limited to timber use and ornamental qualities (APG IV, 2016; cited in Singh, 2019).

These families are important for understanding that Apiales is a large and diverse group including both herbaceous (Apiaceae) and woody (Araliaceae, Pittosporaceae, Myodocarpaceae, Pennantiaceae) forms. The economic core of the order is concentrated in Apiaceae and Araliaceae, while the remaining families play a secondary role.

4.3. Campanulaceae — the Bellflower Family

Campanulaceae — the bellflower family — is a large family of the order Asterales, the closest relative of Asteraceae in the APG IV system (2016). The family includes, according to modern estimates, 84 genera and about 2400 species, distributed worldwide, but most diverse in temperate and subtropical regions, especially in the Mediterranean, southern Africa, the Andes, and the Hawaiian Islands (Lammers, 2007). In the APG IV system, Campanulaceae also include Lobeliaceae — the lobelia family — previously considered a separate family. Thus, the modern understanding of Campanulaceae s.l. unites bellflowers and lobelias into one family.

Morphologically, Campanulaceae are predominantly herbaceous plants, less often subshrubs, shrubs, lianas, and even small trees. The family is characterized by laticifers with white or colored latex, alternate, less often opposite or whorled leaves, simple or pinnately dissected. Flowers are usually bisexual, 5-merous, with an inferior ovary, a sympetalous corolla of bell-shaped, funnel-shaped, or tubular form. Stamens usually 5, free or united by anthers. Style one, stigma lobed. Fruit a capsule, less often a berry. The family is characterized by secondary pollen presentation, in which pollen is shed into the tube of united anthers and brought out by the style — this brings Campanulaceae closer to Asteraceae (Lammers, 2007).

The most economically important genera are Campanula — bellflower. It includes about 420 species, distributed mainly in temperate regions of the Northern Hemisphere. Many species are cultivated as ornamentals: C. persicifolia — peach-leaved bellflower, C. carpatica — Carpathian bellflower, C. glomerata — clustered bellflower, C. medium — Canterbury bells, C. alpestris, C. arvatica, C. waldsteiniana — alpine species popular in rock gardens. C. rapunculus — rampion bellflower, a vegetable plant of Europe whose young leaves and roots were eaten. Some bellflower species are used in folk medicine (Lammers, 2007).

Lobelia — lobelia. It includes more than 400 species, distributed mainly in tropical and subtropical regions, especially in South America. Among economically important species: L. erinus — edging lobelia, a widely cultivated annual ornamental plant for flowerbeds, balconies, and hanging baskets. L. cardinalis — cardinal flower, a perennial ornamental plant of North America with bright red flowers. L. inflata — Indian tobacco, a source of lobeline — a pyridine alkaloid used in medicine as an antitussive, bronchodilator, and anti-smoking agent. L. siphilitica, L. tupa, and other species are used in folk medicine and as ornamentals (Lammers, 2007).

Platycodon — balloon flower. P. grandiflorus — balloon flower, a perennial herbaceous plant of East Asia, widely cultivated as an ornamental. The roots are used in Chinese medicine as jie-geng or kikyo — an expectorant, antitussive, and anti-inflammatory remedy. The active principles are saponins (platycodin, kikyosaponin) (Lammers, 2007).

Codonopsis — codonopsis. C. pilosula — hairy codonopsis, or dang shen, a perennial herbaceous plant of China whose roots are used in traditional Chinese medicine as a tonic and general strengthening remedy, a substitute for ginseng. C. javanica, C. convolvulacea, and other species are also used in medicine. C. clematidea — an ornamental plant (Lammers, 2007).

Other economically important genera: Adenophora — ladybell, ornamental and medicinal plants; Jasione — sheep’s bit, ornamental plants; Michauxia — michauxia, ornamental plants; Phyteuma — rampion, ornamental plants; Trachelium — throatwort, ornamental plants; Wahlenbergia — wahlenbergia, ornamental and medicinal plants; Canarina — canarina, ornamental plants; Cyananthus — bluebell, alpine ornamental plants; Ostrowskia — ostrowskia, ornamental plants; Platycodon — balloon flower, an ornamental and medicinal plant (Lammers, 2007).

