Abstract
We present a study of the diversity, origin and dispersal syndromes of ruderal plants in an urban area in Central-West Brazil. We recorded 266 species in the urban area of Campo Grande, Mato Grosso do Sul state, distributed into 165 genera and 59 families, of which Fabaceae (42 species), Poaceae (32 species) and Asteraceae (26 species) were the most diverse. The surveyed ruderal flora is dominated by native species from the regional Cerrado flora with abiotic dispersal syndromes, through dry fruits and capsules, and by autochory. We present mechanisms of human dispersal of ruderal plants in the city.
Key words:
invaders; Mato Grosso do Sul; naturalized species; spontaneous plants; weeds
Introduction
Ruderals, from the Greek “ruderes” means “ruin” (Font Quer 1953), are plants that occur in environments highly disturbed by human action and can be native or exotic, which can eventually initiate a process of ecological succession (Moro et al. 2012). Ruderal plants can enhance urban ecosystem services by improving nutrient cycling (Sukopp 2004), creating a more robust and resilient urban ecosystem (Mogîldea & Biță-Nicolae 2024). Such plants play a fundamental role as an alternative food resource for pollinators, which, attracted to urban areas, increase fruit productivity, with economic and environmental benefits (Montagnana & Campos 2020). Ruderal species represent a significant resource for the fauna, such as Hymenoptera, birds, and small animals in the urban environment (Luniak 2008). They also represent a potential human food resource once some species have high nutrient contents (e.g., Sukopp 2004).
However, due to the intense movements among urban areas worldwide, exotic species also successfully access new environments far from their origin, widening their geographic distribution. That, allied to gene flow, diaspore dispersal, favorable microclimate conditions and disturbed soil, have contributed to spreading plants and animals from warmer regions (Sukopp 2008). Furthermore, ruderal plants do not avoid little disturbed environments and under low abiotic stress (Schmidtlein et al. 2012). The urban environment can be seen as a mosaic shaped by a tangle of spaces and several landscapes. Though distinct from a natural ecosystem, it is also considered an ecosystem once it depends on and is supported by natural environmental factors such as climate, substrate, water, vegetation and fauna, related and interdependent, constituting an urban ecosystem (Marcondes 2002).
Globalization and the expansion of urban areas worldwide led to marked changes in habitat availability and species assemblages (Bonthoux et al. 2019), suppressing and homogenizing city biodiversity (Ziller 2001). Therefore, urban centers became quickly occupied by generalist native or exotic weed plants, which propagate themselves rapidly, generating a decline of other species (Yalçınalp & Meral 2019) and, as a consequence, exerting relevant ecological role (Kühn 2006; Del Tredici 2010).
Thus, studies on ruderal species can reveal relevant information about their origin and distribution, favoring the understanding of whether such species threaten the native flora or represent native species in the biome where the urban area is inserted (Del Tredici 2010). Furthermore, ruderal plant species are ideal for improving our understanding of the impact of urban ecosystems and the subsequent response of the ecosystem (Dana et al. 2002; Chen et al. 2014), besides their recovery potential of degraded areas (Ranđelović et al. 2024).
We provide an annotated checklist of the ruderal plants from Campo Grande, Mato Grosso do Sul, Brazil, aiming to answer the following questions: 1) What is the diversity and composition of local ruderal florule? 2) What are the types of fruits and dispersal syndromes that favor their dissemination in the urban environment; and 3) What is the origin and distribution pattern of the surveyed ruderal plants?
Material & Methods
Study location
The survey was carried out in Campo Grande city, Mato Grosso do Sul, for eight days, four hours a day, totaling 32 hours. We separated three distinct environments present in urban environments: sidewalks, central beds, and empty lots (Tab. 1; Figs. 1; 2a-m).
