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Fruits, seeds and germination in five species of globose Cacteae (Cactaceae)/ Frutos, semillas y germinacion de cinco especies de Cacteae globosas (Cactaceae)/ Frutos, sementes e germinacao de cinco especies de Cacteae globosas (Cactaceae).

FRUITS, SEEDS AND GERMINATION IN FIVE SPECIES OF GLOBOSE CACTEAE (CACTACEAE)

SUMMARY

The morphological characteristics of fruits and seeds, and the germination responses of freshly matured seeds of five species of Caeteae (Coryphantha bumamma, C. clavata, C. cornifera, Ferocactus histrix and Mammillaria uncinata) were studied at room temperature under laboratory conditions. The aim of the study was to record the macro- and micro-morphology of fruits and seeds of these species and to investigate specific requirements for germination. Variance analysis detected significant differences (p<0.05) for several variables: number of seeds per fruit, weight, and fruit width. Larger fruits with more seeds are observed for F. histrix, whereas smaller fruits with less weight and fewer seeds are seen for C. clavata. Seed germination is a rapid process and usually starts on the third day. High percentages of germination (>80%) are observed on the sixth day in F. histrix and M. uncinata. It is concluded that some morphological characteristics of fruits and seeds can be used to support further systematic studies of Cactoideae genera and will contribute new knowledge for their potential use and conservation.

KEYWORDS / Coryphantha / Ferocactus / Fruits / Germination / Mammillaria / Seeds /

FRUTOS, SEMILLAS Y GERMINACION DE CINCO ESPECIES DE CACTEAE GLOBOSAS (CACTACEAE)

RESUMEN

Las caracteristicas morfologicas de los frutos y semillas, y la respuesta germinativa de semillas maduras y recien cosechadas de cinco especies de Cacteae (Coryphantha bumamma, C. clavata, C. cornifera, Ferocactus histrix, and Mammillaria uncinata) fueron estudiadas bajo condiciones de laboratorio y temperatura ambiente. Se registraron las caracteristicas macro y micromorfologicas de los frutos y semillas de estas especies e investigaron los requerimientos especificos de germinacion. El analisis de varianza mostro diferencias significativas (p<0.05) para algunas variables tales como numero de semi llas por fruto, peso y ancho de fruto. Los frutos mas grandes y con mayor numero de semillas son los de F. histrix, mientras que los frutos mas pequenos y ligeros conel menor numero de semillas son los de C. clavata. La germinacion es un proceso rapido que inicia al tercer dia. El poreentaje de germinacion mas alto (>80%) se observo al sexto dia en E histrix y M. uncinata. Se concluye que algunas caracteristicas morfologicas de los frutos y las semillas pueden apoyar futuros estudios de sistematica en los generos de Cactoideae y que el conocimiento generado contribuira a su uso potencial y conservacion.

FRUTOS, SEMENTES E GERMINACAO DE CINCO ESPECIES DE CACTEAE GLOBOSAS (CACTACEAE)

RESUMO

As caracteristicas morfologicas dos frutos e sementes, e a resposta germinativa de sementes maduras e recem colhidas de cinco especies de Cacteae (Coryphantha bumamma, C. clavata, C. cornifera, Ferocactus histrix, e Mammillaria uncinata) foram estudadas sob condicoes de laboratorio e temperatura ambiente. Registraram-se as caracteristicas macro e miero morfologicas dos frutos e sementes destas especies e investigaram-se os requerimentos especificos de germinacao. A analise de variacao mostrou diferencas significativas (p<0,05) para algumas variaveis, tais como numero de sementes por fruto, peso e largura de fruta. As frutas maiores e com maior numero de sementes sao as de F. histrix, enquanto que os frutos menores e leves com o menor numero de sementes sao as de C. clavata. A germinacao e um processo rapido que inicia ao terceiro dia. A porcentagem de germinacao mais alta (>80%) se observou ao sexto dia em F. histrix e M. uncinata. Conclui-se que algumas caracteristicas morfologicas dos frutos e as sementes podem apolar futuros estudos de sistematica nos generos de Cactoideae e que o conhecimento gerado contribuira a seu uso potencial e conservacao.

