Caryophyllaceae is a medium-sized family composed mainly by herbs from temperate zones or tropical mountains. Fairly homogeneous, the types most distinct in habit are small saplings and some cushions from extreme mountainous regions. Like Brassicaceae, regional listings usually include a large number of naturalized, invasive and introduced species. Many species are grown as ornamental plants, and some species are widespread weeds. Most species grow in the Mediterranean and bordering regions of Europe and Asia. The number of genera and species in the Southern Hemisphere is rather small, although the family does contain Antarctic pearlwort (Colobanthus quitensis), the world's southernmost dicot, which is one of only two flowering plants found in Antarctica.
In Brazil, the diversity of Caryophyllaceae has always been imprecise, mainly due to the fact that the main exclusive Brazilian botanical reference, Reflora (SEE) brings the vast majority of species as 'naturalized', and the largest reference of New World flora, the VPA (SEE), is based on in Reflora and bring the same imprecision. Because most of the Caryophyllaceae in Brazil are small herbs of no ornamental, economic or cultural value, there are few references to them outside the academic world.
With the objective of creating a point of support in this discussion, NSAA made a review of the family in Brazil, using as reference the species cited as Brazilian in POWO (SEE), each one being analyzed within the scope of the VPA and the records of each one. in Species Link (SEE), with eventual complement from other sources, such as Zuloaga et al. (Darwiniana, 2019). Maps were edited based on these records, one per genre, and some referenced images were added. In some genera, we excluded some records due to the fact that they are too disjoint from the occurrence bulk of a given species, suggesting that they are material incorrectly identified, or non-native material.
In this analysis we identify 25 spp. in 10 genera among six tribes native in Brazil, three of them are endemic, highly centered in the mountain forests and grasslands of the southern region, with Rio Grande do Sul leading the national diversity in species (22), but at least two genera (Polycarpon, Polycarpaea) do not occur in most of the region, and are associated with tropical savannas. Only tribe Polycarpeae reaches into northern and center Brazil. In New World, there are 535 spp. within 41 genera.
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| DENSITY OF CARYOPHYLLACEAE GENERA IN BRAZIL EXCEPT DRYMARIA, POLYCARPON AND POLYCARPAEA |
KEY FOR BRAZILIAN GENERA
A dichotomous key for the Brazilian genera, adapted from FNA (SEE) to the North American species, which means that some marked characteristics are not present in species from the country.
(1) Leaves with stipules present - (2)
(2) Fruits utricles; seed 1; petals absent; staminodes petaloid or scalelike - (3)
(3) Sepal awns very stout, spinose; flowers densely woolly ----- Cardionema
(3) Sepals awnless or awns threadlike to somewhat stout, not especially spiny; flowers not woolly ----- Paronychia
(2) Fruits capsules; seeds 3-150+; petals rarely absent or replaced by staminodes
(4) Styles 1, or (2-)3, connate proximally for 2 or more of length or rarely distinct - 5
(5) Petal blades divided into 2-4 lobes; styles 2(-3) ----- Drymaria (in part)
(5) Petal blades entire, emarginate, or irregularly toothed; style 1 - 6
(6) Sepals herbaceous, margins white, apex acuminate to aristate, obscurely to distinctly keeled ----- Polycarpon
(6) Sepals scarious, central portion silvery with brown midrib, margins silvery, apex acute, not keeled ----- Polycarpaea
(4) Styles 3 or 5, occasionally 2, distinct or nearly so - 7
(7) Leaf blades broadly ovate to orbiculate; petal blades with apex divided into 2 linear lobes ----- Drymaria (in part)
(7) Leaf blades linear or subulate to oblong; petal blades with apex entire or petals rudimentary or absent - 8
(8) Leaves opposite but appearing whorled; styles and capsule valves 5 ----- Spergula
(8) Leaves opposite; styles and capsule valves 3 ------ Spergularia
(1) Leaves with stipules absent - 9
(9) Petals absent or rudimentary - 10
(10) Capsules cylindric, opening by 8 or 10 teeth ----- Cerastium
(10) Capsules ovoid to globose, opening by valves or teeth - 11
(11) Capsule valves or teeth 2 times number of styles ------ Sagina (in part)
(11) Capsule valves equal in number to styles ----- Arenaria (in part)
(9) Petals present - 12
(12) Petal apices 2-lobed or 2-fid, often divided nearly to base, or 4-fid - 13
(13) Plants prostrate or sprawling, glabrous, glaucous, and succulent; petals 4-fid ----- Drymaria (in part)
(13) Plants, if prostrate, not glabrous, glaucous, succulent; petals 2-lobed or 2-fid ----- Cerastium (in part)
(12) Petal apices entire, emarginate, jagged, or notched - 14
(14) Styles and capsule valves equal in number ----- Sagina (in part)
(14) Capsule valves or teeth 2 times number of styles - 15
(15) Petals (4-)5, blade apex shallowly emarginate or notched; capsules longer than sepals ------ Cerastium (in part)
(15) Petals 4, blade apex entire; capsules shorter than or equaling sepals ----- Arenaria (in part)
PHYLOGENY among Brazilian Caryophyllaceae, Paronychia is sister of all remaining genera, based on Dillenberger & Kadereit (Taxon, 2014).
