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Articles • Hoehnea 51 • 2024 • https://doi.org/10.1590/2236-8906e1072023 linkcopy

Open-access Interactive key for species of Rudgea Salisb. (Rubiaceae) that occur in Brazil with emphasis on domatia classification1

Chave interativa para espécies de Rudgea Salisb. (Rubiaceae) que ocorrem no Brasil com ênfase na classificação de domácias

AuthorshipSCIMAGO INSTITUTIONS RANKINGS

ABSTRACT

Rudgea Salisb. (Palicoureeae, Rubiaceae) comprises about 150 neotropical species, with one of its diversity centers in the Atlantic Rainforest. Species of Rudgea are usually identified by a combination of characteristics, including the presence or absence of a structure called domatia. Interactive keys, alongside the domatia descriptions, could be excellent tools for the identification of the Rudgea species. Therefore, the objective of this study was to create an interactive key to Rudgea and describe its domatia types. The characteristic with the highest differentiation power was the stipule type, whilst the type of domatia was the second most powerful between the species that possessed it. Excluding this work, other studies involving domatia descriptions in this family and genus are rare, therefore, new studies are necessary, since the structure shows a potential for identification and differentiation of other taxa of this family.

Keywords:
mutualism; Rubioideae; taxonomy

RESUMO

Rudgea Salisb. (Palicoureeae, Rubiaceae) compreende cerca de 150 espécies neotropicais, com um de seus centros de diversidade na Mata Atlântica. Espécies de Rudgea são geralmente identificadas por uma combinação de características, incluindo a presença ou ausência de uma estrutura chamada domácia. Chaves interativas, acopladas às descrições das domácias, podem ser ferramentas excelentes para a identificação de espécies de Rudgea. Dessa forma, o objetivo deste trabalho foi construir uma chave interativa para Rudgea e descrever seus tipos de domácias. A característica com maior poder de diferenciação foi o tipo de estípula, enquanto o tipo de domácia foi o segundo dentre as espécies a possuíam. Apesar deste trabalho, são raros os estudos que envolvem domácias, portanto se faz necessário novos estudos, uma vez que esta estrutura mostra potencial para a identificação e diferenciação de outras espécies nessa família.

Palavras-chave:
mutualismo; Rubioideae; taxonomia

Introduction

Rubiaceae is an angiosperm family comprised of around 13 thousand species, classified in 611 genera (Davis et al. 2009, Delprete & Jardim 2012). In Brazil the family has ca. 1,400 species, with approximately 700 endemic, positioned in around 100 genera, being considered the sixth most diverse family in the country (BFG 2015, BFG 2021, Flora e Funga do Brasil 2024 (cont. updated)). The habit of these plants is variable, with the possibility of being shrubs, trees, herbs, lianas or subshrubs (Delprete & Jardim 2012). The main diagnostic characteristics frequently present on Rubiaceae species are opposite leaves with undivided blades, interpetiolar stipules, actinomorphic flowers with gamopetalous corolla, isostemonous stamens and inferior ovary (Delprete & Jardim 2012). Considering the family’s vast importance to the Brazilian Flora, studies involving groups within this Family will aid to understand its diversity and distribution.

Rudgea Salisb. (Palicoureeae) comprises about 150 species, all neotropical (Zappi 2003, Bruniera et al. 2022). Its distribution goes from Mexico to Argentina, with one of its diversity centers in the Atlantic Rainforest of southestern Brazil (Zappi 2003, Bruniera 2015). In Brazil, Rudgea is represented by 70 species, 48 of them being endemic (Bruniera et al. 2015, Flora e Funga do Brasil 2024 (cont. updated)). The characteristics that define the genus are interpetiolar entire or laciniate stipules with appendages, presence of leaf domatia in several species, frequently corniculate white or cream corolla, white and spongy or orange, red or brown juicy fruits and adaxially furrowed seeds (Zappi 2003, Bruniera 2015) (figure 1). Some species of this genus can be easily misidentified when certain parts of the specimen are missing, as seen with Rudgea recurva Müll. Arg. and Rudgea coronata (Vell.) Müll. Arg., differentiated especially by some calyx characteristics and the shape of the fruits (Zappi 2003, Bruniera 2015). Therefore, the use of modern tools for taxon identification and the inclusion of new characteristics can optimize species recognition.