Introduction

Cacti are a typical component of arid and semiarid environments in the Western Hemisphere, and the second largest plant family restricted to the New World (Anderson, 2001). Mexico is one of the main centers of diversification of the Cactaceae family (Goettsch and Hernandez, 2006); around 560 species belonging to 50 genera are distributed in this country (Guzman et al., 2003). Indeed, 73% of the genera and 78% of the species are estimated to be endemic to Mexico (Hernandez and Godinez, 1994), and the tribes Cacteae and Echinocereae are almost exclusively distributed in Mexico (Anderson, 2001). Members of Caeteae range from globular or depressed to short columnar cacti, varying in size from dwarf (Turbini carpus and some Mammillaria species) to giant (Ferocactus and Echinocactus) genera. Recently, significant progress has been made in understanding various aspects of the biology, phylogeny, and morpho-anatomy of Cactaceae (Nobel, 2002; Godinez-Alvarez et al., 2003; Terrazas and Arias, 2003; Hernandez-Hernandez et al., 2011). The current understanding of cacti, particularly of Cacteae members, is based on scarce analysis of the morphology, anatomy and composition of stems, fruits and seeds. Some species of this tribe produce edible fruits. For example, Ferocactus histrix produces a small and acidic fruit called 'tuna of biznaga', which is traded in regions of Hidalgo, Queretaro, and San Luis Potosi. The small red fruits of Mammillaria species are generally edible and consumed locally (Bravo-Hollis and Sanchez-Mejorada, 1991). However, most of these species are threatened because they have been removed from their natural populations or their habitat has been modified by changes in land usage (Hernandez-Oria et al., 2007). For several genera of Cactaceae, studies on the morphological variability of fruits and seeds are scarce or have focused exclusively on well-known or economically important genera. For example, the composition, phytochemicals, and nutritious elements have been analyzed in Opuntia (Joubert, 1993; Dominguez-Lopez, 1996; Saenz, 1997; Saenz et al., 1998; Butera et al., 2002; Duru and Turker, 2005; Stintzing et al., 2005), Hylocereus (Wybraniec and Mizrahi, 2002), and Myrtillocactus (Barrera et al., 1998). The study of seed morphology and germination processes is important for supporting systematic studies (Arias and Terrazas, 2004; Arroyo-Cosultchi et al., 2006) and may also help explain patterns of population dynamics of Cactaceae species under field conditions. Studies on seed germination in this plant family have focused on the relationship between germination and different attributes of the plant, such as the effect of seed mass and size on regeneration strategies, mechanisms of reproduction, abundance of rare species, and ecophysiological requirements (Rojas-Arechiga et al., 1997; Rojas-Arechiga and Vazquez-Yanes, 2000; Flores and Briones, 2001; Ayala-Cordero et al., 2004; Ramirez-Padilla and Valverde, 2005; Sanchez-Salas et al., 2006; Jimenez-Aguilar and Flores, 2010; Flores et al., 2011). In the present study, differences in fruit and seed macro- and micro-morphology, and the rates of germination processes were investigated in members of Cacteae that co-exist in the scrub of Jalisco, Mexico.

Materials and Methods

Mature and healthy fruits of Coryphantha bumamma (Ehrenb.) Britton et Rose, C. clavata (Scheidw.) Backeb, C. cornifera (DC.) Lem., Feroeaetus histrix (DC.) G.E. Linds., and Mammillaria uncinata Zucc. ex Pfeiff. were collected from their native populations in Jalisco, Mexico. An analytical scale (Precisa XT 220 A) and a Mitutoyo digital caliper were used to record the size and weight of 30 fruits per species. The fruits were dissected in the laboratory and the seeds were then washed in tap water to eliminate pulp remains and mucilage. The seeds were immediately placed on absorbent paper until they dried and were stored in paper envelopes at room temperature.