TAXONOMY the subdivision below follows Harbaugh & al. (International Journal of Plant Sciences, 2010) and Greenberg & Donoghue (Taxon, 2011), although it is still under construction (many generic names unplaced). The descriptions of the clades are mainly according to Harbaugh & al. (2010). For this classification Caryophyllaceae has two subfamilies. Subfamily Telephioideae has two genera and c. 15 spp., nearly cosmopolitan, especially Europe and Africa, also Madagascar. 4 spp. in South America, 3 restricted for Cono Sur, and Corrigiola andina Planch. & Triana disjunct in Mexico, Peru and Bolivia. Subfamily Caryophylloideae has 11 clades, Sclerantheae (9–10/85–100, Northern Hemisphere, Ethiopia, New Guinea, Australia), Eremogoneae (1/90–100, temperate regions on the northern Hemisphere), Rhodalsine clade (1/1, Mediterranean, Canary Islands, Morocco to N Egypt, N Somalia) and Caryophylleae (13/665–685, Eurasia, Africa, North America and Australia and New Zealand) do not occcur in South America; among the seven South American tribes, only Sileneae (2/882) does not occur in Brazil.
TRIBE PARONYCHEAE ‣ six genera, three only in Eurasia, Herniaria from Eurasia with a outlier in Bolivia and Argentina, Phillipiella endemic to Chile and Argentina, and Paronychia in Brazil.
1. Paronychia Mill.
Herbs, annual, biennial, or perennial, sometimes with woody base. Taproots filiform to stout. Stems prostrate, ascending, or erect, simple or branched, terete to angular. Leaves opposite, connate by stipules from adjacent leaves, petiolate (basal) or sessile (cauline); stipules 2 per node, often conspicuous, white or silvery, subulate to lanceolate or ovate, margins entire or fimbriate, apex subobtuse or acute to acuminate, unlobed or sometimes deeply 2-fid; blade 1-veined, linear to elliptic, oblanceolate, or spatulate, sometimes thickened and succulent, apex obtuse or acute to acuminate or spinose. Inflorescences terminal or sometimes axillary, frequently much-branched or congested cymes, or flowers solitary; bracts paired, dimorphic (resembling leaf blades and stipules), often concealing flowers. Pedicels erect in fruit. Flowers bisexual or rarely unisexual, some plants also having staminate unisexual flowers, others also having pistillate unisexual flowers, not woolly, with hairs ± straight or tips coiled, 0.1-0.3 mm; hypanthium cup-shaped, tapering or expanded distally; sepals (3-)5, connate proximally, white or yellowish to green or reddish or purplish brown, subulate to linear-oblong, lanceolate, spatulate, or ovate, 0.4-4.5 mm, margins translucent to white, scarious or papery, apex defined by a usually prominent adaxial hood, ascending to slightly descending, rounded to triangular, sometimes absent, apex obtuse or rounded, usually with terminal or subterminal cusp, crest, mucro, or prominent awn (often thickened-conic proximally, spinose distally); nectar secreted from within hypanthium; stamens usually 5; filaments distinct or connate proximally with alternating staminodes; staminodes absent or 5, arising from hypanthium rim, subulate to narrowly triangular, filiform, or oblong; styles 1-2(-3), distinct or often connate proximally 10- 10 of length, subcapitate to filiform, 0.07-3.2 mm, glabrous proximally; stigmas 2(-3), subterminal or linear along adaxial surface of style branches, obscurely papillate (50×). Utricles ovoid to globose or rarely 4-angular, membranous, indehiscent. Seeds brown, subglobose to ellipsoid, laterally compressed, smooth, marginal wing absent, appendage absent; embryo peripheral, curved. x = 7, 8, 9 (FNA).