The climate is tropical seasonal (Aw), with annual rainfall around 1,500 mm. The predominant soil type is red Latosol, often with earthmoving, landfill, or topsoil removal, compacted by machinery and human trampling. Fertilizers and dog urine add nutrients, and empty plots may receive domestic and construction dumps.
Sampling and data processing
We sampled using the wandering technique and collected fertile specimens in the study areas (Filgueiras et al. 1994) We botanized and identified the plants consulting weed manuals (Kissmann 1991; Kissmann & Groth 1992; Kissmann & Groth 1995; Neto et al. 2010) and websites such as Flora do Brasil and Kew, and deposited them in the CGMS Herbarium, Biosciences Institute of the Federal University of Mato Grosso do Sul (INBio-UFMS). When a species had already been collected in a ruderal area in Campo Grande with complete material deposited in the CGMS herbarium, the Curator of CGMS herbarium opted to not deposit the material again. We analyzed the specimen already deposited in the herbarium CGMS and verified the identification. This material was considered as part of the research when collected in an urbain place in Campo Grande municipality. Data on the collected material can be consulted at the website Jabot (JBRJ continuously updated).
After storing the material in the CGMS Herbarium, we entered the data into a spreadsheet and searched in databases for information on their morphology, vulnerability, occurrences and origin in speciesLink (speciesLink continuously updated), Flora e Funga do Brasil (Flora e Funga do Brasil continuously updated) and Plants of the World Online (POWO continuously updated). The dispersal syndromes were classified according to van der Pijl (1989) into six groups: autochory (explosive dispersal of diaspores or by gravity), anemochory (wind-blown diaspores), zoochory (endozoochory: diaspore dispersal through the animal digestive tract); epizoochory (diaspore dispersed outside the animal) and hydrochory (water dispersal). Although craspedium and lomentum are pod variations, we considered them distinct since their dispersal mechanism differs.
We used the following distribution patterns: American (species with occurrence in South, Central, and North America that reached temperate regions of the continent), South American (species exclusive to South America), Neotropical (species with occurrence in tropical America), African (species native to Africa) and Asian (species native to Asia).
Results
Floristics
We recorded 266 species distributed in 165 genera and 59 families of Angiosperm species (Tab. S1, available on supplementary material <https://doi.org/10.6084/m9.figshare.31126753>). Eudicots (79.1%) were the most collected, followed by Monocots (19.7%) and Magnolids (0.8%). The most diverse families were Fabaceae (41 species), Poaceae (32), Asteraceae (26), Convolvulaceae and Euphorbiaceae (13), Malvaceae (12), Cyperaceae (10), Solanaceae (seven), Amaranthaceae, Cucurbitaceae, Rubiaceae and Verbenaceae (six) (Figs. 3; 4a-m; 5a-m). The three richest families accounted for about 37% of the species, while the ten most diverse families aggregated 62% of diversity within ruderal plants in Campo Grande. On the other hand, although families with one (45%) and two species (16%) represented 61% of the families, these totaled only 17% of the sampled ruderal species diversity. The most diverse genera were Cyperus, Ipomoea, Mimosa (seven species each), Euphorbia, Desmodium (six), Solanum (five), Chamaecrista, Eragrostis, Ludwigia and Sida (four), representing less than 6% of genera, but 20% of species.
Regarding the growth habit, herbs (61%) were predominant, followed by sub-shrubs (18%), shrubs (8%), vines (8%), trees (1.5%), voluble (0.75%) and lianas (0.35%).
Fruits and dispersal syndromes
Dry fruits corresponded to 90% of the diversity in the ruderal flora, in contrast with fleshy structures accounting only for 10% of the species (Fig. 6). Nineteen types of fruits were recorded, with the capsule (34% of the species) the richest, followed by caryopsis (11.8%), cypsela (9.6%) and legume (8.5%), the most representative. Pomid, samara, samarid and siliqua were found in only one species. Furthermore, 54% of the families had capsule fruits, 7% with berries and schizocarps, and 5% with drupes. Families with the highest fruit diversity were Rubiaceae with capsules, schizocarps, and drupes, and Fabaceae with legume, lomentum, and craspedium. Although diverse in number of species and genera, Poaceae (caryopsis), Asteraceae (cypsela) and Fabaceae (legume) had a strongly canalized fruit structure, with their typical and exclusive fruits of each family. Such fruits represented about 30% of the total of ruderal plants.