The number of seeds per fruit for each species was registered. Two hundred seeds per species were used to record quantitative morphological traits (weight, length, width) and qualitative morphological traits (shape, color, structure of the testa) using an analytical balance and a dissecting microscope Leica Zoom 2000 (Z45V) adapted to an image analyzer (Media Cybernetics, 2006). The roundness index (width/length ratio) was calculated from the length and width measurements. For scanning electron microscopy (SEM), three or four seeds per species were washed using ultrasound and 95% ethanol. Dry seeds were fixed to aluminum specimen holders with double-sided adhesive tape and coated with gold in a JEOL-JFC-1100 sputter coater. Morphological observations and micrographs were carried out with a JEOL-JSM-5310LV field-emission scanning electron microscope. The morphological descriptions of the seeds followed the terminology proposed by Barthlott and Hunt (2000).

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The germination experiments were carried out under laboratory conditions using three replicates of 50 seeds per species. Seeds were disinfected by immersion in a solution of 10% commercial bleach during 5min (Vega-Villasante et al., 1996), washed several times with distilled water, and sown on Whatman No 2 filter paper saturated with 10ml of distilled water in 9cm diameter sterile Petri dishes. The mean temperature in the laboratory was measured using a Data-logger Dickson SP125/175 Pro series thermometer. The mean temperature was 25[degrees]C [+ or -] 2[degrees]C during the day, which has been mentioned to be optimal for seed germination of cacti (Nobel, 1988; Rojas-Arechiga and Vazquez-Yanes, 2000), and 12[degrees]C [+ or -] 3[degrees]C during the night. Germination was recorded at three day intervals for 30 days, when germination ceased. A seed was considered to be germinated when the radicle protruded. Interspecific differences in morphological characters of fruits and seeds were evaluated through variance analyses followed by Tukey's pair-wise means comparison analyses (p<0.05). Differences in the number of seeds per fruit were evaluated through covariance analyses. All analyses were performed with SAS software V 9.1.3 (SAS, 2000).

[FIGURE 2 OMITTED]

Results

Morphology of fruits and seeds

Fruits varied in shape, size and color (Table I, Figure 1). Most were juicy berries, varying in color from pale green with light red at the apex in Coryphantha species to red berries in M. uncinata and yellow-green in F. histrix. The fruits exhibited ovoid to claviform shapes, and only F. histrix showed thick walls and was dehiscent at the base. The smallest fruit (1.4cm length) with the lowest number of seeds (56 seeds/ fruit) occurred in C. clavata and the largest fruit (3.2cm length) with the most seeds (>2000) was observed in F. histrix, the species that also had the heaviest fruits (8 [+ or -] 3g, Table I). The seeds were oval to reniform, small to medium-sized (Table II, Figures 2-4), glossy or matte, and light-brown to brown. The anticlinal boundaries were inconspicuous or raised. The boundaries were straight except in M. uncinata, and the microrelief verrucose.

Interspecific comparison of fruit and seed morphology

The analysis of variance detected significant differences for the number of seeds per fruit (F = 17419, df = 4, p<0.0001), weight (F = 3295.67, df = 4, p<0.0001), length (F = 2191.58, df = 4, p<0.0001) and fruit width (F= 1917.22, df = 4, p<0.0001). F. histrix was characterized by larger and heavier fruits, and more seeds. By contrast, C. clavata and M. uncinata had fruits with fewer seeds and were smaller and lighter in weight than the other species (Table I). Significant differences in seeds among species were observed for weight (F= 3797.76, df= 4, p<0.0001), length (F= 9416.14, df= 4, p<0.0001) and seed width (F= 429.19, df= 4, p<0.0001). C. bumamma seeds presented higher weights and lengths than the lighter and smaller seeds of C. cornifera (Table II).

