Piperaceae – the Pepper family

Introducing Piperaceae

Piperaceae is commonly known as the pepper family.  The name Piperaceae comes from the Sanskrit ‘pippali’ which also gave rise to the Greek ‘peperi’, the Latin ‘piper’ and the English ‘pepper’. Originally this referred to Piper longum, the Indian long pepper. These days, the most well-known member of the family is Piper nigrum, the source of the spices black and white pepper (Purseglove). Don’t get confused with the sweet, chilli and cayenne peppers which are all species of Capsicum, a genus in the family Solanaceae. There is more on the economic and other uses of  Piperaceae plants later on.

The Bigger Picture

Before we look at Piperaceae in detail, it will be helpful to understand where this family fits in the plant kingdom and what are its closest relatives. Starting at the top, Piperaceae are angiosperms, that is, flowering plants, but more than this, they are in one of the groups which are considered to be basal angiosperms. In evolutionary terms, basal angiosperms branched off from other flowering plants at various times before the appearance of either the true dicots, more properly called the eudicots, or the monocots.

Piperales

Under the APG III system of botanical classification, Piperaceae is placed in the order Piperales along with three other families, namely Aristolochiaceae, Hydnoraceae and Saururaceae. The circumscription of Piperales has undergone many changes over the years, see table below, however Piperaceae and Saururaceae have been linked since at least 1964 when, in the update of the Engler system, they were placed in Piperales.

Engler (1964 update)     Chloranthaceae, Lactoridaceae, Piperaceae, Saururaceae

Cronquist (1981)              Chloranthaceae, Piperaceae, Saururaceae

APG I (1998)                     Aristolochiaceae, Lactoridaceae, Piperaceae, Saururaceae

APG II (2003)                  Aristolochiaceae, Hydnoraceae, Lactoridaceae,
                                                 Piperaceae, Saururaceae

APG III (2009)                Aristolochiaceae (including Lactoridaceae), Hydnoraceae,
                                                   Piperaceae, Saururaceae

Family Photos – similarities and differences

Looking at the pictures below, you can see why, at a time when much classification was based on morphology, the families Aristolochiaceae, Lactoridaceae and Hydnoraceae were not grouped with Piperaceae and Saururaceae.

Aristolochiaceae species often have showy flowers with the calyx fused into a tube.

Lactoris fernandeziana, the sole member of family Lactoridaceae, which is now considered to be part of Aristolochiaceae, has small, green flowers.

Hydnoraceae is a small family consisting of 2 genera and 7 species of leafless, achlorophyllous root parasites.

 

 

In contrast, Piperaceae and                                     Saururaceae have spikes of
flowers which lack a perianth
(Heywood et al., 2007).

 Family relationships within Piperales

So what is the evidence that all 5 families should be in the same order, Piperales? It comes mostly from phylogenetic studies. Cronquist, in 1981, along with previous authors, recognised that Hydnoraceae was linked to Aristolochiaceae, so it is perhaps a little surprising that Hydnoraceae was not included in Piperales, along with Aristolochiaceae, in the APG system of 1998.

The relationships between Lactoridaceae, Aristolochiaceae and Hydnoraceae were investigated in 2002. The conclusion was that they were closely related, and were in a sister clade to Piperaceae and Saururaceae, but that there was not enough evidence to decide whether they should be separate families or not (Nickrent, 2002).

A study entitled Phylogenetic Relationships in Aristolochiaceae asserted that Lactoris was not nested within Aristolochiaceae (Kelly and Gonzalez, 2003). A further study found that Lactoris was closely related to Aristolochiaceae but that including this genus in Aristolochiaceae made the family paraphyletic (Wanke, Jaramillo et al., 2007). However, in the APG III system of 2009, Lactoridaceae, which contained the single species Lactoris fernandeziana, was subsumed into Aristolochiaceae (Angiosperm phylogeny website).

Piperaceae or Saururaceae?