Thus, Campanulaceae is a large family of the order Asterales, whose economic importance is connected mainly with ornamental (bellflowers, lobelias, balloon flower, codonopsis, ladybells) and medicinal (lobelia, balloon flower, codonopsis) plants. Some species are used as vegetables (rampion bellflower). Campanulaceae demonstrate the transition from insect-pollinated entomophilous forms to specialized systems with secondary pollen presentation, which brings them closer to Asteraceae.

4.4. Goodeniaceae, Menyanthaceae — an Overview

Goodeniaceae — the goodenia family — is a small family of the order Asterales, including 11 genera and about 440 species, distributed mainly in the Southern Hemisphere, chiefly in Australia (Carolin, 2007). These are herbaceous plants, subshrubs, shrubs, less often lianas. The family is characterized by an indusium — a cup-shaped structure at the tip of the style that collects pollen and presents it to pollinators — a unique structure not found in other flowering plants. Flowers are usually 5-merous, with an inferior or semi-inferior ovary, often zygomorphic. Fruit a capsule, nut, or drupe. Economic importance is small: some species of Scaevola (for example, S. taccada — beach naupaka, S. aemula — fan flower) are cultivated as ornamentals and used to stabilize coastal sands. Brunonia australis — an ornamental plant. Lechenaultia — ornamental plants of Australia. The economic importance of Goodeniaceae is limited to ornamental use and some medicinal uses in Australian traditional medicine (Carolin, 2007).

Menyanthaceae — the buckbean family — is a small family of the order Asterales, including 5 genera and about 60 species, distributed worldwide, but mainly in temperate and subtropical regions. These are aquatic or marsh herbaceous plants with creeping rhizomes and alternate, less often opposite leaves. Flowers are usually 5-merous, with an inferior ovary, often heterostylous. Fruit a capsule or berry. The most important genus is Menyanthes — buckbean. M. trifoliata — bogbean, or buckbean, a marsh plant of the temperate Northern Hemisphere, used in folk medicine as a bitter, tonic, and choleretic remedy. Nymphoides — floating heart, aquatic plants with floating leaves and yellow or white flowers, cultivated as ornamentals in ponds and aquariums. Villarsia — villarsia, aquatic and marsh plants of Australia and Asia. Liparophyllum — liparophyllum, aquatic plants. The economic importance of Menyanthaceae is limited to ornamental (Nymphoides) and medicinal (Menyanthes) use (Carolin, 2007; APG IV, 2016; cited in Singh, 2019).

Goodeniaceae and Menyanthaceae are important for understanding phylogenetic relationships within Asterales: they demonstrate a gradual transition from insect-pollinated forms with open flowers to specialized systems with secondary pollen presentation, characteristic of Asteraceae. Their economic importance is small, but they play an important role in understanding the evolution of the order.

4.5. Dipsacaceae — the Teasel Family (Included in Caprifoliaceae)

Dipsacaceae — the teasel family — is a small family of the order Dipsacales, in the APG III–IV system included in Caprifoliaceae s.l. (Mayer, 2016; APG III, 2009; cited in Singh, 2019). The family includes 14 genera and about 300 species, distributed mainly in the Mediterranean, Irano-Turanian region, less often in Central and Southern Africa, Central and Eastern Asia (Mayer, 2016). Morphologically, Dipsacaceae are predominantly herbaceous plants, less often subshrubs and shrubs. They are characterized by opposite leaves, often pinnately dissected, and capitulum-like inflorescences — capitula, surrounded by an involucre of bracts. Flowers are usually 4- or 5-merous, with an epicalyx — a cup-shaped structure of fused bracts surrounding the ovary. The calyx is often reduced to bristles or teeth. The corolla is sympetalous, often zygomorphic. The fruit is a cypsela, enclosed in the epicalyx, often with a pappus of bristles (Mayer, 2016).

The most economically important genera are Dipsacus — teasel. D. sativus — cultivated teasel, or fuller’s teasel, a biennial herbaceous plant cultivated for its inflorescence heads, which were used for fulling cloth (hence the name). D. fullonum — fuller’s teasel, also used for fulling. D. sylvestris — wild teasel, a wild plant, medicinal and honey-bearing. D. asper — rough teasel, or xu duan, a Chinese medicinal plant whose roots are used to treat lower back pain, fractures, and osteoporosis (Mayer, 2016).