116 spp., Canada to Terra del Fuego except some places in tropical South America, Central America and Caribbean, also from Mediterranean Basin up to Somalia and Iran; 53 spp. in New World, 26 in South America, 22 from Venezuela to Cono Sur, six in Brazil (all South American exclusives).
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| Paronychia communis Cambess. in Santa Catarina state, Brazil © Luiz A. Funez/floradesantacatarina.wordpress.com |
Zuloaga et al. (2019) recognizes for southern Brazil the species P. brasiliana DC., P. camphorosmoides Cambess., P. chilensis DC., P. communis Cambess., P. revoluta C.E. Carneiro & A. Furlan, P. setigera (Gillies ex Hook. & Arn.) F. Herm.; all are fully recognizes native in Brazil by POWO (SEE), with P. camphorosmoides Cambess. under as Cardionema camphosmoides (Cambess.) A.Nelson & J.F.Macbr., plus a species absent in southern country, P. fasciculata Chaudhri, endemic to Minas Gerais state.
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| Paronychia revoluta C.E. Carneiro & Furlan in Santa Catarina state, Brazil © Luiz A. Funez/biodiversity4all.org/photos/108928075 |
P. revoluta C.E. Carneiro & A. Furlan was described as endemic to Rio Grande do Sul (Carneiro & Furlan, Novon, 2004), but there are records of it in Santa Catarina (SEE); Species Link cites P. fasciculata Chaudhri native in Rio Grande do Sul, but this occurrence is here rejected; P. setigera (Gillies ex Hook. & Arn.) F. Herm. in Brazil was collected only in Quaraí region, Rio Grande do Sul state; remainig species are widely distributed in S & SE country. Records of the genus in Ladário (MS), Pico das Almas (BA) and Luziania (GO) are highly doubtful and are here rejected.
TRIBE POLYCARPAEAE ‣ c. 20 genera and c. 160 spp., mainly in Mediterranean Basin, Africa, North America to Caribbean, 7 genera in South America, 4 in Brazil, e 3 remaining (Augustea, Microphyes, Pycnophyllum) from Peru, Bolivia, Chile, Argentina and Uruguay.
2. Cardionema DC
Herbs, perennial. Taproots often stout. Stems prostrate, much-branched proximally, terete to angular. Leaves opposite, not connate, connected by thickened margins from which stipules arise, sessile; stipules 2 per node, silvery, lanceolate to ovate, margins entire to irregularly fimbriate, apex 2-lobed; blade prominently 1-veined, needlelike, not succulent, apex spinose. Inflorescences axillary, 1-3-flowered clusters; bracts paired, resembling stipules, smaller. Flowers sessile; hypanthium cup-shaped, not abruptly expanded distally; sepals 5, distinct, olive-green, oblong to obovate, 1.2-2.8 mm (excluding awns), indurate, herbaceous, margins whitish, scarious, apex spinose, prominently hooded, awned, hoods projecting inward and enclosing developing fruit, awns arising from near apex, widely divergent, very stout, spinose, 3 awns and associated sepals well developed, alternating with 2 reduced sepals; nectaries not apparent; stamens 3-5; filaments distinct to base; staminodes 5, adnate basally with alternating filaments, ovate-triangular, apex acuminate; styles 2, distinct, subcapitate, 0.2 mm, glabrous proximally; stigmas 2, subterminal, obscurely papillate (50×). Utricles ellipsoid to cylindric, teeth absent, indehiscent. Seeds tan with red spot at hilum, narrowly ovate, not compressed, smooth, marginal wing absent, appendage absent; embryo peripheral, straight to slightly curved (FNA).