Suaruraceae, the Lizard’s Tail family, is a small family consisting of 4 genera and six species of herbs. As stated above, Piperaceae and Saururaceae have long been linked, and this close relationship has been confirmed by a phylogenetic study (Nickrent, 2002). They look similar, as shown by the photographs below, sharing the characteristics of spicate inflorescences, the absence of a perianth and actinomorphic flowers.

 

 

 

 

 

 

 

 

 

So how do you tell them apart? They differ in that plants in Suaruraceae have parietal placentation with several ovules and dry fruits while plants in Piperaceae have a single basal ovule and fleshy berries or drupes (Heywood et al., 2007).

Members of the Family

Now that we have seen where the family Piperaceae fits in the APG system of classification, we can look at the groups of plants within it. The number and circumscription of genera in Piperaceae has varied considerably over the years, with recent changes mostly due to phylogenetic research. In some instances genera have been combined and in others they have been split. The APG III system recognises 5 genera: Piper, Peperomia, Zippelia, Manekia and Verhuellia (Angiosperm phylogeny website).

Say Goodbye to three well-known genera XXX

Sarcorhachis disappeared as a genus in 1996 after its similarities to Manekia were recognised (Bornstein, 1996). The two genera were combined as Manekia, as this name had priority, being published two months earlier than Sarcorhachis (Arias, Posada and Bornstein, 2006).

More recently, there have been changes between APG II and APG III. Firstly, it was shown that Piperanthera was a synonym of Peperomia (Semain, Mathieu et al., 2007). Secondly, genus Macropiper was merged into Piper  (Wanke, Jaramillo et al., 2007). It is now recognised as one of 10 major clades in this huge genus (Jaramillo et al., 2008).

And Hello to a new one √

In contrast, Verhuellia is now considered to be a separate genus and not part of Peperomia as previously thought (Wanke et al., 2007). All species of Verhuellia are prostrate with three to four leaves in a whorl at one side of the stem, a pattern not found in any Peperomia species. Another difference from Peperomia is that Verhuelllia flowers have 3 or 4 stigmas, whereas those of Peperomia only have 1, or, rarely, 2 (Semain et al., 2008). A subsequent morphological, developmental and anatomical study provided further evidence that Verhuellia was indeed a separate genus (Semain et al., 2010).

The sub-families change too

Piperaceae was traditionally subdivided into two subfamilies, Peperomioideae and Piperoideae (Semain et al. 2008). However, a 2006 study confirmed that the two major clades actually consisted of Piper and Peperomia in one, and Zippelia and Manekia in the other (Wanke et al., 2006). Note that this was before Verhuellia became a genus in its own right. Following this, a three sub-family classification was proposed: Verhuellioideae, containing the single genus Verhuellia, Piperoideae containing the genera Piper and Peperomia and Zippelideae containing Zippelia and Manekia (Semain et al. 2008).

Genus by genus

While Piper and Peperomia are two of the largest angiosperm genera, the other three genera in Piperaceae only contain about 10 species between them.

 

 

Zippelia

The placing of Zippelia in Piperaceae was confirmed in 1995 by a study of its floral ontogeny, that is, how it developed. Prior to this it had been placed, at various times, in both Piperaceae and Saururaceae (as Circaeocarpus saururoides), most recently in the latter by Heywood in 1993 . The sole species is Zippelia begoniaefolia, an erect herb found in  southeast Asia (Han-Xing and Tucker, 1995).

 

 

 

Manekia

Manekia had been a monotypic genus, containing just Manekia urbanii, until the species within Sarcorhachis were transferred to it. It is unclear how many species of Sarcorhachis there were (Bornstein, 1996), but new combinations were made for 4 species. Manekia are climbing plants found in Central and South America, the Lesser Antilles and Haiti (Arias, Posada and Bornstein, 2006).

 

Verhuellia

The taxonomy of the genus Verhuellia was revised in 2008 and descriptions published of 3 species. 3 other species were found to be synonyms of Peperomia species, the genus of which Verhuellia was previously considered to be part. The genus contains 3 species of small, perennial, prostrate herbs endemic to Cuba and Hispaniola (Semain et al., 2008).

Piper

Piper is an enormous genus containing over 2000 species (Samain et al., 2008). Species show a variety of habits including herbs, shrubs and climbers.