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Figure 1
Examples of the diagnostic characteristic of Rudgea. a. Interpetiolar stipule of R. jasminoides (Cham.) Müll. Arg. subsp. corniculata. b. Domatia of R. gardenioides (Cham.) Müll. Arg. c. Inflorescence of R. coriacea (Spreng.) K.Schum. d. Fruits of R. cornifolia (Kunth.) Standl. Photos by Bruniera, C.P. (a and d) and Marcon, S.F. (b and c).

One of the characteristics used to differentiate Rudgea species is the presence of domatia on the underside of the leaves (Zappi 2003). Domatia are structures that serve as a shelter for small animals, generally linked to mutualistic interactions (Barros 1961, Gonçalves & Lorenzi 2011). The structure is present among Eudicotyledons and it is widely distributed in Rubiaceae (Robbrecht 1988). These structures vary greatly in shape, size, diameter of the opening and trichome occurrence (Robbrecht 1988). In the Rubiaceae, domatia are mostly restricted to the axils of the secondary veins of the leaf blade (Robbrecht 1988). As for Rudgea species, these structures are described in some taxonomic treatments (e.g.Zappi 2003) and in some recently described species (Bruniera et al. 2015, 2022, Torres-Leite et al. 2016, 2018, Taylor & Bruniera 2021), however, there is no standardization for the description of domatia found in this genus. Robbrecht (1988) made a brief revision of the types of domatia found in Rubiaceae, and some of them have been identified in some Rudgea species. The types include cavern, crypt, pit, pocket and tuft of hair, all present in Rubiaceae (figure 2) (Robbrecht 1988). A precise identification of the domatia type, when present, found in each of the Rudgea species can exhibit high taxonomic value and can help with the correct identification of these species.

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Figure 2
Illustration of the types of domatia found in Rubiaceae. a. Cavern domatia. b. Crypt domatia. c. Pit domatia. d. Pocket domatia. e. Tuft of hair domatia. f. Example of an intermediary type domatia, with characteristic of both crypt and pit type. Illustration by Marcon, S.F.

A digital tool that can help with species identification and can, also, be integrated within Flora e Funga do Brasil platform are the interactive identification keys (BFG 2018). Interactive keys can be easily updated, accessed, and maintained throughout the years (Dallwitz et al. 2018). As for the process of identification, these keys give the observer freedom to choose the objects characters in any order and gives the opportunity to skip characteristics that are hard to see with the given material (Dallwitz et al. 2018). Interactive keys can also include field photos, photos of exsiccates and illustrations of the characters, states and specimens within the system, facilitating the comparison between the observed specimen and the options present in the keys data bank (Dallwitz et al. 2018). Given the complete digital and intuitive nature of interactive keys, they are great tools to be integrated into digital data banks facilitating the identification of specimens collected on field, for example, Rudgea species.

The objective of this study was to manufacture an Interactive key for Rudgea species that occur in Brazil, with the intent of incorporating it into the Flora e Funga do Brasil platform. Besides, we also investigated, in detail, the type of domatia within the Brazilian species, as it is rarely described in detail. So, the study of vegetative characters, as the leaf domatia, and the incorporation of this information into the Interactive key, will facilitate the recognition of Rudgea species without flowers or fruits.

Material and methods

List of species and descriptors - The list of species analyzed during this study was derived from Flora e Funga do Brasil (2024, cont. updated). According to this platform, Rudgea has 70 species, however, four of the names are being considerate as synonyms by C.P. Bruniera et al. (in prep.) in the largest revision of the genus, therefore, those species were excluded from the final list. The descriptor list was composed with macroscopic characters, prioritizing those that can be analyzed without the use of equipments or a magnifier, and vegetative characters. The characteristics listed, with their respective states, in brackets, include: presence of domatia (present or absent), type of domatia (pocket, cavern, crypt, pit and/or tuft of hair), type of stipule (laciniate, keeled with appendages, basal appendages without a keel, apical appendages without a keel or covered with appendages), length of the leaf blade, texture of the leaf blade (membranaceous, coriaceous or chartaceous), presence of petiole (present or absent), development of the inflorescence (pedunculate capitate, pedunculate and branched or sessile), shape of the pedunculate and branched inflorescence (corymbiform, pyramidal or cylindrical), length of the inflorescence peduncle, shape of the sepals [without lobe differentiation (lobes truncate) or with lobe differentiation (lobes separate)], size of the sepal remnants in the fruit, length and shape of the corolla (infundibuliform, tubular or campanulate) and color (yellow to orange, red or black or light green to white), and texture of the mature fruit (fleshy, spongy or hardened). The morphological terms used for the descriptors followed the following literatures: Radford (1974) and Gonçalves & Lorenzi (2011), in addition to Robbrecht (1988) for the morphology of the domatia and stipules.

