Capsicum tovarii

Overview

Capsicum tovarii is a wild, small-fruited, pungent pepper endemic to Peru, best documented from the deep Río Mantaro drainage and the surrounding inter-Andean country. It is not a variety of a familiar supermarket chilli: its narrow geographic range, small seasonal shrub habit, unusual flower structure and molecular evidence support recognition as a distinct species. The Royal Botanic Gardens, Kew, accepts the scientific name and identifies its native range as central Peru in seasonally dry tropical vegetation.

The flowers offer the first surprise. They are often violet or purple with pale cream areas and green markings, but some are nearly cream-coloured. The fruit provides the second: a tiny rounded berry, green when young and red and hot when ripe, on a stalk that changes its orientation as the flower develops. An especially important anatomical character is the mixture of ordinary hairs and minute dark-headed glandular hairs on parts of the plant.

Botanists first formally described the species in 1983, using a preserved collection made by Oscar Tovar in April 1954. Earlier botanical discussions sometimes called the plant Capsicum tovari, but that spelling had not been validly published for the species and tovarii is the established name. Modern DNA research places the species alone in the Tovarii clade, an evolutionary lineage separate from the usual Annuum, Baccatum and Pubescens groups.

It also has clear conservation significance. Kew’s taxon record references an IUCN Red List assessment of Endangered (EN). A separate geographic modelling study published in 2020 classified its model-based risk as Vulnerable and identified major conservation shortfalls. These are different assessments using different evidence, not interchangeable labels.

Accepted name

Capsicum tovarii Eshbaugh, P.G.Sm. & Nickrent

Family / tribe

Solanaceae / Capsiceae

Native range

Central Peru, Mantaro basin

Recorded departments

Ayacucho, Huancavelica, Junín

Evolutionary group

Tovarii clade

Growth habit

Seasonally deciduous shrub/subshrub; often 0.5–1 m

Type gathered

14 April 1954; Oscar Tovar 1867

Original publication

Brittonia 35: 55–60 (1983)

Flowers

Bell-shaped; violet/cream with green marks

Ripe berries

Round, red, pungent; about 4.4–7.6 mm

Seeds

Cream, ear-shaped, approximately 3.5–4.5 mm

Chromosomes

2n = 24 (n = 12)

Type-site elevation

2,000–2,200 m

Conservation

IUCN Endangered (EN), via Kew

Discovery & naming

The name begins with Oscar Tovar Serpa, whose collecting in Peru’s central Andes helped document a distinctive pepper occurring on the steep slopes of the Mantaro valley. On 14 April 1954, Tovar collected specimen 1867 in Huancavelica, Tayacaja Province, in the Mantaro valley between Colcabamba and Surcubamba. His label described a deciduous shrub approximately 0.5–1 m tall, with blue-purple or violet flowers, at 2,000–2,200 metres. This collection would eventually become the holotype, the fixed reference for the species name.

Additional collections followed, including material from 1962 and 1963 in Huancavelica, and plants from Ayacucho in 1971. The specimens were unusual among known peppers. Their flowers were variably cream and purple, their stems formed spreading, sometimes clambering branches, and the plants inhabited a relatively dry Andean landscape instead of the wet forests often associated with rare wild Capsicum.

The chilli had appeared under the spelling Capsicum tovari in Charles Heiser’s 1976 treatment and in several studies from the late 1970s. However, the later authors explained that the 1976 appearance lacked the description or diagnosis needed to establish a species name under botanical nomenclature; they treated it as a nomen nudum, an unpublished or invalidly published name in that technical sense. The correct, formally established spelling is Capsicum tovarii, honouring Tovar.

W. Hardy Eshbaugh, Paul G. Smith and Daniel L. Nickrent published the valid species description in the first 1983 issue of Brittonia, volume 35, pages 55–60, under the title Capsicum tovarii (Solanaceae), a new species of pepper from Peru. They recorded the holotype at USM, the herbarium of the Universidad Nacional Mayor de San Marcos in Lima, with isotypes at MU and US. These herbarium abbreviations indicate institutions holding preserved reference material, not separate types of pepper.

Later research changed the interpretation of the plant’s closest relatives without changing its identity. Field botanists and breeders initially grouped it with various purple-flowered peppers. Controlled crosses in the 1990s suggested another relationship, with C. baccatum. By 2016, a broad DNA study showed that C. tovarii instead occupied a recognizable single-species evolutionary clade. The detailed 2022 monograph retained the species and provided a modern morphological treatment, photographic plate and distribution map.