- Map of Brazil, highlighting the collection points within the limits of the city of Campo Grande, Mato Grosso do Sul, Brazil.
Abiotic mechanisms dominated the diaspore dispersal forms of ruderal flora, accounting for 78% of the species, while biotic processes represented 22%. The most frequent dispersal syndrome in ruderal flora was autochory (63%), followed by anemochory (14%), zoochory (22%), and hydrochory (1%) (Fig. 7a-c). Regarding zoochory, there was a prevalence of endozoochory (12%) over epizoochory (9%). Autochory occurred especially in Fabaceae (19% of the syndrome), Poaceae (10%), Euphorbiaceae (8%) and Convolvulaceae (7%), and this is exclusive in Convolvulaceae, Cyperaceae, Lamiaceae, Loganiaceae, Onagraceae, Oxalidaceae, Sapindaceae and Talinaceae. On the other hand, anemochory was predominant in Asteraceae (52%) and Poaceae (28%), representing 80% of this syndrome in ruderal flora.
- a-m. Ruderal places in the city of Campo Grande, Mato Grosso do Sul, Brazil - a-f. urban environments - a. sidewalk; b. empty lots; c.empty lots; d. central beds; e. empty lots; f. central beds; g-m. examples of occurrence ruderal plants in urban environments.
Zoochory was also concentrated in Fabaceae (18% epizoochory), Solanaceae (10% endozoochory) and Cucurbitaceae (7% endozoochory). Endozoochory was also exclusive to families such as Moraceae, Passifloraceae, and Piperaceae, and epizoochory was exclusive to Caryophyllaceae and Phytolaccaceae. Seeds dispersed by ants (myrmecochory) occurred only in Turneraceae and Polygalaceae.
Within ruderal flora, the capsule was the fruit type that presented all five dispersal syndromes, followed by caryopsis and cypsela with three dispersal forms. Autochory, zoochory and anemochory were the dispersal mechanisms observed in the highest diversity of fruits (Fig. 8). Fleshy fruits, with 83% of the syndrome, were directly linked to endozoochory.
Origin and distribution pattern
The ruderal flora was represented by 72% of native and 28% of exotic species of Brazil (Fig. 9). The diversity found has a predominance of species widely distributed in the Americas, with 78% native to the continent and 71% exclusive to the Americas. The distribution patterns with the highest number of species were American (27% of species), South American (20%), Neotropical (18%), and African-Asiatic (6%), while 12 species were endemic to Brazil.
- Most diverse families and genera of ruderal plants in the city of Campo Grande, Mato Grosso do Sul, Brazil.
Regarding native species from Brazilian Phytogeographic Domains, 66% of the diversity was from Cerrado, followed by the Atlantic Forest (64%). Concerning species exclusive to one of these domains, 50% were from the Cerrado and 37% from the Atlantic Forest. Twenty-five per cent of species occurred in all Brazilian Phytogeographic Domains (Amazon, Caatinga, Cerrado, Atlantic Forest, Pampa and Pantanal); 71% occurred, at least, in four of these domains, and 6% were exclusive to one domain. We highlight that 13% of the ruderal plants from Campo Grande are not mentioned in the Flora do Brasil for Mato Grosso do Sul state. Stachytarpheta sprucei, Malvastrum coromandelianum, Piriqueta carnea, Mimosa chaetosphaera
Fabaceae (33 species), Asteraceae (18), and Poaceae (12) were the families with the highest number of native species; however, they also had more exotics, Poaceae (19 species), Fabaceae (8) and Asteraceae (7) (Fig. 10). In contrast, Cucurbitaceae (100%), Poaceae (60%), and Lamiaceae (50%) were the families with the highest percentage of exotic species. Among the ten most diverse families, Cucurbitaceae presented all exotic species, while Solanaceae, Verbenaceae, Turneraceae, Acanthaceae, and Onagraceae had exclusively native diversity.