[FIGURE 3 OMITTED]

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Seed germination response

All species reached 100% germination in the three repetitions per species. Seed germination was a rapid process that started 3-6 days after sowing (DAS). In C. cornifera and C. clavata, germination started on the third day, reaching the highest percentages (24-28%) at 6 and 15 das (Figure 5). Seed germination of C. bumamma began on the sixth DAS, also reaching the highest percentage (24%) 9 DAS (Figure 5). F. histrix and M. uncinata were characterized by higher rates of seed germination (>80%) on the sixth DAS. These two species concluded their germination 12 and 8 DAS respectively. By contrast, C. bumamma concluded its germination 21 DAS with 10% of its seeds germinated in that rime (Figure 5).

[FIGURE 5 OMITTED]

Discussion

Morphology of fruits and seeds

The studied species showed considerable variation in fruit and seed morphology. The Coryphantha species had similarities in fruit morphology with other species of the genus (C. echinoidea Britton & Rose, C. greenwoodii Bravo, C. scheeri (Muehlenpf.) L.D. Benson, C. suleata (Engelm.) Britton & Rose, and C. werdermannii Boed.) (Bravo-Hollis, 1978; Bravo-Hollis and Sanchez-Mejorada, 1991). Characteristics, such as red and claviform fruits of 1.5 x 0.5cm size, in M. uncinata were similar to other Mammillaria species like M. standleyi (Britton & Rose) Orcutt, M. johnstonii (Britton & Rose) Orcutt, M. heyderi Muehlenpf., M. miegiana W.H. Earle, M. ortegae (Britton & Rose) Orcutt, M. pachycylindrica Backeb., M. serippsiana (Britton & Rose) Orcutt and M. magnimamma Haw. (see Bravo-Hollis and Sanchez-Mejorada, 1991). Other Mammillaria species (M. gigantea Hildm. ex K. Schum. and M. melanocentra Poselg) can be distinguished by their claviform fruits, which are pink or purple in color. Noticeable differences were observed in the fruit size (length) of M. uncinata and other species such as M. gigantea. In the latter, the fruits were 2.5-3.0cm long (Bravo-Hollis and Sanchez-Mejorada, 1991), whereas the fruits of M. uncinata were smaller (1.5 [+ or -] 0.3cm).