Plant & foliage

The original field description presents C. tovarii as a perennial, partially woody shrub or subshrub reaching about half a metre to one metre in height. The shoots can be sprawling or clamber through adjacent vegetation rather than forming a neatly compact, self-supporting chilli bush. The plant may lose its leaves seasonally, consistent with its native dry-valley climate. That makes a leafless wild plant during a dry spell a poor basis for declaring it dead.

Branches divide in the characteristic sympodial pattern of peppers. New growth may be green and somewhat angular, while older stems become more woody. Leaves are ovate to lanceolate, tapering toward the pointed tip, with a smooth, toothless margin. The original 1983 measurements for leaf blades were approximately 3.5–8.2 cm long by 1.5–3.8 cm wide, with petioles approximately 1–3 cm long. Younger leaves may look more triangular or trowel-shaped than mature foliage.

Leaf texture and hairs are useful identification features. Eshbaugh and co-authors described the upper leaf surface as nearly smooth or sparsely hairy; beneath, the surface is lightly hairy but can carry longer tufts in the angles where side veins meet the main vein. The 2022 identification key adds a more subtle character: small, dark glandular trichomes mixed with ordinary non-glandular hairs on young stems, leaves and the calyx. These glandular hairs are easy to miss without close inspection or magnification.

The leaves are not uniformly woolly in the way some other wild peppers appear. The original authors specifically noted that hairiness varied among the limited material they examined. Consequently, a relatively smooth-looking individual can still be C. tovarii if its flowers, fruit and diagnostic microscopic traits agree. The overall impression is a fine-branched, seasonally leaf-shedding valley shrub with small, pointed leaves rather than a large, broad-leaved tropical chilli.

Flowers & calyx

The flowers of C. tovarii are unusually variable in colour for a pepper species. The 1983 description gives corollas that are purple and cream or predominantly cream, typically with two green spots near the base of each corolla lobe. The latest botanical photographic plate shows blossoms whose inner surfaces have dark violet-purple patches across a pale cream or yellowish background; the exact distribution of the pigment differs from flower to flower.

In shape, the corolla is campanulate (bell-shaped) or somewhat bell-star-shaped, not broadly flat like a wheel. In the 2022 identification key, its lobes are divided by less than one-third of the total corolla length. The original 1983 account reported a flower width around 4.4–8.5 mm, reflecting the small flowers of the material then measured. Variation in measurement ranges between the original species account and the later monograph should not be mistaken for a change of taxonomic identity.

Inflorescences occur in the leaf axils. The modern key characterises them as relatively small groups of two to eight flowers. A striking change occurs as flowers develop: the flowering pedicels are typically short, roughly 3–10 mm, and hang down, while the stalks supporting berries become upright and green. This contrast between the position of the buds and the mature fruits is especially helpful in identifying a living plant.

The calyx forms a small cup at the flower’s base. Early material described five variably developed teeth, sometimes so short that they are scarcely visible. The 2022 key emphasizes no obvious appendages or five extremely short teeth, generally no more than 0.5 mm, rather than the long projections seen in some other wild species. Under a lens, small glandular hairs may be visible among ordinary hairs on the calyx surface.

Inside, five stamens alternate with the corolla lobes. The style length is not fixed across all flowers, which leads to one of the most unusual aspects of the species: two strongly differing style forms occur, tied to questions about flower function and mating biology. A botanist examining it should therefore inspect several flowers rather than using a single style length as proof of identity.

Reproductive biology: two style forms

The 1983 authors reported flowers that could function as bisexual or effectively unisexual, and noted individuals that appeared to act as male or female at particular times. They treated this as a subject needing further experimental study, not a fully resolved description of a fixed mating system. The distinctions matter: having flowers of different functional sexes is not automatically proof that the species is strictly dioecious, with permanently separate male and female plants.

The mature taxonomic key published in 2022 highlights style dimorphism: a short form approximately 1–1.6 mm long and a long form approximately 4.1–4.2 mm long. The earlier 1983 paper recorded a much broader combined style range, about 1.5–7.6 mm, among its specimens. Both accounts communicate the same essential phenomenon: C. tovarii cannot be described adequately by the single, consistent style shape typical of many familiar garden chillies.