Six spp., four only in W South America, C. camphorosmoides (Cambess.) A.Nelson & J.F.Macbr. endemic to E Brazil, and C. ramosissimum (Weinm.) A. Nelson & J.F. Macbr. widely distributed from North America to Cono Sur and SE Brazil, a aggressive spine fruited common in coast of S Brazil. Zuloaga et al. (2019) recognizes for southern Brazil the species C. ramosissima (Weinm.) A. Nelson & J.F. Macbr. and C. rosetta (Cambess.) A.Nelson & J.F. Macbr. POWO synonymize the second over the first, and recognize it in Brazil (SEE), and recognizes in this genus Cardionema camphorosmoides (Cambess.) A.Nelson & J.F.Macbr. (endemic to Brazil), which Zuloaga et al. (2019) and Reflora puts as Paronychia camphorosmoides Cambess. Reflora considers C. ramosissima naturalized in the country (SEE), and Cardionema camphorosmoides/Paronychia camphrosmoides as native (SEE).
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| Cardionema ramosissimum (Weinm.) A. Nelson & J.F. Macbr. in Santa Catarina state, Brazil © Luiz A. Funez/Flora de Santa Catarina |
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| Cardionema camphorosmoides (Cambess.) A.Nelson & J.F.Macbr. in Santa Catarina state, Brazil © Luiz A. Funez/Flora de Santa Catarina |
In Brazil C. ramosissima (Weinm.) A. Nelson & J.F. Macbr. occur only southern regions, mainly in coastal places, while C. camphorosmoides (Cambess.) A.Nelson & J.F.Macbr. was collected in vast area from Bahia to Rio Grande do Sul state. A record of C. ramosissima in SE Mato Grosso state was rejected by high inconsistency of records so far from the center of diversity.
3. Drymaria Willd. ex. J. A. Schults.
Herbs, annual or perennial, caudices often branched. Taproots slender, elongate. Stems sprawling to erect, simple or branching proximally or throughout, terete. Leaves opposite or appearing whorled, connate by membranous to thickened line, petiolate or sessile; stipules 2 per node, white to tan, simple or divided into segments, subulate to filiform, often minute, margins entire, apex acute to acuminate; blade 1-5-veined, linear to lanceolate, spatulate, ovate, reniform, or orbiculate, not succulent, apex rounded to acuminate. Inflorescences terminal or axillary, open to congested, bracteate cymes or umbelliform clusters or flowers solitary, axillary; bracts paired, scarious or central portion herbaceous. Pedicels erect to spreading or reflexed. Flowers: perianth and androecium hypogynous; sepals 5, distinct, white, lanceolate to oblong, ovate, or orbiculate, 1.5-4.8(-5) mm, herbaceous, margins white to purple, scarious, apex acuminate to rounded, hooded or not; petals (3-)5, sometimes absent, white, claw narrow, tapering distally or with oblong or expanded, sessile or short-clawed trunk, auricles absent, blade apex divided into 2 or 4 lobes; nectaries at base of filaments opposite sepals; stamens 5; filaments distinct or briefly connate proximally; styles 3, occasionally 2, connate proximally for 2 of length, rarely to nearly distinct (D. cordata), filiform, 0.1-0.3 mm, glabrous proximally; stigmas 3, occasionally 2, linear along adaxial surfaces of styles (or branches), obscurely papillate (30×). Capsules ellipsoid to globose, opening by (2-)3 spreading to recurved valves; carpophore absent. Seeds 3-25, tan, reddish brown, dark brown, black, or transparent (white embryo visible), horseshoe-, snail-shell- or teardrop-shaped, compressed laterally, at least somewhat, tuberculate, marginal wing absent, appendage absent. x = (11), 12 (FNA).
57 spp., U.S.A. to Argentina, tropical Africa, tropical Asia to Australia; 56 spp. are exclusive to New World, 22 in South America, only one in Brazil. Zuloaga et al. (2019) recognizes for southern Brazil the species D. cordata (L.) Willd. ex Roem. & Schult. POWO only recognizes this one in Brazil (SEE), and Reflora gives it as naturalized (SEE).
