The Flora of Paraíba State , Brazil : subfamilies Ajugoideae and Viticoideae ( Lamiaceae )

The Lamiaceae family includes about 7,000 species allocated in 236 genera, with almost cosmopolitan distribution, currently divided into nine subfamilies. In Brazil 46 genera and 525 species occur, distributed in four subfamilies. Among these subfamilies, Ajugoideae and Viticoideae were originally parts of the Verbenaceae family, being transferred to Lamiaceae as a result of several systematic studies on the two families. Ajugoideae is characterized by having drupes with four mericarps, non-persistent styles and pollen grains with exine formed by granular and branched columns. On the other hand, Viticoideae is characterized to have drupes with 4-locular pyrenes or 2 2-locular pyrenes and exine microreticulate or slightly rough. We present here the results of a taxonomic survey of the subfamilies Ajugoideae and Viticoideae (Lamiaceae) for Paraíba State, Brazil, based on excursions to collect fertile material, analyses of specimens incorporated into herbaria in Paraíba (ACAM, CSTR, EAN and JPB) and Pernambuco State (IPA and PEUFR), and consultations of the SpeciesLink and Herbário Virtual REFLORA databases. Identifications were based on the specialized literature and the examination of type and protolog specimens. Ten species belonging to three genera, five belonging to the subfamily Ajugoideae (Aegiphila integrifolia (Jacq.) Moldenke, A. luschnathii Schauer, A. pernambucensis Moldenke, A. verticillata Vell. and Amasonia campestris (Aubl.) Moldenke) and five to the subfamily Viticoideae (Vitex gardneriana Schauer, V. megapotamica (Spreng.) Moldenke, V. orinocensis Kunth, V. rufescens A. Juss. and V. schaueriana Moldenke) were recorded. The taxonomic treatment includes a key to identify genera and species, descriptions, illustrations, taxonomic comments, geographic distribution, habitats and phenological data of the species.