The biological consequences remain important research questions. Differences in style length could influence pollen deposition, the likelihood of self-pollination and the success of particular crosses. However, the older published observations do not by themselves establish the exact frequency of successful self-fertilisation, the genetics of flower forms or a universal pollination mechanism across all wild populations. These questions would require controlled observations of identified plants and repeated flowering seasons.

For anyone maintaining an authenticated living collection, these records have a practical implication: do not assume that one plant, one flower or one pollen donor tells the whole reproductive story. Tracking the flowering stage, which plant supplied pollen, and whether seeds actually germinate is more informative than recording fruit set alone.

Fruit & seeds

The ripe berries are globose, glossy red and distinctly pungent, rather than dark or purple like the flowers. The original protologue reports fruit approximately 4.4–7.6 mm long and 4.2–7.3 mm wide. They begin green, often in noticeable clusters, and turn bright red as they mature. The stalk carrying a berry is usually around 1–2 cm long in the original account, with the fruit generally held upright.

This orientation is taxonomically useful. The flower buds are commonly pendent and short-stalked, yet the ripening berry is presented above its erect stalk. The persistence of the small calyx beneath the berry, and the absence of long calyx spurs, help separate the species from purple-flowered lookalikes. A red berry without flowers is a weaker identification clue than a branch showing both flower and fruit stages.

The original paper describes cream-coloured seeds, about 3.5–4.5 mm long, with an ear-like or curved outline, termed auriform. Under magnification the seed coat shows fine lines to a net-like pattern and minute bumps. Their light colour is notably different from the dark or black seeds of C. pubescens, one of the familiar purple-flowered peppers with which C. tovarii was once grouped.

Despite documented pungency, there is no defensible single Scoville Heat Unit figure for the wild species as a whole in the principal references used for this account. The botanical papers establish that the fruit is hot, but a numerical SHU would need quantitative capsaicinoid measurement from a clearly documented accession. A value quoted for an unverified commercial line should not be presented as a species-wide laboratory measurement.

The berries are very small compared with cultivated chillies, so they have not been developed as an agricultural vegetable pepper. In their native region, however, the combination of strong heat and a small, collectable fruit has made them locally useful as a condiment, an observation already published in the species’ earliest scientific description.

Range & habitat

Capsicum tovarii is a Peruvian endemic, closely tied to the dramatic topography of the Mantaro River basin. The 1983 paper documented collections in the departments of Huancavelica and Ayacucho, particularly around the Tayacaja and Huanta areas. The USDA GRIN distribution adds Junín to its accepted Peruvian departmental records. The 2022 genus monograph and Kew both retain its occurrence in central Peru; there is no verified native record from Bolivia or Ecuador in those taxonomic accounts.

The habitat is notably unlike a wet equatorial mountain forest. Eshbaugh and colleagues described the species as characteristic of a low montane xerophytic zone within the Mantaro basin, with columnar cacti and the tree then referred to as Bombax ruizii among the associated vegetation. In plain terms, this is a seasonally dry inter-Andean valley where hardy shrubs and drought-adapted plants occur on exposed slopes.

The type specimen was collected at 2,000–2,200 m. Other original collections reported sites near 850 m, 1,800 m and 2,000 m. These are documented label elevations, not a claim that the plant occupies every intermediate elevation or that an exact lower or upper ecological limit has been measured. The large elevation differences also show why local slope position, rainfall and season may be as relevant as elevation alone.

One 1971 collection came from agricultural plots and stony scree slopes near Huanchuy, upstream of a bridge across the Mantaro River. Such records are evidence that the plant can occur in or adjacent to disturbed country; they do not establish that the species is a widespread field weed or that its wild populations are secure. Much of the river basin is steep, and small pockets of suitable vegetation can be separated by quite different slopes and land uses.

Documented collecting dates include March, April and May, with flowering explicitly noted in May by the 1983 authors. It would be inaccurate to convert these limited herbarium dates into a strict flowering calendar for every population. Flowering and fruit set in a seasonal valley can be sensitive to rain and to the timing of individual collections.

See the 2022 monograph’s Figure 119 for a published distribution map, and USDA GRIN for the departmental range.

Historical field collections

The original published records make it possible to trace the species long before its formal 1983 description. These are vouchers — identifiable specimens placed in herbaria — rather than an estimate of all populations existing in the countryside. Collector numbers such as 1867 or 4114 are references to gatherings, not seed accession numbers or counts of plants.