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| Drymaria cordata (L.) Willd. ex Roem. & Schult. in Santa Catarina state, Brazil © Luiz A. Funez/Flora de Santa Catarina |
In Brazil D. cordata occur in over country in more than 20 states; It is the most distributed species in the country, occurring in several environments, being well adapted to the tropical climate.
4. Polycarpaea Lam.
Herbs [small shrubs], annual [perennial]. Taproots slender to stout. Stems erect, branched, terete. Leaves opposite, sometimes appearing whorled, not connate, short-petiolate (basal leaves) or sessile (cauline leaves); stipules 2 per node, white, lanceolate, margins entire, apex hairlike; blade 1-veined, oblong-ovate to suborbiculate (basal leaves), subulate to linear (cauline leaves), not succulent, apex acute or hairlike. Inflorescences terminal, compact to loose cymes; bracts paired, scarious. Pedicels erect or spreading. Flowers: perianth and androecium perigynous; hypanthium cup-shaped, not abruptly expanded distally; sepals distinct, silvery with brown midrib, ovate to lanceolate, not keeled, 2.2-3.1 mm, scarious, margins scarious, apex acute; petals 5, pink, blade apex irregularly toothed or entire; nectaries between filament bases; stamens 5; filaments distinct; style 1, filiform, minute, less than 0.1 mm, glabrous proximally; stigmas 3, subcapitate, smooth to obscurely papillate (50×). Capsules ellipsoid, opening by 3 recurved valves; carpophore present. Seeds 5-8, tan, translucent, pyriform, not compressed, rugulose, marginal wing absent, appendage absent. x = 9 (FNA).
74 spp., mainly Old World, only two in New World, P. hassleriana Chodat from Bolivia and Paraguay, and P. corymbosa (L.) Lam. from over South America (collected in several Brazilian states), Panamá, tropical Africa south to Saara, India and E China to Thailand.
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| Polycarpaea corymbosa (L.) Lam. in Pará state, Brazil © André Cardoso/flickr.com |
Zuloaga et al. (2019) recognizes for southern Brazil the species Polycarpaea corymbosa (L.) Lam., as well as POWO (SEE), but no recognized in Brazil as native by Reflora (SEE); for Brazilian range of this genus, see Guterres et al. (Harvard Papers in Botany, 2021).
In Brazil this genus is widelt distributed in all regions, higly centered in savannas from Tocantins, Goiás, Bahia and Minas Gerais state. Is the only Caryophyllacae from Roraima state.
5. Polycarpon Loef.
Herbs, annual. Taproots slender. Stems prostrate to erect, branched, terete to finely ridged. Leaves opposite or in whorls of 4, not connate, petiolate; stipules 2 per node, silvery, lanceolate to triangular-ovate, margins entire or irregularly cut, apex acuminate to aristate; blade 1-veined, spatulate or oblanceolate to ovate or elliptic, not succulent, apex obtuse, sometimes mucronate. Inflorescences terminal or axillary, dense or lax cymes; bracts paired or absent. Pedicels erect. Flowers: perianth and androecium perigynous; hypanthium minute, cup-shaped, not abruptly expanded distally; sepals distinct, green, lanceolate to elliptic or ovate, often keeled, 1-2.5 mm, herbaceous, margins white, scarious, apex acute, ± hooded, ± awned; petals often fugacious, 5, white, blade apex emarginate; nectaries between filament bases; stamens 3-5; filaments shortly connate distally around ovary; style 1, obscurely 3-branched, filiform, 0.1-0.3 mm, glabrous proximally; stigmas 3, linear along adaxial surface of style branches, papillate (30×). Capsules ovoid to spherical, opening by 3 incurved or twisting valves; carpophore present. Seeds ca. 8-15, whitish, ovoid to lenticular or triangular, laterally compressed to angular, papillate or granular, marginal wing absent, appendage absent. x = [7], 8, 9 (FNA).