Cactoideae seeds show considerable variation in their shape, size, structure and testa color (Barthlott and Hunt, 2000). Indeed, the studied species exhibited morphological variability. The seeds of C. bumamma and C. cornifera were curved or reniform, and similar to those of C. durangensis (Runge ex K. Schum) Britton & Rose. C. clavata and M. uncinata had oval seeds, similar to other species of Mammillaria like M. gigantea, M. petterssonii Hildm. and Coryphantha like C. gracilis L. Bremer & A. B. Lau, C. pseudoechinus Boed., C. recurvata (Engelm.) Britton & Rose, C. robustispina (Ant. Schott ex Engelm.) Brittion & Rose (Bravo-Hollis, 1978; Bravo-Hollis and Sanchez-Mejorada, 1991; Barthlott and Hunt, 2000; Dicht and Luthy, 2005). Great variability occurs in the color of the seed testa in Cactaceae. A black or brown color is characteristic in most species, whereas a redor brown color is common in other species, such as C. bumamma and M. uncinata. C. clavata and C. cornifera exhibited dark brown or brown seed testas, which is characteristic of C. duranguensis (Runge ex K. Schum) Britton & Rose, C. pseudonickelsiae Backeb., C. pallida Britton & Rose, C. macromeris (Engelm.) Lem., and C. sulcata (Engelm.) Britton & Rose, according to Bravo-Hollis and Sanchez-Mejorada (1991). Glossy seeds were only observed in C. bumamma, in agreement with seeds of some columnar species (Arias and Terrazas, 2004; Arroyo-Cosultchi et al., 2007). In Cactaceae, studies on the macro- and micro-morphology of the seeds have been successfully applied to different taxonomic levels (Friedrich and Glaetzle, 1983; Glaetzle and Prestle, 1986; Arias and Terrazas, 2004; Arroyo-Cosultchi et al., 2006, 2007). Buxbaum (1958) recognized four subtribes, particularly in the Cacteae tribe, based on seed morphology: Echinocactinae exhibits a perisperm and hard, black seed testas; Thelocactinae has black seed testas and a mostly warty surface; Ferocactinae exhibits dotted or reticulated seed testas; and Coryphanthinae has soft, smooth, and brown testas, similar to the Coryphantha and Mammillaria species studied herein. More recently, Taylor and Clark (1983) recognized that the Ferocactus section includes species with a larger hilum-micropylar region (HMR), dried fruit dehiscing by a basal pole and glossy seed testas. By contrast, the Bisnaga section includes species with seeds whose HMR is narrow with bright seed testas (Taylor and Clark, 1983). Cota and Wallace (1997) used chloroplast DNA evidente to determine that the members of one lineage (Sect. Bisnaga) are related to F. flavovirens (Scheidw.) Britton & Rose and include species distributed mainly in Central Mexico and areas in the putative center of the origin of the genus (Tehuacan Valley). The morphological characteristics of the fruit, seed characteristics and distribution of F. histrix support its inclusion in the Bisnaga section along with other species like F. glaucescens (DC.) Britton & Rose, F. lindsayi Bravo, F. macrodiscus (Mart.) Britton & Rose, F. recurvus (Mill.) Y. Ito ex G. E. Linds., F. latispinus (Haw.) Britton & Rose and F. flavovirens, as suggested by Cota and Wallace (1997).

Potential importance of Cacteae fruits

Most cactus fruits are edible; however, information on their morphology and nutritional value is scarce (Kiesling, 2001; Pardo, 2002). According to Esquivel (2004), most studies have focused on the fruit of prickly pears (Opuntia spp.). Little has been investigated on the composition and aspects of fruit cultivation of other species of cacti, despite the high potential of these fruits for industrial use (Ortega-Nieblas et al., 2001). For example, the presence of red-purple pigments in the fruit epidermis of M. uncinata is ah important source of natural dyes in foods and in various applications (Stintzing et al., 2001; Wybraniec and Mizrhi, 2002; Stintzing and Carle, 2005; Emaldi et al., 2006). Antihyperglycemic and antihyperlipidemic effects found in the fruits of F. latispinus and F. histrix have been demonstrated, suggesting their application in the prevention of diabetes (Perez-Gutierrez and Mota-Flores, 2010). Further studies involving analyses of the nutritional composition of fruits of Cacteae species as well as their morphological characterization may contribute significantly in the search for alternative medicinal products.

Ecological aspects of seed germination response

Differences in the number of seeds per fruit were found between the species studied. A high variability was observed in this trait even within the same genus. For example, C. bumamma and C. cornifera had 84-99 seeds, whereas C. clavata contained 56 seeds per fruit on average. F. histrix was distinguished by larger fruits and more seeds than the other Cacteae species studied. Mcintosh (2005) found 1727-3064 seeds per fruit in F. cylindraceus (Engelm.) Orcutt and F. wislizeni (Engelm.) Britton & Rose, so F. histrix is near the average for the genus. According to Rojas-Arechiga and Vazquez-Yanes (2000), the number of seeds per fruit in Cactaceae species is highly variable; these differences may serve as strategies for reproductive efforts as mentioned by Harper et al. (1970). The number of seeds per fruit may also depend on the age and size of the plant, the number of flowers produced, and the origin (wild or cultivated species), as demonstrated for Cactoideae (Parker, 1987; Leon de la Luz and Dominguez-Cadena, 1991; Rojas-Arechiga et al., 2001; Guillen et al., 2011), or may reflect environmental heterogeneity (Harper et al., 1970). However, more detailed studies are needed on intra-and interspecific variability for this characteristic.