Oscar Tovar 1867 — the name-bearing type

Gathered 14 April 1954 near Huaya, between Colcabamba and Surcubamba, Tayacaja Province, Huancavelica, at approximately 2,000–2,200 m. The field note described a deciduous shrub with violet flowers. Deposited as the holotype at USM, with isotypes cited at MU and US.

Additional Huancavelica collections

O. Tovar 4114 was gathered on 21 April 1963 at Quintabamba, Huachocolpa District, Mantaro valley, with an elevation given as about 850 m. The 1983 description also cited a specimen from the valley between Pampas and Salcabamba, dated 16 April 1962 and labelled P. G. Smith accession 2017, at approximately 1,800 m.

Ayacucho material

Barbara Pickersgill 401 was made 28 March 1971 on agricultural ground and rocky slopes near Huanchuy, around eight kilometres upstream from the point where the Huanta road crossed the Mantaro River. The original authors also studied material from the hills north of Huanta, gathered in April 1971, associated with the collection identifier Eshbaugh E-1137.

The authors additionally mentioned a report of material from Andahuaylas, Apurímac, conveyed to them by another researcher. That report is part of the historical discussion, but is not equivalent to a newly verified departmental range in the modern accepted lists. It is better regarded as a historical lead requiring voucher-level verification rather than definitive proof of a current Apurímac population.

Newer scientific photographs were based on Barboza 5044, the specimen credited for the 2022 monograph’s Figure 118. The visual comparison between these modern photographs and the original 1983 botanical drawings is particularly informative: the same species may be represented by cream-dominated or strongly purple-patterned flowers.

Local names & traditional use

The species has a human history beyond its scientific name. The authors of its 1983 description recorded the names mukúru and mucuru-uchu, reporting that local Indigenous people collected the fruits and used them as a spice. The same broad name forms are retained by USDA GRIN as the Quechua names mucuru and mucuru uchu; GRIN also lists the Spanish phrase ají silvestre, meaning wild chilli.

These records document local gathering and consumption, not a wide commercial industry or the existence of a standardized cultivated variety. The labels are useful when looking through older botanical references, but spellings and accents may vary between transcriptions and sources. The generic expression ají silvestre is also not specific enough to identify an unknown plant without its botanical characters.

The red berries are small and pungent, making a condiment use plausible, but the published sources cited here do not establish a detailed traditional recipe, a specific cultural preparation method, a widespread market trade or a quantified harvest rate. It would be misleading to invent any of those details merely because similar wild peppers are used in pickles or sauces elsewhere in South America.

Recognition of local names is important because research centred exclusively on scientific nomenclature can overlook the people who already know and use a plant. At the same time, conservation planning should avoid assuming that local use necessarily threatens the species: the effects of fruit gathering on population regeneration have not been quantified in the sources examined for this account.

DNA, chromosomes & evolution

Capsicum tovarii has 24 chromosomes in its ordinary somatic cells (2n = 24), or 12 chromosomes in its reproductive cells (n = 12). The original 1983 description recorded n = 12, and later cytogenetic studies reported the diploid number. This chromosome count is shared with many cultivated chillies, but having the same chromosome number does not mean that two species are close relatives or freely cross-compatible.

The wider Capsicum genus includes lineages with 24 and 26 chromosomes. In particular, many species of the early-diverging Andean group have 2n = 26. C. tovarii belongs among the 24-chromosome lineages, but its combination of dark-headed glandular hairs, purple-cream flowers and independently supported DNA position separates it from more familiar species.

The major multilocus study by Carrizo García and colleagues (2016) was decisive in resolving a longstanding evolutionary argument. Earlier authors alternately considered C. tovarii related to purple-flowered peppers or to members of the C. baccatum complex. The molecular tree instead placed it in its own monotypic Tovarii clade, near the base of the branch that leads to the Baccatum and Annuum groups. The authors reported limited statistical support for the exact branching order at that point, so its relative position should not be overstated.

The 2022 genus monograph continued to recognise eleven informal evolutionary clades and retained Tovarii as one of them. A later genome-wide RAD-seq study in 2022 further examined chilli diversification through time and space and estimated a branch divergence associated with C. tovarii at approximately 3.3 million years ago. That is a model-dependent evolutionary divergence estimate, not a measured age for a living specimen or a claim that the modern species arose at one exact moment.