8 spp., 4 from Europe to N Africa and India, make up a complex of highly similar, annual or perennial taxa mainly distributed in Mediterranean region, P. depressum Nutt. in California to Mexico (Baja California), P. prostratum (Forssk.) Asch. & Schweinf. in over tropical Old World, and P. apurense Kunth in Venezuela, Colombia, Peru, Paraguay, Brazil, and Argentina. Zuloaga et al. (2019) recognizes for southern Brazil the species Polycarpon apurense Kunth; however, POWO does not recognize this species in Brazil (SEE), neither Reflora (SEE). Gereau e Marticorena (Novon, 1995) cites this species in Brazil. Here we accept the species in Brazil, based on this reference.
In Brazil this species occur in few places among riversides in savannas from C Brazil, from Pará to São Paulo state.
TRIBE SPERGULARIEAE ‣ three genera, two in South America (both in Brazl), Thylacospermum monotypic only in central Asia.
6. Spergula Dill. ex. L.
Herbs, annual or winter annual. Taproots slender to ± stout, especially proximally. Stems spreading or ascending to erect, simple or branched, terete to somewhat angular. Leaves opposite but appearing whorled, as 8-15 per axillary cluster, 2 clusters per node, connate proximally by often-prominent ridge from which stipules arise, sessile; stipules 4 per node, white, ovate to triangular, margins entire but splitting variously with age, apex obtuse to acuminate; blade 1-veined, linear or filiform, sometimes succulent, apex blunt to apiculate. Inflorescences terminal, open to diffuse cymes; bracts paired, minute. Pedicels erect to ascending, spreading or usually reflexed and sometimes secund in fruit. Flowers usually bisexual, sometimes pistillate by stamen abortion; perianth and androecium briefly perigynous; hypanthium dish- or cup-shaped, not abruptly expanded distally; sepals distinct, silvery, elliptic to nearly ovate, 2.5-5 mm, herbaceous, margins scarious, apex acute to obtuse; petals 5, white, blade apex entire; nectaries at adaxial base of broader filaments opposite sepals; stamens 5 and opposite sepals, or 10 and arising from distally tapered rim of hypanthium; filaments distinct; styles 5, distinct, filiform, 0.4-0.6 mm, glabrous proximally; stigmas 5, linear along adaxial surface of styles, obscurely papillate (30×). Capsules ovoid, opening by 5 spreading to somewhat recurved valves; carpophore absent. Seeds 5-25, blackish, circular, subglobose or lenticular and laterally compressed, nearly smooth or finely papillate, membranous, entire marginal wing often present, appendage absent; embryo peripheral, annular to spirally curved. x = 9 (FNA).
12 spp., Algeria to Siberia, tropical Africa, and southern South America (9), two up to S Brazil. Zuloaga et al. (2019) recognizes for southern Brazil the species Spergula grandis Pers., Spergula levis (Cambess.) D. Dietr. and Spergula ramosa (Cambess.) D. Dietr. POWO recognizes the three native to Brazil (SEE), but points out the third in another genus, Spergularia L. (S. ramosa Cambess., SEE). Reflora recognizes Spergula ramosa within Spergula grandis (SEE), as naturalized, and rejects fully S. levis.
Brazilian members occur in three states of southern region, but S. levis in known in Brazil only by one collection, of unknown exact location, but assinaled for Rio Grande do Sul state.
7. Spergularia (Pers.) J.Presl & C.Presl
Herbs, annual or strongly perennial with branched, woody caudex. Taproots filiform to stout. Stems erect to sprawling, simple to freely branching distally or throughout, terete, sometimes woody. Leaves opposite, axillary clusters of leaves often present, distinct, sessile; stipules 2 per node, white to tan, lanceolate and acuminate to widely triangular, margins entire, apex entire to variously split; blade 1-veined, threadlike to linear, mostly succulent, apex acute to acuminate. Inflorescences terminal cymes, branching symmetrically or to 1 side (monochasium), simple to 8+-compound or sometimes flowers solitary and axillary; bracts usually paired or sometimes single, smaller, foliaceous, distalmost sometimes with scarious margins. Pedicels ascending to erect, divergently spreading, reflexed, or arching downward in fruit. Flowers: perianth and androecium hypogynous, briefly perigynous; hypanthium dish- or cup-shaped, not abruptly expanded distally; sepals connate in proximal 1/5, green, lanceolate to ovate, 0.9-8 mm, herbaceous, margins scarious, apex acute to obtuse; petals 5, white to pink, blade apex entire; nectaries as lateral expansion of bases of filaments opposite sepals; stamens 1-10, arising from rim of hypanthium; filaments distinct; styles 3, distinct or nearly so, filiform, 0.2-3 mm, glabrous proximally; stigmas 3, linear along adaxial surface of styles, obscurely papillate (30×). Capsules ovoid, opening by 3 spreading valves with recurved tips; carpophore absent. Seeds 30-150+, light to dark brown, reddish brown, or black, circular to angular, plump or laterally compressed, smooth to variously sculptured to papillate, complete or partial, membranous, laciniate, marginal wing often present, appendage absent. x = 9 (FNA).