With the exception of Opuntioideae seeds, cacti seeds germinate quickly (Bregman and Bouman, 1983; Potter et al., 1984; Del Castillo, 1986; Rojas-Arechiga and Vazquez-Yanes, 2000; Mandujano et al., 2005). The species analyzed in this study began the germination process between 3 and 6 DAS. Two studied species, F. histrix and M. uncinata, showed the highest percentages of germination at 6 DAS (>80%), suggesting that this process is synchronized with available water. In contrast, the species of Coryphantha showed a wider period of germination with their higher values at 9 DAS or even 15 DAS (22 or 28% respectively). Although there was a differential time of germination all species reached 100% germination after 22 DAS. The germination in the species studied is greater than those reported by Sanchez-Soto et al. (2010), Jimenez and Flores (2010), and Flores et al. (2011) for other species of Ferocactus (62-91%), Mammillaria (42-99%) and Coryphantha (72-98%) However, the high germination percentages did not include any pregermination treatment, in contrast with other studies in which specific requirements were needed as for light, water availability, and mechanical scarification (Nolasco et al., 1996, 1997; Godinez-Alvarez and Valiente-Banuet, 1998; De la Barrera and Nobel, 2003; Martinez-Mendoza et al., 2004; Larrea-Alcazar and Lopez, 2008). The pregermination treatments include light-dark treatment, different temperatures, addition of growth regulators (Cancino et al., 1993; Flores et al., 2006; Ortega-Baes and Rojas-Arechiga, 2007; Ortega-Baes et al., 2010; Sanchez-Soto et al., 2010), chemical scarification (Rosas-Lopez and Collazo-Ortega, 2004), and various stress conditions and water potentials (Guillen et al., 2009). In the present study, seeds recently collected were included. Moreover, laboratory conditions during the experiment (20-25[degrees]C) were sufficient to promote seed germination. Temperature is an important factor that can significantly affect the germination percentage. Trujillo (1982) mentioned that cactus seeds germinate at temperatures between 24 and 27[degrees]C. There are species of cacti that require temperatures ranging from 15 to 35[degrees]C for germination, such as T. terscheckii (J. Parra. ex Pfeiff.) Britton & Rose (Ortega-Baes and Rojas-Arechiga, 2007), Eehinoeaetus platyacanthus Link & Otto (Quintana, 1994) and Epiphyllum phyllanthus (L.) Haw. (Simao et al., 2010). Other authors suggest that alternating temperatures increase germination rates of globose species such as Echinocactus platyacanthus, F. flavovirens, F. robustus (Pfeiff.) Britton & Rose, and M. mazatlanensis (Rojas-Arechiga et al., 1998; Sanchez-Soto et al., 2010). In addition to temperature, germination also depends on the species and seed characteristics (thickness of the seed coat, dormancy, etc.). In some species, characteristics of the testa, in particular cuticular secretions, contribute to improve the collection and distribution of water during imbibition and increase germination (Bregman and Graven, 1997). Specific features of the seed (thin seed testa, absence of dormancy and others) in the species studied herein probably influenced significantly the higher germination rate, as has been observed in germinating seeds of columnar cacti (Loza-Cornejo et al., 2008). Further work on the mechanisms of germination is needed, particularly on the requirements for the germination of seeds of other Cactoideae, with the aim of contributing to their use, management and conservation. Some species, such as F. histrix, are subject to special protection in regions of Jalisco and other Mexican states, and other species, such as C. clavata, are considered to be rare species in Jalisco (Chavez-Martinez et al., 2007; Harker et al., 2008).