Published chromosome studies include evidence of karyotype variation between sampled lines. Such observations are relevant to hybrid fertility and the history of chromosomal rearrangements, but should not be used to invent a new chromosome count: both sampled cytotypes documented in the monograph retain 2n = 24. This is a good example of why chromosomes, crossing tests and DNA sequence data provide different, complementary evidence.

Crossability & pepper breeding

Its experimental crossing history explains why the relationships of C. tovarii were debated. The 1983 authors described early observations of crosses with C. eximium and C. baccatum, including reports of first-generation hybrids from some combinations. They also recorded attempts that produced fruits but failed to yield dependable germination or mature offspring. They explicitly said that later generations needed further study.

In 1999, Tong and Bosland published a dedicated crossability study in Euphytica. They reported that crossing C. baccatum with C. tovarii produced F1 offspring, F2 offspring and backcross progeny in the successful direction investigated. A cross involving the species then known as C. praetermissum also succeeded; that name is now generally treated as C. rabenii.

Microscopic examination of the C. baccatum hybrid’s meiosis showed many chromosomes pairing normally as bivalents, but also abnormalities including multivalents, chromosome bridges and lagging chromosomes. Tong and Bosland interpreted this as evidence that at least one reciprocal chromosome translocation distinguished the parental genomes. Their results supported assigning C. tovarii to the Baccatum breeding complex for crossing purposes.

That does not contradict its recognition as a separate species or require moving it into the Baccatum DNA clade. A breeding complex groups plants by experimental compatibility, while a phylogenetic clade represents inferred evolutionary ancestry. Partial hybridisation can occur across separately named branches, and crossing success may depend on the parent used as female, genetic accession, pollination conditions and subsequent generations.

The potential value to breeding is therefore scientific, not a guaranteed commercial trait package. An authenticated wild accession may introduce genetic diversity into experimental populations, but its small fruit, reproductive variation and unusual chromosome interactions would all require careful evaluation. The existence of a cross in a paper does not mean seed from arbitrary garden plants will reliably produce fertile hybrids.

Identification & related species

A strong identification is based on several characters seen together: a small seasonally deciduous Peruvian shrub; pointed, moderately hairy leaves with tufts in vein junctions; fine dark glandular hairs; short, commonly pendent flower stalks; small bell-shaped purple-and-cream flowers; distinctly different short and long styles; and small upright bright-red pungent berries with cream seeds. No single one of these traits is entirely safe by itself.

Compared with Capsicum cardenasii

Both plants are relatively small Andean peppers with purple-pigmented flowers and glandular hairs, but C. cardenasii is a Bolivian species. In the 2022 key it has five longer calyx appendages, about 1–2 mm, flower pedicels generally 8–18 mm and bent near the blossom, a more consistent style form, and pendent fruiting stalks. In C. tovarii, the calyx teeth are tiny or absent, the flower pedicels are shorter and generally hanging, the styles are dimorphic and fruiting stalks are erect.

Compared with Capsicum eximium

C. eximium is another wild purple-corolla pepper and featured in early crossing experiments. Its flowers can also vary from cream to purple, which makes colour alone an unreliable discriminator. The best approach is to examine the complete floral structure, calyx, stalk orientation, pubescence and geographic source and use the modern species key. Molecular evidence places C. eximium in the Purple Corolla clade, whereas C. tovarii has its own lineage.

Compared with Capsicum pubescens

The familiar C. pubescens (rocoto) also bears striking purple flowers and often hairy leaves, but usually grows as a larger, cultivated or semi-cultivated chilli with substantial fruits and conspicuously dark seeds. C. tovarii has small red berries and pale cream seeds. DNA analyses additionally distinguish the Pubescens and Tovarii clades, despite their superficial floral resemblance.

Compared with Capsicum baccatum and C. rabenii

Controlled crosses link C. tovarii with the Baccatum breeding complex, but C. baccatum usually has a much more openly spreading, pale flower with yellow-green spots instead of the characteristically bell-shaped purple-and-cream form. C. rabenii typically has a purple or lilac corolla margin and noticeable woolly hairs beneath the leaves. Crossability and a 24-chromosome count are not adequate identification tests.