Material analysis - For descriptor data and, mostly, for the domatia identification, digital herbariums, like SpeciesLink (speciesLink network), Reflora virtual herbarium (Reflora - Herbário Virtual) and Jstor (plants.jstor.org) were consulted for high-definition photos of the selected species. In addition, herborized materials from Herbarium of Federal University of São Paulo (HUFSP) were also analyzed. The identification of these specimens was confirmed or modified with the use of relevant literature (Bruniera 2015, Bruniera et al. 2015, Delprete et al. 2005, Jung-Mendaçolli 2007, Taylor et al. 2007, Torres-Leite et al. 2016, 2018, Zappi 2003). After identification, the data selected for the key were acquired via observation and comparison with other specimens of the same species. Taxonomic treatments and floristics studies of brazilian localities were also taken in consideration, extracting the data required from descriptions of the desired species made by other authors (e.g.Woodson et al. 1980, Zappi & Lucas 2001, Zappi 2003, Pereira et al. 2006, Jung-Mendaçolli 2007, Taylor et al. 2007, Oliveira et al. 2014, Zappi et al. 2014, 2017, Bruniera 2015, Bruniera et al. 2015, Junior & Vieira 2015, Taylor 2015, Torres-Leite et al. 2016, 2018, Carmo & Simões 2017, Taylor & Bruniera 2018, Fonseca et al. 2020).

Key construction - The interactive key construction began with the selected character codification in the descriptor list. Each character was assigned a number from 1 to 15 and each of its states was assigned a number from 0 to 5, depending on the number of states (appendix 1). With the codification done, a matrix was created with the characteristics represented by the columns, the species represented by the lines, and the determined states, represented by the intersections between the columns and lines (appendix 2 and 3).

Program used - Using the table described above as a guide, the program Xper2 ver. 2.3.2, free software, available for download in <http://www.infosyslab.fr>;, was utilized for the making of the interactive key. The file was then uploaded and made available on the Xper3 digital database.

Results and Discussion

Sixty-five species of Rudgea were analyzed during this study, with only one, Rudgea nivosa Linden ex Berkeley, not having digitized or physical material available for analysis. According to Bruniera (2015) R. nivosa may refer to the same biological entity named as R. coriacea (Spreng.) K.Schum. or R. parquioides (Cham.) Müll. Arg. based on its original description and illustration, but the type material could not be located. Regarding domatia inclusion in the key, the presence of this structure can reduce around 50% of the species present on the databank. Within the species that have the structure (Table 1), some can present variation in the presence and absence of domatia, data represented by the (Yes/No) on the “Presence” column of table 1. Some individuals can present domatia in some leaves, but not on others, in contrast, other specimens can have domatia in all leaves, but others of the same species will have none. When the type is taken into consideration, each of the different categories is almost equally distributed among the Brazilian species (Table 1). The most frequent types were: pit (50%) (figure 3 c), followed by crypt (40,6%) (figure 3 b), cavern (37,5%) (figure 3 a), tuft of hair (34,3%) (figure 3 e) and pocket (25%) (figure 3 d). Note that the sum of the given percentages exceeds 100%, that is because some species can exhibit more than one type of domatia, and others can present an intermediary type, and for both cases the species were marked on table 1, and deserves further attention and specific studies (figure 3 f). However, the domatia of some specimens could not be classified, as shown by the (?) present in some cells of table 1, due to the lack of material available of analysis, both in the form of physical or digitalized exsiccates. In addition, some species, as R. jasminoides (Cham.) Müll. Arg., R. minor (Cham.) Standl. and R. sessilis (Vell.) Müll. Arg. have more than one subspecies, each of them possessing a different type of domatia, or no domatia at all, which contributes to the percentage of domatia occurrence being higher than 100%.