The photo on the right is of a young plant of P. nigrum growing in the tropical greenhouse. It is a climber and in the wild it could easily grow to 10m, although when cultivated a size of 4m x 1.5m is considered optimum (Purseglove).

 

As this is such a large group, many attempts have been made at infrageneric classification over the years. A molecular phylogenetic study has provided evidence for three main lineages of distinct geographic distributions, namely Neotropics (8 major clades), Asian Tropics (Piper s.s. major clade) and the South Pacific (Macropiper major clade) (Jaramillo et al., 2008).

Peperomia

Peperomia is another huge genus with between 1600 and 1800 species (Semain et al., 2007). Many are epiphytes or succlents and narrowly endemic (Wanke et al., 2007).

 

 

 

 

 

 

 

 

As with Piper, various systems of infrageneric classification have been proposed. The most commonly used is that by Dahlstedt (1900), although there have been later versions. Dahlstedt divided Peperomia into 9 subgenera, 7 sections and 4 subsections. In recent years, work has been carried out to name and typify these (Samain et al., 2007).

How do you recognise a Piperaceae plant?

That’s not an easy question to answer and there isn’t really a ‘typical’ Piperaceae plant as, although many of the most common and well-known species are herbs, this family also contains small trees, shrubs and lianas. To make things even more complicated, this variety of form is seen both between and within individual genera – see genus descriptions below. Having said that, there are a few key features that all members of this family have in common.

One of the most obvious of these is that they bear dense flower spikes ……..

 

 

 

 

 

 

 

 

 

…… of minute, radially symmetrical flowers without either petals or sepals, each with a bract below.

 

 

The leaves of Piperaceae are often soft, fleshy or succulent, of a simple shape and with an entire edge (Heywood et al., 2007) as can be seen in many of the photos above.

Piper or Peperomia?

OK, so you have identified a plant as belonging to Piperaceae and want to know what genus it belongs to. You’ve ruled out Zippelia, Manekia and Verhuellia so need to decide between Piper and Peperomia. What do you look for?

Piper often have swollen nodes and prophylls, which are stipule-like structures (Heywood et al., 2007).

 

 

 

 

 

 

Piper also frequently have leaves with a cordate (heart-shaped) base, as in this photo of a Piper nigrum leaf, below, but, as can be seen in the photos of Piper apiculatum above, this is not always the case.  Many Peperomia have succulent leaves, like this Peperomia clusiifolium in the photo below.

 

How do people use Piperaceae?

Having discovered some of the botany of Piperaceae, we will now consider its ethnobotany.

A Valuable Spice

Pepper has been an important economic crop for more than two millennia and was written about by Theophrastus, (327-287 BC), a Greek philosopher sometimes known as the Father of Botany. It is recorded that the ancient Greeks and Romans valued Piper longum more than Piper nigrum, but in modern times, the use of the Indian long pepper has become restricted to the areas where it grows. Pepper had reached Britain by the time of Ethlered and there was a Pepperers Guild in London from 1180. It was a major part of the spice trade and the wealth of various European cities was founded on it.

 

 

 

With Piper longum,  the whole spike of berries is used. It is picked before the berries ripen and then dried. As mentioned in the introduction, Piper nigrum is the source of both black and white pepper. P. nigram is a native of Southern India and Sri Lanka and requires the sort of wet tropical climate found there. You may wonder how both black and white pepper can be made from the same berry. The answer is quite simple. To get black pepper, the unripe peppercorns are sun-dried causing the outer skin to turn black and wrinkle. For white pepper, the ripe berries, having turned red, are soaked, then the outer covering is rubbed off (Purseglove, 1981).

 

Other uses of Piperaceae species

The leaves of Piper betle are used to wrap slices of betel nuts (Areca catechu), smeared with lime and then chewed. They are a gentle stimulant. They are also used locally for medicinal purposes as are some other species of Piper (Purseglove, 1981).

 

 

 

 

Piper methysticum is used to make the Polynesian national beverage, known as kava. Although not an intoxicant, kava is a narcotic, with sedative, soporific and hypnotic effects (Purseglove, 1981).