Compared with Capsicum caatingae

In the modern key, the Brazilian species C. caatingae is an especially helpful structural comparison because both can possess tiny dark glandular hairs. But C. caatingae typically carries many more flowers in a cluster, develops more deeply star-shaped corollas and has longer flower stalks with pendent, rather than erect, fruiting stalks. Its Brazilian dryland distribution is also far from the Mantaro valley.

Identification from purchased seed or an internet photograph should be regarded as provisional unless the plant’s origin and a suite of morphological characters can be examined. For a species of conservation concern, an accession traceable to a documented collection is far more useful scientifically than a name alone on a seed packet.

Conservation

Capsicum tovarii is recorded as Endangered (EN) on the IUCN Red List, as linked and reported by Kew’s Plants of the World Online in its taxon information. This is an actual Red List category reflected by the Kew record, not simply an informal assessment made by a website author. The species is geographically restricted, making the condition and connectivity of the Mantaro basin’s suitable dry-slope vegetation particularly important.

A different study by Khoury and colleagues, published in Diversity and Distributions in 2020, modelled the ranges and conservation coverage of wild chilli relatives. Its model-derived risk category for C. tovarii was Vulnerable (VU), not Endangered. That paper’s classification was based on occurrence-derived geographical estimates and did not replace the official IUCN assessment. Both findings are valuable when properly labelled, and their difference is a reason for clearer data rather than a reason to discard one.

The same study ranked the species a high priority for conservation action. Its published dataset identified very limited ex situ representation compared with the diversity predicted in its distribution, and inadequate protection of suitable habitat within established protected-area networks. These results concern geographic coverage of genetic resources and habitat; they are not a direct census of how many wild plants are alive.

Suitable dry inter-Andean vegetation can be affected by agricultural expansion, grazing, roadbuilding, settlement and changes in land management, but the magnitude of each pressure on C. tovarii specifically would need site-level evidence. Its documented occurrence near cultivated land should not be interpreted as proof that it thrives under every type of disturbance. Small, isolated populations may be vulnerable to local changes even while the species persists elsewhere in its valley network.

Priority conservation steps suggested by the evidence include repeat surveys of verified localities, updated georeferenced herbarium records, protection of remaining seasonally dry habitats, and well-documented seed or living-plant accessions maintained with appropriate permission. The USDA GRIN database currently lists the species among its genetic resources, but a database entry does not imply that seeds are commercially purchasable or available to the public.

For clarity, the Endangered listing is a formal taxon risk classification; the 2020 Vulnerable estimate is one research group’s modelling result; and Kew’s separate computer-based plant-risk predictions provide another kind of evidence. They are not three interchangeable descriptions of an identical assessment.

What remains unknown

For all its scientific importance, C. tovarii is much less extensively documented than C. annuum, C. baccatum or C. pubescens. The formal description was based on a limited set of wild material, and much of the better-known experimental literature concerns accessions maintained for crosses. That creates useful questions rather than grounds for filling the gaps with confident-sounding guesses.

Population size and current distribution

A high-quality, current census is needed to determine how many occupied sites remain, how strongly they are connected and which old collection sites can still support the species. The known departmental records and distribution modelling do not constitute an exhaustive survey of every ravine or hillside in central Peru.

Flower function and natural mating

Studies could test the frequency of short- and long-styled flowers, whether individuals change floral function over time, the degree of self-compatibility, which pollinators visit the flowers and whether pollen movement is restricted between isolated stands. The interesting 1983 observations should be investigated rather than over-interpreted as a fully described sexual system.

Seed biology and cultivation

Scientific reports of difficult or inconsistent germination in certain interspecific hybrid crosses do not automatically establish that pure C. tovarii seeds require a specific pretreatment. Published, repeatable germination protocols for authenticated accessions, storage longevity and seasonal dormancy responses would be valuable. A cultivation guide should not invent exact germination temperatures or guaranteed timelines without a tested source.

Fruit chemistry and breeding value

Quantitative measurements of capsaicinoids, aroma components, stress tolerance and disease resistance would help determine which traits are actually present in different natural populations. The species’ occurrence in dry habitats is not proof that a particular accession is drought resistant in commercial agriculture; similarly, its pungency is not a substitute for a species-wide Scoville survey.

The main lesson from C. tovarii is that a rare plant can be important at once as a locally known food resource, a distinct evolutionary lineage, an experimental crossing partner and a conservation priority. The botanical record is unusually rich in questions about relationships and reproductive development, and those questions are best answered by a combination of fieldwork, verified specimens and careful laboratory experiments.