Scabiosa — scabious. It includes more than 30 species, distributed in Europe, the Mediterranean, Eastern Asia, and Africa. Many species are cultivated as ornamentals: S. columbaria — small scabious, S. caucasica — Caucasian scabious, S. atropurpurea — sweet scabious, or Sicalix atropurpurea, popular garden plants. S. columbaria is used in South African folk medicine for colic and heartburn. S. tschiliensis, S. comosa, and other species are ornamental plants (Mayer, 2016).

Cephalaria — cephalaria. It includes about 80 species, distributed in the Mediterranean, Eastern and Southern Africa, Russia, and Persia. C. gigantea — giant cephalaria, a large ornamental plant of the Caucasus, cultivated in gardens. C. syriaca — Syrian cephalaria, a weed and oilseed plant. C. ambrosioides, C. transsilvanica — medicinal plants containing triterpene saponins with antimicrobial activity (Mayer, 2016).

Knautia — knautia. It includes about 60 species, distributed in Europe and Asia Minor. K. arvensis — field scabious, or blue buttons, a perennial herbaceous plant of meadows and forest edges, honey-bearing and ornamental. K. macedonica — Macedonian scabious, a popular ornamental plant with dark purple flowers. K. orientalis — Oriental knautia, an annual plant of the Mediterranean (Mayer, 2016).

Other economically important genera: Succisa — devil’s bit. S. pratensis — devil’s bit scabious, a perennial herbaceous plant of wet meadows, medicinal and honey-bearing. Succisella — succisella, an ornamental and medicinal plant. Pterocephalus — pterocephalus, ornamental and medicinal plants. Lomelosia — lomelosia, ornamental plants, some species used in folk medicine. Pycnocomon — pycnocomon, ornamental and medicinal plants of the Mediterranean (Mayer, 2016).

Thus, Dipsacaceae is a small but economically important family included in Caprifoliaceae s.l. Its importance is connected mainly with ornamental (scabious, knautia, cephalaria, lomelosia), medicinal (teasel, scabious, cephalaria), and technical (teasel) plants. Dipsacaceae demonstrate a unique structure of inflorescences and fruits, which makes them an important object for studying the morphology and systematics of Dipsacales.

4.6. Caprifoliaceae s.str. — the Honeysuckle Family

Caprifoliaceae s.str. — the honeysuckle family — is a family of the order Dipsacales, in the narrow sense including 13 genera and about 230 species, distributed mainly in northern temperate regions, less often in the tropics (Hofmann & Bittrich, 2016). In the APG III–IV system, Caprifoliaceae are understood broadly and also include Diervillaceae, Linnaeaceae, Morinaceae, Dipsacaceae, and Valerianaceae (APG III, 2009; cited in Singh, 2019). In this section we consider Caprifoliaceae s.str., that is, honeysuckles in the narrow sense.

Morphologically, Caprifoliaceae s.str. are shrubs, lianas, less often subshrubs and herbaceous plants. They are characterized by opposite leaves, often fused at the bases, without stipules, but sometimes with interpetiolar stipule-like structures. Flowers are usually 5-merous, bisexual, zygomorphic or actinomorphic, with an inferior ovary. The corolla is sympetalous, tubular, bell-shaped, or funnel-shaped, often bilabiate. Stamens 5, less often 4. Fruit a berry, drupe, or capsule (Hofmann & Bittrich, 2016).

The most economically important genera are Lonicera — honeysuckle. It includes about 180 species, distributed in temperate and subtropical regions of the Northern Hemisphere. L. caprifolium — Italian honeysuckle, L. periclymenum — European honeysuckle, L. japonica — Japanese honeysuckle, L. sempervirens — trumpet honeysuckle — ornamental lianas widely cultivated in gardens and parks. L. caerulea — blue honeysuckle, L. edulis — edible honeysuckle, L. kamtschatica — Kamchatka honeysuckle — berry crops whose fruits are rich in vitamins and used fresh and for processing. L. tatarica — Tatarian honeysuckle, an ornamental shrub, honey plant. L. xylosteum — fly honeysuckle, or wolfberry, a poisonous plant, ornamental shrub (Hofmann & Bittrich, 2016).

Symphoricarpos — snowberry. It includes about 15 species, distributed in North America, Mexico, and China. S. albus — common snowberry, or wolfberry, an ornamental shrub with white berries, widely cultivated in gardens and parks. S. orbiculatus — coralberry, an ornamental shrub. The fruits of snowberry are poisonous to humans but serve as food for birds (Hofmann & Bittrich, 2016).