Significant differences were observed for the number of seeds per fruit, weight, and fruit width in the cacti studied. Seed germination was a rapid process, usually starting on the 3 DAS, and high percentages of germination were seen. Some morphological characteristics of fruits and seeds can be used to support further systematic studies of Cactoideae genera and will contribute to knowledge for the potential use and conservation of these plants.

Received: 10/28/2011. Modified: 02/17/2012 Accepted: 02/23/2012.

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Teresa Terrazas. Ph.D. in Biology, University of North Carolina at Chapel Hill, USA. Professor, UNAM, Mexico. Address: Instituto de Biologia, Universidad Nacional Autono ma de Mexico, Apartado Postal 70-233, Mexico, D.F. 04510. Mexico. e-mail: tterrazas@ibiologia.unam.mx

Lauro Lopez-Mata. Ph.D. in Biology, University of North Carolina at Chapel Hill, USA. Professor, COLPOS, Montecillo, Mexico.
TABLE I
SEED NUMBER, MASS AND SIZE OF FRUITS OF FIVE
GLOBOSE SPECIES OF CACTEAE

Species                    Seeds/fruit             Weight
                             (number)                (g)

Coryphantha bumamma       84 [+ or -] 35 b   2.0 [+ or -] 0.56 c
Coryphantha clavata       56 [+ or -] 8 a    0.5 [+ or -] 0.18 a
Coryphantha cornifera     99 [+ or -] 2 b    1.4 [+ or -] 0.22 b
Ferocactus histrix      2100 [+ or -] 90 c   8.0 [+ or -] 2.80 d
Mammillaria uncinata      70 [+ or -] 13 b   0.5 [+ or -] 0.20 a

Species                       Length                Width
                               (cm)                  (cm)

Coryphantha bumamma     2.7 [+ or -] 0.5 b   0.6 [+ or -] 0.40 a
Coryphantha clavata     1.4 [+ or -] 0.3 a   0.6 [+ or -] 0.15 a
Coryphantha cornifera   2.7 [+ or -] 0.9 b   1.2 [+ or -] 0.80 ab
Ferocactus histrix      3.2 [+ or -] 0.6 b   2.2 [+ or -] 0.40 b
Mammillaria uncinata    1.5 [+ or -] 0.3 a   0.5 [+ or -] 0.01 a

Means [+ or -] SD. n= 30 fruits. Different letters in columns mean
significant differences (p<0.05).

TABLE II
MASS, SIZE AND ROUNDNESS INDEX OF FEEDS
OF FIVE GLOBOSE SPECIES OF CACTEAE

                        Weight               Length
Species                 (mg)                 (mm)

Coryphantha bumamma     1.9 [+ o -] 0.10 d   2.3 [+ o -] 0.12 d
Coryphantha clavata     1.8 [+ o -] 0.28 d   1.4 [+ o -] 0.12 c
Coryphantha cornifera   0.2 [+ o -] 0.07 a   0.9 [+ o -] 0.04 a
Ferocactus histrix      0.6 [+ o -] 0.20 b   1.4 [+ o -] 0.10 c
Mammillaria uncinata    1.1 [+ o -] 0.15 c   1.0 [+ o -] 0.02 b

                        Width                 Roundness
Species                 (mm)                    index

Coryphantha bumamma     0.7 [+ o -] 0.50 ab      0.3
Coryphantha clavata     0.9 [+ o -] 0.13 b       0.6
Coryphantha cornifera   0.6 [+ o -] 0.10 a       0.7
Ferocactus histrix      0.9 [+ o -] 0.10 b       0.6
Mammillaria uncinata    0.7 [+ o -] 0.14 a       0.7

Means [+ o -] SD. n= 200 seeds. Different letters in columns mean
significant differences (p<0.05).
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Title Annotation:REPORTS/COMUNICACIONES/COMUNICACOES
Author:Loza-Cornejo, Sofia; Terrazas, Teresa; Lopez-Mata, Lauro
Publication:Interciencia
Date:Mar 1, 2012
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