The ruderal flora in Campo Grande presents some commercial crop species, such as sesame (Sesamum indicum L.), papaya (Carica papaya L.), watermelon (Citrullus lanatus (Thunb.) Matsum. & Nakai), pumpkin (Cucurbita pepo L.), loofah (Luffa aegyptiaca L.), castor-bean (Ricinus communis L.), basil (Ocimum basilicum L.) and guava (Psidium guajava L.). Ornamental plants are also found, such as king’salad (Cosmos caudatus Kunth), fortune flower (Kalanchoe laetivirens Desc.), begonia (Begonia cucullata Willd.), lemon-balm (Lippia alba (Mill.) N.E. Br. ex Britton & P. Wilson), shamrock (Oxalis latifolia Kunth.), marvel-plant (Mirabilis jalapa L.), and the guarujá flower (Turnera subulata Sm.).
Tithonia diversifolia (Hemsl.) A. Gray, Cenchrus echinatus L., Mimosa caesalpiniaefolia Benth., Urochloa decumbens (Stapf) R.D.Webster are listed on the online platform Brazilian Biodiversity Information System as invasive species (<https://specieslist.sibbr.gov.br>).
Seedlings of urban arborization trees with wind-blown seeds are abundant, e.g., native trumpet trees (Handroanthus Mattos and Tabebuia Gomes ex DC. species), also the exotic Tabebuia rosea (Bertol.) DC., besides the native trees Albizia niopoides (Spruce ex Benth.) Burkart and Astronium urundeuva (F. Allemão) Engl. Seedlings of the exotic tree Leucaena leucocephala (Lam.) de Wit are highly frequent, besides Bauhinia variegata L. and Moquilea tomentosa Benth.
- a-m. Ruderal flora from Campo Grande (MS) Brazil - a. Arachis glabrata; b. Chamaecrista rotundifolia; c. Clitoria ternatea; d. Commelina erecta; e. Cucurbita; f. Desmodium incanum; g. Dichondra sericea; h. Emilia fosbergii; i. Celosia argentea; j. Euploca procumbens; l. Indigofera spicata; m. Ipomoea cairica.
Discussion
Floristics
We present a survey of ruderal plants with a high species richness (264 species) considering other studies of ruderal plants in some Brazilian regions (e.g., Cervi et al. 1988; Carneiro & Irgang 2005; Silva et al. 2008). The richness of those ruderal floras varied from 53 species (Souza et al. 2012) to 344 species (Neto 2010; Neto et al. 2010) (Tab. S2, available on supplementary material <https://doi.org/10.6084/m9.figshare.31126753>). Discrepancies in the diversity of floristic studies are generally linked to the different methodologies adopted, sampling effort, area size and Phytogeographic Domain of the vegetation fragment of the city (e.g., Cervi et al. 1988; Carneiro & Irgang 2005; Silva et al. 2008; Souza et al. 2012). Another influencing factor is the history of disturbance in the area and whether cities maintain forest fragments around (Ratter et al. 2003; Bridgewater et al. 2004).