Weigela — weigela. It includes about 10 species, distributed in Eastern Asia. W. florida — flowering weigela, W. floribunda — Japanese weigela, W. praecox — early weigela — ornamental shrubs with bright tubular flowers, widely cultivated in gardens and parks. Weigelas are valued for abundant and long flowering and diversity of flower color (Hofmann & Bittrich, 2016).

Diervilla — bush honeysuckle. It includes 3 species, distributed in eastern North America. D. lonicera — northern bush honeysuckle, D. sessilifolia — southern bush honeysuckle, D. rivularis — mountain bush honeysuckle — ornamental shrubs with yellow flowers, cultivated in gardens. Bush honeysuckles are also used for soil stabilization and as honey plants (Hofmann & Bittrich, 2016).

Linnaea — twinflower. L. borealis — twinflower, an evergreen creeping subshrub with small fragrant pink flowers, distributed in subarctic, subalpine, and cool-temperate regions of the Northern Hemisphere. Twinflower is an ornamental plant used in rock gardens and as a ground cover (Hofmann & Bittrich, 2016).

Kolkwitzia — beautybush. K. amabilis — beautybush, an ornamental shrub of Central and Northern China with pink bell-shaped flowers, widely cultivated in gardens. Dipelta — dipelta, ornamental shrubs of China. Heptacodium — seven-son flower. H. miconioides — seven-son flower, an ornamental shrub or small tree of China with fragrant white flowers, cultivated in gardens. Abelia — abelia, ornamental shrubs of Asia. Zabelia — zabelia, ornamental shrubs of Asia (Hofmann & Bittrich, 2016).

Thus, Caprifoliaceae s.str. is a family whose economic importance is connected mainly with ornamental (honeysuckle, weigela, snowberry, bush honeysuckle, beautybush, abelia) and berry (edible honeysuckle) plants. Some species are used in folk medicine and as honey plants. Caprifoliaceae s.str. is an important component of temperate forest and shrub communities of the Northern Hemisphere.

4.7. Valerianaceae — the Valerian Family (Included in Caprifoliaceae)

Valerianaceae — the valerian family — is a small family of the order Dipsacales, in the APG III–IV system included in Caprifoliaceae s.l. (Weberling & Bittrich, 2016; APG III, 2009; cited in Singh, 2019). The family includes 5 genera and about 400 species, distributed mainly in the Northern Hemisphere and mountain regions of South America; it is absent from Australia and Polynesia (Weberling & Bittrich, 2016). Morphologically, Valerianaceae are predominantly herbaceous plants, less often subshrubs, shrubs, and lianas. They are characterized by opposite leaves, often pinnately dissected, and corymbose, paniculate, or capitate inflorescences. Flowers are usually 5-merous, less often 3-merous, zygomorphic, with an inferior ovary, often with a spur or saccate outgrowth at the base of the corolla. The calyx is often reduced to teeth or bristles, sometimes enlarged into a pappus. Fruit a cypsela, often with a pappus of plumose bristles (Weberling & Bittrich, 2016).

The most economically important genera are Valeriana — valerian. It includes about 270 species, distributed in Eurasia, North and South America, and Africa. V. officinalis — common valerian, or cat’s root, a perennial herbaceous plant of Europe and Asia whose rhizomes and roots contain valepotriates — iridoid compounds with sedative, antispasmodic, and hypnotic action. Valerian is widely used in medicine as a calming remedy. V. wallichii — Indian valerian, or jatamansi, a Himalayan plant used in Ayurvedic medicine. V. hardwickii — Hardwick’s valerian, a medicinal plant of the Himalayas. V. edulis — edible valerian, whose roots were used as food by Native Americans. V. celtica — Celtic nard, or spica nardi, used as a perfume and medicinal plant since antiquity. V. capensis — Cape valerian, a South African medicinal plant. V. scandens — climbing valerian, a liana of tropical America. V. pycnantha, V. globularis, V. rigida, and other high-Andean species form cushion-like forms (Weberling & Bittrich, 2016).

Valerianella — corn salad. It includes 60–70 species, distributed mainly in the Mediterranean, Southwest Asia, Europe, and North America. V. locusta — common corn salad, or field salad, mâche, rapunzel, an annual vegetable plant whose leaves are used fresh as a salad. V. cornucopiae — horned corn salad, a vegetable and ornamental plant. V. eriocarpa, V. dentata, V. carinata — other species used as salad and medicinal plants. Corn salad is an important vegetable crop of the Mediterranean and Western Europe (Weberling & Bittrich, 2016).