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Table 1
Rudgea species analyzed that present domatia at least in some individuals (Column 1), presence of domatia (Column 2) and the type of domatia (Columns 3 to 7).

thumbnail of Figure 3
Figure 3
Types of domatia present in the Rudgea a. Cavern type of R. guyanensis (A. Rich.) Sandwith. b. Crypt type of R. loretensis Standl. c. Pit type of R. gardenioides (Cham.) Müll. Arg. d. Pocket type of R. minor (Cham.) Standl. e. Tuft of hair type of R. allophyla Standl. f. Intermediary type of R. cornifolia (Kunth.) Standl. Photos by Marcon, S.F.

Of the characteristics analyzed (appendix 1, 2 and 3), the ones with the highest potential for differentiation between species were: the stipule type, followed by the corolla length and the leaf blade length, therefore, we recommend you start the identification by these descriptors. Although “Presence of domatia” is a good differentiator its type is more descriptive. While the “Presence of domatia” is the fourth strongest descriptor, the “Type of domatia” is the second strongest. This seems to contradict the claim, made above, that the second strongest descriptor is the corolla length. That is because the “Type of domatia” is a conditional step, meaning it is only available for choosing after you’ve selected the “Present” in the “Presence of domatia” step. Therefore, this descriptor’s strength cannot be compared to the other non-conditional characters on the key, which results in the “Type of domatia” only being the second strongest descriptor for the species that have said structure. These statistics show that the type of domatia is important for the correct identification of these species. Additionally, species like R. jasminoides, have a clear difference between the domatia of each subspecies, as stated above. This information shows that this characteristic can also be used for subspecies identification. In the final key, we also included definitions and/or descriptions of each characteristic, to facilitate the use of the key. Each species in the databank has its own individual set of characteristics, enabling a more precise identification of any Brazilian Rudgea species. The interactive key is available for public use under the Xper3 digital databank with:

<https://app.xper3.fr:443/xper3GeneratedFiles/publish/identification/7518395130273810344/>.

One advantage to these keys are numerical characters. In this category the observer can input the measurement found in the specimen and the program will automatically exclude the species in which the given number does not fall in between the species size range. As for Rubiaceae, some interactive keys were already produced, like the ones made by Alejandro (2007), for the Rubiaceae genera in the Philippines, and Carmo & Simões (2017), for the Rubiaceae that occurs in the municipality of Camanducaia. However, specifically for Rudgea, any interactive key had been developed prior to this one.

Conclusion

During data collection, it became clear that sorting of domatia into types is not as simple as it appears. Although the types are well described, more often than not, the domatia seen in the specimens available for analysis could be sorted into more than one type. For example, most species classified with the pit or crypt type domatia are dubious and can be seen as intermediate states. This confusion may be cleared with more flexible categories. Besides the difficulties during domatia classification for the inclusion in the key, the interactive key produced during this study is a great tool for easier differentiation of Brazilian Rudgea species. The inclusion of the type of domatia made a significant difference in the separation of species within this genus, indicating that a more precise domatia description can be even more efficient to the identification of Rudgea and the use of this key. Because the interactive key is available at a digital database, it can be constantly updated with new descriptors, new states, include new species, exclude synonyms or reclassified species, increasing the longevity of this study’s results. Descriptive papers regarding domatia are scarce, thus, research into these structures is necessary, including infraspecific variation and if the shape and size of domatia can be linked to environmental variation. New data can also help improve identification and description of new Rubiaceae taxa, since domatia are widely distributed in this family.

  • 1
    Part of the Course Completion Work of the first Author

Acknowledgements

The authors are thankful to Prof. Dr. Marcelo José Sturaro, Dr. João Afonso Martins do Carmo and Prof. Dr. Cristiano Feldens Schwertner, for contributing with the revision of former versions of this manuscript; to Universidade Federal de São Paulo, campus Diadema, for the permission to use the institution’s Virtual Private Network during the coronavirus pandemic, for without it, this study would not be possible; and to HUFSP herbarium, for allowing the use of herbarium specimens and infrastructure for data analysis.

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Appendix 1
Table of codified descriptors (Column 1) and its respective codified states (Column 2).
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Appendix 2
Characters (descriptors) 1-8 used to construct the interactive key and the respective state for each species.
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Appendix 3
Characters (descriptors) 9-15 used to construct the interactive key and the respective state for each species.

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Edited by

  • Editor Associado:
    Renata Sebastiani

Publication Dates

  • Publication in this collection
    16 Dec 2024
  • Date of issue
    2024

History

  • Received
    24 Nov 2023
  • Accepted
    22 May 2024
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