Although Fabaceae was the most diverse family of ruderal plants of Campo Grande, Asteraceae was the richest in most studies in Brazil, followed by Poaceae and Fabaceae, respectively (e.g.,Carneiro & Irgang 2005; Schneider & Irgang 2005; Hassemer & Trevisan 2012). Asteraceae is mentioned as the most diverse family in 63% of surveys, and Poaceae appears in 27% and Fabaceae in 23% (Tab. S1, available on supplementary material <https://doi.org/10.6084/m9.figshare.31126753>). These three families were highlighted in all national surveys of ruderal flora (Brandão et al. 1995; Silva et al. 2008; Rolim 2013; Neto et al. 2015; Vichiato & Vichiato 2016) and among the four most diverse families in Brazil (Forzza et al. 2010; BFG 2018; Gomes da Silva et al. 2021), representing about 25% of the floristic diversity of the Cerrado Phytogeographic Domain, predominantly from poor and open environments (Sano et al. 2008).
- a-m. Ruderal flora from Campo Grande (MS) Brazil - a. Ipomoea procurrens; b. Ipomoea quamoclit; c. Mollugo verticillata; d. Passiflora pohlii; e. Pectis odorata; f. Plantago tomentosa; g. Portulaca oleracea; h. Portulaca oleracea; i. Richardia grandiflora; j. Tithonia diversifolia; l. Turnera orientalis; m. Turnera subulata.
- a-c. Mechanisms and dispersal syndromes of the ruderal flora at Campo Grande, MS, Brazil.
Fabaceae is the family that most contributes to the richness of the Neotropical flora (Gentry 1988), has the highest richness in Brazil (Forzza et al. 2010; BFG 2018; Gomes da Silva et al. 2021) and is highlighted in the floristic composition of all Phytogeographic Domains (Pott & Pott 1999; Ribeiro et al. 1999; Bridgewater et al. 2004; Felfili et al. 2002; Ratter et al. 2003; Battilani et al. 2005; Sano et al. 2008), including ruderal floras (Gavilanes & D’angieri Soares-Filho et al. 2016; Vichiato & Vichiato 2016). Asteraceae and Poaceae are also among the most diverse families in Brazil (Gomes da Silva et al. 2021) and are relevant components of the invasive species diversity of anthropic areas on account of preference for open environments, rapid growth and efficient dispersal (Funk et al. 2009; Vichiato & Vichiato 2016; Souza & Lorenzi 2019). The ruderal florule in this central area of the state is similar to that found in the eastern part (Neto 2010; Neto et al. 2010), and many species also occur in the western area (Pott & Pott 1985, 1986, 1999).
We point out that Mazaceae, with the tiny Mazus pumilus (Burm. f.) Steenis, had never been collected and mentioned for Mato Grosso do Sul state. The species is from Asia and Oceania, adapting in Brazil to cracks in sidewalks and walls (Souza & Lorenzi 2019).
Fruits and dispersal syndromes
Abiotic dispersal mechanisms in ruderal flora, as found here, are directly related to the high proportion of dry fruits (e.g.,Silva & Rodal 2009; Silva et al. 2013) in agreement with the anemochoric and autochoric syndromes of dispersal (van der Pijl 1982). There is a direct relationship between the proportion of dry fruits and strong seasonality, with the opposite occurring with fleshy fruits (Silva & Rodal 2009). Therefore, dry fruits, especially capsules, are also the most abundant type in the Cerrado Phytogeographic Domain (Peres 2016).
- Types of fruits and their respective dispersal syndromes of the ruderal flora from Campo Grande (MS) Brazil.
Abiotic dispersal mechanisms are common in pioneer plants and increase in importance in more open environments and regions with low rainfall (Gentry 1982; van der Pijl 1982; Peres 2016). Consequently, abiotic syndromes, such as autochory and anemochory, are dominant in the Caatinga (Silva & Rodal 2009; Silva et al. 2013; Lima & Melo 2015), Chaco (Andrella et al. 2023), rupestrian cerrado (Dutra et al. 2009), outcrop rocky areas (Araújo et al. 2008; Costa 2014), deserts (van der Pijl 1982; van Oudtshoorn & van Rooyen 1999; Liu et al. 2014), grasslands (Peres 2016) and in ruderal environments, as we have seen here. For this reason, abiotic dispersal tools are typical of dry fruits (Silva & Rodal 2009; Silva et al. 2013), such as dry fruits are dominant (90%) in ruderal flora.