Centranthus — centranthus. It includes about 10 species, distributed in Southern Europe, the Mediterranean, and Southwest Asia. C. ruber — red valerian, or red spur valerian, an ornamental plant with bright red flowers, widely cultivated in gardens. C. macrosiphon — long-tubed centranthus, an ornamental plant. C. calcitrapae — spurred centranthus, a medicinal plant of the Mediterranean. Centranthus is an ornamental and medicinal plant (Weberling & Bittrich, 2016).

Patrinia — patrinia. It includes about 25 species, distributed in Central Asia, the Himalayas, and Eastern Asia. P. scabiosifolia — scabious-leaved patrinia, an ornamental and medicinal plant. P. villosa — hairy patrinia, a medicinal plant. P. intermedia, P. heterophylla — medicinal plants containing saponins. Patrinia is used in Chinese and Tibetan medicine (Weberling & Bittrich, 2016).

Nardostachys — nardostachys. N. jatamansi — jatamansi, or spikenard, a perennial herbaceous plant of the Himalayas whose rhizomes contain essential oil with patchoulol, valeranone, and other compounds. Nardostachys has been used as a perfume and medicinal plant since ancient times and is mentioned in the Bible. It is included in CITES Appendix II. It is used in Ayurvedic medicine as a stimulating, tonic, and anticonvulsant remedy (Weberling & Bittrich, 2016).

Thus, Valerianaceae is a small but economically important family included in Caprifoliaceae s.l. Its importance is connected mainly with medicinal (valerian, nardostachys, patrinia), vegetable (corn salad), and ornamental (centranthus, patrinia) plants. Valerianaceae is an important component of temperate and high-mountain floras, especially in the Mediterranean, the Himalayas, and the Andes.

4.8. Conclusion

The related groups of Asteraceae and Apiaceae considered in this chapter demonstrate the diversity of economically important plants within the orders Asterales, Apiales, and Dipsacales. In the order Apiales, the closest relative of Apiaceae is Araliaceae — a family yielding medicinal (ginseng, aralia, eleuthero) and ornamental (ivy, fatsia, schefflera) plants. Other families of Apiales — Pittosporaceae, Myodocarpaceae, Pennantiaceae — have limited economic importance but are important for understanding the phylogeny of the order.

In the order Asterales, the closest relatives of Asteraceae are Campanulaceae — a large family yielding ornamental (bellflowers, lobelias, balloon flower), medicinal (lobelia, balloon flower, codonopsis), and vegetable (rampion bellflower) plants. Goodeniaceae and Menyanthaceae are small families with limited economic importance but important for understanding the evolution of Asterales.

In the order Dipsacales, economically important groups include Dipsacaceae (teasel, scabious, cephalaria, knautia), Caprifoliaceae s.str. (honeysuckle, weigela, snowberry, bush honeysuckle), and Valerianaceae (valerian, corn salad, centranthus, nardostachys). In the APG III–IV system these families are united in Caprifoliaceae s.l., reflecting their close kinship. The economic importance of Dipsacales is connected mainly with ornamental, medicinal, vegetable, and technical plants.

Thus, the study of related groups of Asteraceae and Apiaceae allows one to see that economically important plants are not limited to the two largest families but are distributed across several orders of the Campanulids clade. Understanding the phylogenetic relationships of these groups is necessary for correct identification, breeding, and rational use of plant resources.

5.1. The Capitulum Inflorescence in Asteraceae

5.2. The Umbel Inflorescence in Apiaceae

5.3. Fruits of Asteraceae — the Cypsela

5.4. Fruits of Apiaceae — the Schizocarp

5.5. Comparative Analysis of Inflorescences and Fruits of Asteraceae and Apiaceae

5.6. Use of Inflorescence and Fruit Morphology in Identification Manuals

5.7. Conclusion

The morphology of inflorescences and fruits is a key source of diagnostic characters for the systematics of Asteraceae and Apiaceae. The capitulum of Asteraceae and the compound umbel of Apiaceae are unique structures that allow these families to be easily distinguished. The cypsela of Asteraceae and the schizocarp of Apiaceae carry many characters used to delimit genera and species. Understanding the morphology of inflorescences and fruits is necessary for correct plant identification, breeding, plant protection, and rational use of plant resources. In identification manuals, these characters are used at all levels: from family to species. Knowledge of the morphology of inflorescences and fruits is the basis for working with herbarium material, seed collections, and living collections.