Introduction
Lamiaceae Martinov is the largest family of the order Lamiales, comprising approximately 7200 species subordinated to 236 genera occurring in tropical and temperate regions throughout the world (Harley et al. 2004).A total of 46 genera and 525 species are known to be native to Brazil, although those numbers are almost certainly underestimations (BFG 2015).
Numerous morphological, chemical, and phylogenetic studies have had enormous impacts on the classification of the family, being possible to conclude that Lamiaceae would be a monophyletic group only with the transfer of about 50 genera were traditionally belonging to Verbenaceae (Cantino et al. 1992, Yuan et al. 2010).
Ajugoideae (Teucrioideae) contains 24 genera of woody or herbaceous plants, with single (rarely compound), opposite, alternate or spiral leaves and unilabial (rarely bilabial) corolla flowers (Harley et al., 2004).According to Judd et al. (2009), is considered monophyletic and is delimited by the following synapomorphies: drupes with four mericarps, non-persistent styles and pollen grains with exine formed by granular and branched columns.
Viticoideae includes 10 genera of arboreal, shrub and lianescent habit (rarely herbaceous), with single or composite, decussate or spiral leaves and frequently bilabial corolla flowers (rarely unilabiate) (Harley et al., 2004).This subfamily is considered as a polyphyletic group and is characterized by having drupes with a 4-locular pyrenes or two 2-locular pyrenes and pollen grains with microreticulated or slightly rough exine (Judd et al., 2009).Bentham (1832Bentham ( -1836Bentham ( , 1848Bentham ( , 1876) ) was the first author to undertake floristic/taxonomic studies of the family Lamiaceae, describing various genera and species, although the principal work focusing on representatives of that family in Brazil is still the Flora Brasiliensis produced by Schmidt (1858).
Some taxonomic studies on the family were carried out in the country, as Harley (2007), for the Ducke Reserve; Harley (2012), for the Brazilian Amazon; Silva-Luz et al. (2012), for the Serra do Cipó, Minas Gerais; Harley (2016), for Serra do Carajás, Pará and Mota et al. (2017), for Serra Negra, Minas Gerais.For the Brazilian northeast, the floristic and taxonomic information about Lamiaceae are scarce, and it is limited to records of new species, evidencing the importance of new studies in the region to contribute to the knowledge of the wealth and distribution of this family (Almeida & Albuquerque 2002;Soares et al. al. 2017).For the Paraiba state, in addition to mentions to their representatives in floristic lists, an annotated list of the species of Lamiaceae was produced by Rocha & Agra (2001), with a record of 23 species and 10 genera.
For the subfamily Ajugoideae in Brazil, studies that deal with the taxonomy of the group are scarce and there is only the study of Santos et al. (2012) andFrança (2003).For the subfamily Viticoideae the scenario is even more precarious, having only the study of Lima & França (2009).
We present here a taxonomic survey of the subfamilies Ajugoideae and Viticoideae (Lamiaceae) for Paraiba State, in northeastern Brazil, presenting descriptions of native species, data concerning flowering, fruiting and geographic distributions, a list of the materials examined, illustrations, and a key for the identification of the species.
The vegetation of Paraiba state reflects the different environmental conditions found in its distinct geomorphological regions, presenting different vegetation formations (Figure 2): restinga forests, mangroves, sawdust and remnants of Atlantic Forest and in the hinterland, caatinga and enclaves of mountain forests (Carvalho & Carvalho 1985).
The climate is considered semi-arid (BSh), with annual rainfall less than 800 mm, extending throughout the Borborema plateau, where in the Paraíba river valley, the annual rainfall is about 400 mm, considered one of the driest places of Brazil (Alvares et al. 2013).
Fertile specimens (with flowers and/or fruits) were collected.The study was based on fieldwork carried out between May/2016 and June/2017.The collected material was incorporated in the ACAM herbarium at Campus I, Universidade Estadual da Paraíba (UEPB).Complemented analyses were made with specimens housed at various herbaria in northeastern Brazil (CSTR, EAN, IPA, JPB and PEUFR), and with images available from virtual herbaria (B, G, HUEFS, HUNEB, K, MAC, NY, RB, SP, and UFP) (Thiers continuously updated) available on the JSTOR and Species Link platforms.
The terminological designations to the morphological descriptions follow Lawrence (1973), Radford et al. (1974), and Harris & Harris (2001).We present keys to the identifications of genera and species, and illustrations of the diagnostic characteristics of the species.Also included are commentaries concerning taxonomic affinities based on vegetative and reproductive morphological characteristics, information concerning geographic distributions, the environments where the taxa are encountered, and the flowering and/or fruiting of the species.
This study is part of the "Flora of Paraíba" project and seek to update the current list of species occurring in the state since, to date, the only work of the taxonomy of the family is Rocha & Agra (2001), which contains exclusively the list of species found in Paraíba.

Results and Discussion
Ten species belonging to three genera were encountered, five belonging to Lamiaceae subfam.Ajugoideae [ Aegiphila Jacq.comprises about 150 Neotropical species, of which 32 occur in Brazil and four species occur in Paraíba state (França & Giulietti 2012, Flora do Brasil 2020, in constr.).The species is distributed through Ecuador, Bolivia, Paraguay, Argentina, and Brazil (all of its states), being associated with all of the phytogeographical domains, being found in anthropic areas, rupestrian fields, ciliary forests, seasonal forests, ombrophilous forests and Amazonian savannah (BFG 2015, Harley et al. 2015).Encountered flowering in May in the study area, present in environments of Atlantic Forest and anthropic areas.
The taxon is distributed in Bolivia, and in all regions of Brazil, associated with Caatinga, Cerrado, Atlantic Forest, Pantanal, and Amazonian vegetation (BFG 2015, Harley et al. 2015, Salimena & Silva 2009).It was found with flowers in July in areas of restinga and Caatinga enclaves.

Figure 1 .
Figure 1.Map of the study area, Paraiba State, Brazil.
is located in northeastern Brazil, bordering Rio Grande do Norte State to the north; Pernambuco State to the south; the Atlantic Ocean at Ponta do Seixas to the east; and Ceará State to the west.