In contrast, zoochoric dispersal, especially endozoochory, intensifies in more humid vegetation (Gentry 1982), dominating in the forested formations of the Amazon and Atlantic Forests (Yamamoto et al. 2007; Stefanello et al. 2010; Silva et al. 2012) besides in Cerrado riparian forests (Stefanello et al. 2010; Silva et al. 2012; Ramos & Sartori 2013) and restingas (Amaral et al. 2015). The prevalence of endozoochory over epizoochory increased by the occurrence of species with fleshy native and cultivated fruits, i.e., Carica papaya, Citrullus lanatus, Citrus spp., Cucurbita pepo L., Cucumis anguria L., Psidium guajava, and Solanum spp. Endozoochory has birds as the preferential dispersal agent in cities (Soares-Filho et al. 2016; Yang et al. 2021), a more diverse group in the urban environment compared with native mammals (Yang et al. 2021).
The deforested open environment in ruderal areas, the anthropic and wind stresses, and the virtual absence of mammals (except bats, pets, and humans) as dispersers selected species of dry fruits, autochory, and anemochory (Vittoz & Engler 2007), which may be a characteristic that enabled a high number of native species are present. Since autochory does not require biotic dispersers, such species may show an aggregated distribution pattern (Vittoz & Engler 2007).
Although hydrochory in the survey only occurred in Nymphoides humboldtiana (Kunth) Kuntze, this mechanism may also occur in Ipomoea carnea Jacq. due to trichomes on the seeds as a flotation tool (Pott & Pott 2000). Nevertheless, Pilea microphylla (L.) Liebm. abundant tiny seeds are also carried by rain and irrigation water, besides being wind-blown, growing in shaded, moist, rich soil. Five species showed dispersal by zoochory-myrmecochory (Asemeia rhodoptera (Mart. ex A.W. Benn.) J.A. Pastore & J.R. Abbott, A. violacea (Aubl.) J.F.B. Pastore & J.R. Abbott, Turnera orientalis (Urb.) Arbo, T. pumilea L., and T. subulata). Asemeia rhodoptera is restricted to Brazil, perhaps for a dispersal mechanism favorable to its survival in the urban environment (Leal et al. 2015). The dispersal of T. subulata by myrmecochory shows that this type of dispersal is not restricted to small-sized species (Leal et al. 2015). The highly frequent leaf-cutting ants (Atta laevigata (Smith 1858)) can carry such seeds.
Poaceae species, such as Aristida riparia Trin, Digitaria insularis (L.) Fedde, Melinis minutiflora P. Beauv. and M. repens (Willd.) Zizka can be dispersed by both anemochory or epizoochory. This phenomenon results from long periods of contact of the invasive species with agriculture, generating a change in its form of dispersal, originally anemochoric, gradually becoming epizochoric (anthropochoric) by adhesion (Müller 1955).
Important factors in the dispersal of ruderal plant species are the domestication (Zohary & Hopf 2000), the closer relationship between man and wild animals (Povolny 1963; Kenward & Allison 1994), making them cosmopolitan, soil use in the city (Sukopp 2008) and dispersal by residents and domestic animals through epizoochory associated with anemochory (Vittoz & Engler 2007).
Human actions for anthropochory are a direct dispersal agent of ruderal plants, specially through aspects of his daily life. Below, we describe the potential by which people in cities disperse diaspores of ruderal plants by anthropochory.