6.1. Two Families — Two Poles of Diversity

In this lecture we considered the two largest and economically important families of flowering plants — Asteraceae (composites) and Apiaceae (umbellifers, celery family) — as well as their closest relatives within the Campanulids clade. Asteraceae is the largest family of flowering plants, including more than 1600 genera and about 23,000 species (Anderberg et al., 2007). Apiaceae is a family considerably smaller in number of species (466 genera and about 3800 species), but with an exceptionally high concentration of food, spice-aromatic, medicinal, and poisonous plants (Reduron, 2021). These two families represent two different evolutionary paths, two different strategies of survival and dispersal.

Asteraceae bet on the pseudanthium — the capitulum functioning as a single flower — on an enormous number of small flowers, on secondary pollen presentation, on a diversity of cypsela fruits with pappi, and on broad biochemical radiation. This allowed them to become the largest family of flowering plants and to occupy almost all terrestrial habitats except Antarctica (Anderberg et al., 2007). Apiaceae took the path of the compound umbel, aromatic secretory canals, the schizocarp with vittae, and a comparatively narrow but very effective food and medicinal specialization. They are not so rich in species, but it is among them that carrot, celery, parsnip, parsley, coriander, dill, fennel, caraway, anise, and many other crops are concentrated (Reduron, 2021).

6.2. Phylogenetic Position and Related Groups

In the APG IV system (2016), both Asteraceae and Apiaceae belong to the clade Asterids, but belong to different orders: Asteraceae to the order Asterales, Apiaceae to the order Apiales (APG IV, 2016; cited in Singh, 2019). Both orders belong to the informal clade Campanulids. This is a fundamentally important point: the external similarity of umbel-like and capitulum-like inflorescences does not mean close kinship. Before us are two different evolutionary lines that independently achieved high economic importance.

The order Asterales includes, besides Asteraceae, such families as Campanulaceae — the bellflowers, including lobelias (Lammers, 2007), Goodeniaceae — the goodenia family (Carolin, 2007), Menyanthaceae — the buckbean family, Calyceraceae — the calycera family, and a number of others. Campanulaceae is a large family yielding ornamental (bellflowers, lobelias, balloon flower), medicinal (lobelia, balloon flower, codonopsis), and vegetable (rampion bellflower) plants. Goodeniaceae and Menyanthaceae are small families with limited economic importance but important for understanding the evolution of Asterales (Lammers, 2007; Carolin, 2007).

The order Apiales includes, besides Apiaceae, Araliaceae — the aralia family (ginseng, aralia, eleuthero, ivy), Pittosporaceae — the pittosporum family, Myodocarpaceae, Pennantiaceae, and some other groups (APG IV, 2016; cited in Singh, 2019). Araliaceae is the closest relative of Apiaceae, a family yielding medicinal (ginseng, aralia, eleuthero, devil’s club) and ornamental (ivy, fatsia, schefflera) plants. The remaining families of Apiales have limited economic importance but are important for understanding the phylogeny of the order (Singh, 2019).

The same large branch Campanulids also includes the order Dipsacales, in its modern broad sense including Caprifoliaceae s.l., into which in APG III–IV were also included Dipsacaceae — the teasel family — and Valerianaceae — the valerian family (Mayer, 2016; Hofmann & Bittrich, 2016; Weberling & Bittrich, 2016). Dipsacaceae yield ornamental (scabious, knautia, cephalaria), medicinal (teasel, scabious), and technical (teasel) plants. Caprifoliaceae s.str. — ornamental (honeysuckle, weigela, snowberry, bush honeysuckle) and berry (edible honeysuckle) plants. Valerianaceae — medicinal (valerian, nardostachys, patrinia), vegetable (corn salad), and ornamental (centranthus, patrinia) plants (Mayer, 2016; Hofmann & Bittrich, 2016; Weberling & Bittrich, 2016).

6.3. Morphology of Inflorescences and Fruits — the Key to Systematics

The morphology of inflorescences and fruits is a key source of diagnostic characters for the systematics of Asteraceae and Apiaceae. The capitulum of Asteraceae and the compound umbel of Apiaceae are unique structures that allow these families to be easily distinguished. The cypsela of Asteraceae and the schizocarp of Apiaceae carry many characters used to delimit genera and species (Anderberg et al., 2007; Reduron, 2021).