 

 

Some species of Peperomia are grown as ornamentals and there is a wealth of information on the internet about how to grow these attractive plants.

 

 

 

 

 

 

 

 

All unattributed photos are by Irene Lucas

References:

The Angiosperm Phylogeny Group (1998). An ordinal classification for the families of flowering plants. Annals of the Missouri Botanical Garden 85 (4): 531–553

Angiosperm Phylogeny Group (2003). An update of the Angiosperm Phylogeny Group classification for the orders and families of flowering plants: APG II. Botanical Journal of the Linnean Society 141 (4): 399-436

Arias, T., Calleias, R. & Bornstein, A., (2006). New Combinations in Manekia, an Earlier Name for Sarcorhachis (Piperaceae). Novon 16 (2): 205-208

Bornstein, A. J., (1996). Proposal to Conserve the Name Sarcorhachis against Manekia (Piperaceae). Taxon 45 (2): 323-324

Cronquist, A. (1981). An Integrated System of Classification of Flowering Plants. New York, Guildford: Columbia University Press

Engler, A., Melchior, H., Werdermann, E. (1964). Syllabus der Pflanzenfamilien edition 12 Berlin-Nikolassee: Gebr. Borntraeger

Han-Xing, L & Tucker, S.C. (1995). Floral Ontogeny of Zippelia begoniaefolia and its Familial Affinity: Saururaceae or Piperaceae? American Journal of Botany 82 (5): 681-689

Heywood, V. H., Brummitt, R.K., Culham, A., Seberg, O. (2007). Flowering plant families of the world.  Richmond: Royal Botanic Gardens, Kew

Jaramillo, M. A., Callejas, R., Davidson, C., Smith, J. F., Stevens, A.C. & Tepe, E. J. (2008). Phylogeny of the Tropical Genus Piper Using ITS and the Chloroplast Intron psbJ-petA. Systematic Botany 33 (4): 647-66 

Kelly, M. K. & Gonzalez, F. (2003). Phylogenetic relationships in Aristolochiaceae. Systematic Botany 28 (2): 236-249

Nickrent, D. L., Blarer, A., Qiu, Y., Soltis, D. E., Soltis, P. S. & Zanis, M. (2002). Molecular data place Hydnoraceae with Aristolochiaceae. American Journal of Botany 89 (11): 1809-1817

Purseglove, J.W., 1981. Spices Vol.1. Longman: New York

Semain, M. S., Mathieu, G.,Vanderschaeve, L., Wanke, S., Neinhuis, C. & Goetghebeur, P. (2007). Nomenclature and typification of subdivisional names in the genus Peperomia {Piperaceae). Taxon 56 (1): 229-236

Samain, M. S., Mathieu, G., Wanke, S., Neinhuis, C & Goetghebeur, P (2008) Verhuellia Revisited-Unravelling Its Intricate Taxonomic History and a New SubfamilialClassification of Piperaceae. Taxon 57 (2): 583-587

Samain, M. S., Vriidaghs, A., Hesse, M., Goetghebeur, P., Jimenez, R., Stoll, A., Neinhuuis, C. & Wanke, S. (2010) Verhuellia is a segregate lineage in Piperaceae: more evidence from flower, fruit and pollen morphology, anatomy and development. Annals of Botany 105 (5): 677-688

Wanke, S., Jaramillo, M. A., Borsch, T., Samain, M. S., Quandt, D., Neinhuis, C. (2007) Evolution of Piperales–matK gene and trnK intron sequence data reveal lineage specific resolution contrast. Molecular Phylogenetics and Evolution 42: 477–497

Stevens, P. F. (2001 onwards). Angiosperm Phylogeny Website. Version 12, July 2012 [and more or less continuously updated since]

Wanke, S., Vanderschaeve, L., Mathieu, G., Neinhuis, C., Goetghebeur, P. & Samian, S. (2007) From Forgotten Taxon to a Missing Link? The Position of the Genus Verhuellia (Piperaceae) Revealed by Molecules. Annals of Botany 99: 1231–1238