Fortuitous transport of diaspores (burrs) by human epizoochory, which adhere to clothes and shoes, especially the lower limbs, and objects that come into contact with plants. These diaspores are transported to other places in the city and removed deliberately or left around houses and backyards. Examples: Bidens subalternans DC., Boerhavia difusa L., Cenchrus echinatus L., Desmodium incanum (Sw.) DC., D. tortuosum (Sw.) DC.,, Drymaria cordata (L.) Willd. ex Roem. & Schult., Priva bahiensis A.DC., Triumfetta semitriloba Jacq. and Zornia latifolia Sm. Earthworks favor vegetative propagation of perennial species, e.g., grasses and sedges. Under frequent lawn mowing and human trampling, plants tend to regrow from the base or underground, and their size becomes reduced. Sometimes, fires occur in empty plots overgrown by grass.
- Distribution of exotic (black) and native species (green) in the main botanical families of the ruderal flora of Campo Grande, MS, Brazil.
2) Soil transfer between locations carrying diaspores from seed banks is potentially a dispersal route for all ruderal plant species. Examples: Ipomoea cairica (L.) Sweet, Cyperus rotundus L., Ricinus communis, Solanum sisymbriifolium Lam., Turnera subulata and Stachytarpheta cayennensis (Rich.) Vahl. Besides autochory to the soil seed bank, the pods of Leucaena leucocephala are wind-blown, and the seeds are also carried by water. The propagules of Arachis glabrata Benth. are rhizomes since ir rarelely fructifies, and Cyperus rotundus L. has bulbs and tubers connected by rhizomes, besides seeds. Present in lawns kept short, Cipura paludosa also has bulbs.
3) Public agents weeding, pruning and mowing urban ruderal areas remove excessive vegetation and dropped leaves and transfer diaspores to garbage dumps. Examples: Cyperus esculentus L., Euphorbia heterophylla L., E. hirta L., E. hyssopifolia L. and Tithonia diversifolia.
4) Disposal of residential organic waste with seeds of ruderal food species to inappropriate areas and vacant lots. Examples: Acrocomia totai Mart., Carica papaya, Citrullus lanatus, Cucumis anguria, Cucurbita pepo, Mangifera indica L., Psidium guajava and Syagrus romanzoffiana (Cham.) Glassman.
5) Bagged and potted soil and lawn bought from flower shops and nurseries carry many seedlings and diaspores, which germinate and disperse their seeds, e.g., Cyperus spp.
6) Domestic animals (dogs and cats) help to disperse ruderal species by transporting diaspores attached to their fur.
7) House-to-house transport of diaspores by exchange of ornamental ruderal plants, enabling their dissemination in nearby areas. Examples: Mirabilis jalapa, Petiveria alliacea L. and Lantana fucata Lindl.
8) Dispersal of seeds of potential ruderal plants to feed and attract birds to houses (ornithophily). E.g., Carica papaya, Eugenia uniflora L., Helianthus annuus L., Panicum milliaceum L. (proso millet) and Psidium guajava. Many fleshy and dry fruits feed native birds, such as blue-and yellow macaws.
9) Introduced birds, such as pigeons and sparrows, help to transport diaspores of ruderal plants attached to their body or when they feed on seeds, especially Poaceae.
10) Wind and air displacement by vehicles, besides adherence to wheels, help disseminate anemochoric species in gardens around the streets or central beds. Examples: Aristida riparia, Eragrostis tenuifolia (A.Rich.) Hochst. ex Steud. (Guglieri Caporal et al. 2011), Emilia fosbergii Nicolson, Pectis brevipedunculata (Gardner) Sch.Bip., Porophyllum ruderale (Jacq.) Cass., Tridax procumbens L. and Youngia japonica (L.) DC.
Origin and distribution pattern
Native species accounted for the majority of ruderal surveys, reaching an average of 74% (Tab. S2, available on supplementary material <https://doi.org/10.6084/m9.figshare.31126753>), close to the about 72% found here. However, the percentage (28%) of exotic species here was higher than the national average (23%) (Tab. S2, available on supplementary material <https://doi.org/10.6084/m9.figshare.31126753>). The results of native species are similar to research in Bolivia (80%) and China (69%) (Guo et al. 2018), but different from that found in Chile (20%), Australia (S) (11%) and New Zealand (19%) (Meurk et al. 2016).