In identification manuals, these characters are used at all levels: from family to species. For Asteraceae, the key characters are the type of capitulum, number of flowers, presence and shape of paleae, number of rows of phyllaries, type of pappus, and shape of cypsela. For Apiaceae — the type of umbel, number of rays, presence and shape of involucre and involucel, shape of schizocarp, number and shape of ribs, and number and arrangement of vittae. Understanding the morphology of inflorescences and fruits is necessary for correct plant identification, breeding, plant protection, and rational use of plant resources (Anderberg et al., 2007; Reduron, 2021).

6.4. Economic Importance

The economic importance of Asteraceae and Apiaceae is exceptionally great and diverse. Asteraceae yield the most important oilseed (sunflower, safflower, niger seed), vegetable (lettuce, artichoke, cardoon, chicory, endive, black salsify, salsify, Jerusalem artichoke, yacón), medicinal (calendula, chamomile, yarrow, wormwood, echinacea, dandelion, beggarticks, burdock, milk thistle, stevia), essential-oil and spice-aromatic (tarragon, wormwood, tansy, chamomile), insecticidal (pyrethrum), ornamental (asters, chrysanthemums, dahlias, marigolds, zinnias, gerberas, gazanias, daisies, calendula, yarrow, echinacea, black-eyed Susan, goldenrod, gaillardia, cosmos), fodder (sunflower, Jerusalem artichoke, fodder varieties of lettuce, chicory, dandelion, yarrow), weed (thistle, sow thistle, plumeless thistle, knapweed, Russian knapweed, cotton thistle, rush skeletonweed, ragweed, cocklebur, groundsel, dandelion), and toxic (groundsel, cocklebur) plants (Anderberg et al., 2007).

Apiaceae yield the most important root vegetables (carrot, root celery, parsnip, root parsley, alexanders, great earthnut, earth chestnut, skirret, turnip-rooted chervil, arracacha), leafy vegetables and spice-aromatic plants (leaf and petiole celery, leaf parsley, chervil, cilantro, dill, vegetable fennel, lovage, angelica, sweet cicely, alexanders, sawtooth coriander), spice-aromatic plants (caraway, coriander, dill, fennel, anise, cumin, ajwain, zira, celery, parsley), medicinal (anise, caraway, fennel, coriander, dill, lovage, angelica, bishop’s weed, Asiatic pennywort, bupleurum, Phloidocarpus sibiricus, Visnaga daucoides, Dorema ammoniacum, Ferula), essential-oil (anise, caraway, fennel, coriander, dill, lovage, angelica, ammi, ferula), poisonous (poison hemlock, hemlock water dropwort, water hemlock, hogweeds, wild parsnip), weed (ground elder, Sosnowsky’s hogweed, poison hemlock, water dropwort, water hemlock, wild parsnip, wild carrot, wild chervil, cow parsley), and ornamental (angelica, astrantia, bupleurum, sea holly, hogweed, hare’s ear) plants (Reduron, 2021).

6.5. Importance of Systematics for the Agrosphere

For a specialist in the agrosphere, knowledge of the systematics of these families is necessary for several reasons. First, it allows one to correctly identify species, genus, and family, which is critical for phytosanitary control, quarantine, and plant protection. Second, understanding phylogenetic relationships helps predict potential pests and diseases, as well as select related species for breeding. Third, the morphology of inflorescences and fruits — the capitulum and cypsela in Asteraceae, the compound umbel and schizocarp in Apiaceae — underlies identification manuals and diagnostic keys. Fourth, knowledge of systematics is necessary for rational use of plant resources, conservation of biodiversity, and sustainable development of agriculture (Anderberg et al., 2007; Reduron, 2021).

Of particular importance is the ability to distinguish cultivated and medicinal umbellifers from poisonous ones. Poison hemlock (Conium maculatum), hemlock water dropwort (Oenanthe crocata), and water hemlock (Cicuta spp.) are deadly poisonous plants that are often confused with parsley, celery, and dill, which leads to severe poisoning. Accurate identification based on morphological characters is the only reliable way to avoid mistakes (Reduron, 2021).