Exotic ruderal species (28%) originate mainly from the American continent, followed by Africa and Asia. Human action since the colonization of Brazil has introduced many exotic species through agriculture (Schneider & Irgang 2005). The invasive potential of ruderal plants is remarkable, as these species often thrive in disturbed environments, facilitating their spread into natural ecosystems. Ruderal areas serve as critical entry points for invasive species, which can negatively impact local biodiversity and ecosystem health (Vanessa et al. 2019; Dubyna et al. 2022). Invasive species accounted for 20% of all ruderal species observed in Beijing’s urban areas, indicating their widespread distribution (Guo et al. 2018). A study in Bratislava, Slovakia (i.e., Rendeková et al. 2017) revealed an increase in invasive neophytes over 30 years, indicating a growing trend of invasions in ruderal areas. The results indicated a trend of biodiversity loss and homogenization of plant communities in urban areas, driven by increasing anthropogenic pressures. Therefore, it is important of monitoring alien species dynamics in urban environments to understand ecological changes and inform conservation efforts (Rendeková et al. 2017).
Immigrants from different continents have contributed to the cultural exchange of varied eating habits and the tradition of plant cultivation. Campo Grande is one of the largest Japanese (Okinawa) and Paraguayan communities in Brazil, besides Arabs and Lebanese, and a large number of migrants from the South and Southeast of the country. Indigenous communities and quilombolas also occur in the municipality.
Our data show that the ruderal plants generally present wide distribution, with occurrence records in various Brazilian biomes (Flora e Funga do Brasil, continuously updated). Distinct biomes such as Amazon, Caatinga, Cerrado, Atlantic Forest, Pampa and Pantanal have influenced the flora of Mato Grosso do Sul, and this we also observed in the recorded ruderal native species. The native vegetation areas around the sampled sites, with floristic elements from Cerrado and gallery forest, may have favored the prevalence of native species and the high percentage of species exclusive to the Cerrado. Considered one of the most arborized cities in Brazil, Campo Grande keeps 20% of native vegetation protected in linear parks and conservation units (Planurb 2020). Besides, native species add up to 49% of those used in the landscaping of the city (Pestana et al. 2011), and their planting has been stimulated in urban planning.
We highlight a new species record in our study in the Flora of Brazil. Among the exotic species stands out Mazus pumilus, from the Asian continent, a new occurrence for Mato Grosso do Sul. We point out that the state has one of the lowest collection indices in Brazil (Peixoto 2003; Shepherd 2003; Alves et al. 2018). In the last decades, the official incentive of research policies for cataloguing the Brazilian biodiversity has been reflected in the gradually growing botanical knowledge about the distinct Brazilian regions, e.g., the project Flora and Funga of Brazil (Flora e Funga do Brasil, continuously updated). However, despite all efforts, the extensive territory and megadiversity of the country have hindered the cataloguing of the flora.
In this study, the richest families in ruderal species were Fabaceae, Asteraceae, and Poaceae. The different uses of the species of these families go beyond their food value, such as soil cover (rain infiltration and erosion control), ornamental, medicinal, and religious. Fabaceae and Asteraceae also correspond to the families with the highest number of species among Angiosperms and are well represented worldwide. Besides, these three families have a high reproductive potential, diverse reproductive patterns, and seeds that are fit for wind dispersal, mainly in Asteraceae and Poaceae. Most Fabaceae fix nitrogen and, thus, are favored by their capacity to accumulate nutrients and colonize new areas.
Acknowledgements
This work was supported by the Fundação de Apoio ao Desenvolvimento do Ensino, Ciência e Tecnologia do Estado de Mato Grosso do Sul (FUNDECT) (grant number 71/018.907/2021).
Data availability statement
In accordance with Open Science communication practices, the authors inform that there is no data sharing of this manuscript
Data availability statement
See supplementary material at Figshare
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