6.6. Prospects for Further Study

The systematics of Asteraceae and Apiaceae continues to develop. Molecular-phylogenetic studies refine the boundaries of genera and subfamilies, reveal cases of paraphyly and polyphyly, and propose new classifications. For Asteraceae, the phylogeny and evolution of subfamilies, especially Asteroideae and Cichorioideae, are actively studied, and the boundaries of large genera (Senecio, Artemisia, Helianthus, Centaurea, Cirsium) are being refined. For Apiaceae, study of the phylogeny of the subfamilies Apiodeae, Saniculoideae, Azorelloideae, and Mackinlayoideae continues, and the boundaries of genera (Daucus, Apium, Pastinaca, Ferula, Heracleum, Bupleurum) are being refined (Anderberg et al., 2007; Reduron, 2021).

Modern research includes not only morphology and molecular phylogeny but also genomics, transcriptomics, and metabolomics, which opens new possibilities for understanding evolution, breeding, and rational use of these economically important families. Knowledge of systematics is the basis for working with genetic resources, breeding material, herbarium collections, and databases (Anderberg et al., 2007; Reduron, 2021).

6.7. Summary Statements

1. Asteraceae is the largest family of flowering plants (more than 1600 genera, about 23,000 species), distinguished by the unique capitulum inflorescence, secondary pollen presentation, an inferior ovary with one ovule, and a cypsela fruit, often with a pappus. The family includes five main subfamilies: Barnadesioideae, Mutisioideae, Carduoideae, Cichorioideae, and Asteroideae. Among Asteraceae are the most important oilseed, vegetable, medicinal, essential-oil, insecticidal, ornamental, fodder, and weed plants (Anderberg et al., 2007).

2. Apiaceae is a large and economically important family (466 genera, about 3800 species), distinguished by the compound umbel, schizocarp, secretory canals with essential oils, and the characteristic structure of fruits with ribs and vittae. The family includes four subfamilies: Apiodeae, Saniculoideae, Azorelloideae, and Mackinlayoideae. Among Apiaceae are the most important root, vegetable, spice-aromatic, medicinal, essential-oil, poisonous, and weed plants (Reduron, 2021).

3. Related groups of Asteraceae and Apiaceae include Araliaceae (ginseng, aralia, ivy), Campanulaceae (bellflowers, lobelias, balloon flower), Goodeniaceae, Menyanthaceae, Dipsacaceae (teasel, scabious), Caprifoliaceae s.str. (honeysuckle, weigela), and Valerianaceae (valerian, corn salad). These families are important for understanding the phylogeny and evolution of economically important plants (Singh, 2019; Lammers, 2007; Carolin, 2007; Mayer, 2016; Hofmann & Bittrich, 2016; Weberling & Bittrich, 2016).

4. Morphology of inflorescences and fruits is a key source of diagnostic characters for the systematics of Asteraceae and Apiaceae. The capitulum and cypsela in the former, the compound umbel and schizocarp in the latter, are unique structures that allow these families to be easily distinguished and their genera and species identified (Anderberg et al., 2007; Reduron, 2021).

5. Economic importance of Asteraceae and Apiaceae is enormous and diverse. They yield oilseed, vegetable, medicinal, essential-oil, insecticidal, ornamental, fodder, weed, and poisonous plants. Knowledge of the systematics of these families is necessary for identification, breeding, plant protection, and rational use of plant resources (Anderberg et al., 2007; Reduron, 2021).

6.8. Closing Remarks

Asteraceae and Apiaceae are two examples of successful but fundamentally different evolutionary strategies. Asteraceae is the largest family of flowering plants, setting the “quantitative” pole of diversity. Apiaceae is a family of moderate volume, but with exceptional economic value, setting the “qualitative” pole of concentration of useful plants. Both groups belong to the clade Asterids, but belong to different orders — Asterales and Apiales (APG IV, 2016; cited in Singh, 2019). Their study in the module “Systematics of Agricultural Plants” of the discipline “Systematics and Taxonomy of Organisms of the Agrosphere” forms a system of knowledge necessary for working with plant resources, breeding, plant protection, and sustainable development of agriculture.

Knowledge of systematics is not an end in itself but a tool. It allows one to see, behind the external diversity of plants, deep evolutionary connections, to understand patterns of distribution and economic use, and to predict the properties of related species. For a specialist in the agrosphere, systematics is the language in which the entire diversity of cultivated and wild plants is described, and mastery of this language is necessary for professional activity.