63 spp., cosmopolitan except Central America, several tropical areas in South America, Africa and southern Asia; 24 spp. in New World, 19 in South America, only S. ramosa Cambess. in Brazil. Zuloaga et al. (2019) recognizes for southern Brazil the species Spergula ramosa (Cambess.) D. Dietr. POWO points in another genus, Spergularia L. (S. ramosa Cambess., SEE). Reflora recognizes Spergula ramosa within Spergula grandis (SEE), as naturalized.
Spergularia ramosa Cambess. in known in Brazil only by few collections in southern Rio Grande do Sul state, at grasslands.
TRIBE SAGINEAE ‣ c. 10 genera and c. 200 spp., mainly Mediterranean Basin to India, three genera in New World, all in South America: Colobanthus in South America, Antarctic region and Australia, with disjuntc records in Mexico; Sabulina from northern Hemisphere with one outlier in Chile; and Sagina, which occur in Brazil.
8. Sagina L.
Herbs, annual, winter-annual, or perennial, often cespitose or matted. Taproots slender. Stems ascending, decumbent, or procumbent, simple or branched, terete to slightly angular. Leaves: basal and secondary rosettes present in perennial species, usually connate proximally, sometimes forming conspicuous, scarious cup, sessile, with or without axillary fascicles of leaves; blade 1-veined, linear to subulate, succulent or not, apex acute to mucronate or apiculate (or long-aristate in S. subulata). Inflorescences terminal or axillary cymes, or flowers solitary; bracts paired, foliaceous. Pedicels erect or spreading. Flowers: perianth and androecium hypogynous; sepals 4 or 5, distinct, green or sometimes purple, lanceolate to elliptic or orbiculate, 1-5.5 mm, herbaceous, margins white or purple, scarious, apex obtuse or rounded to somewhat acute, frequently hood-shaped in bud; petals 4 or 5, sometimes absent (frequently absent or soon dropping in annual species), white, claw absent or minute, blade apex entire; nectaries at base of filaments opposite sepals; stamens 4, 5, 8, or 10, arising from base of ovary; filaments distinct; staminodes absent; styles 4 or 5, clavate to filiform, 0.5-1.5 mm, glabrous proximally; stigmas 4 or 5, subterminal to linear along adaxial surface of styles, minutely papillate (30×). Capsules globose to ovoid, opening by 4 or 5 valves, sutures running to base, but in some species dehiscing only ca. 1/ 1/ 2 capsule length; carpophore absent. Seeds ca. 125, light tan to dark or reddish brown, obliquely triangular with abaxial groove, or reniform to nearly globose without abaxial groove, laterally compressed or plump, smooth to tuberculate, marginal wing absent, appendage absent. x = 6, 7, 11. (FNA).
36 spp. (POWO), Canada to Mexico (6), Caribbean, southern South America, Africa, Eurasia to SE Australia. Two species in South America, both in Brazil. Zuloaga et al. (2019) recognizes for southern Brazil the species S. chilensis Naudin ex Gay and S. humifusa (Cambess.) Fenzl & Rohrb., both also accepted in Brazil by POWO (SEE); Reflora recognizes the latter native (SEE) and the former naturalized (SEE).
Both species in Brazil cited above occur only in Rio Grande do Sul and SE Santa Catarina state, with the latter known only from the former state.
TRIBE ARENARIEAE ‣ two genera and c. 200 spp., Moehingia has c. 30 spp. from northern Hemisphere, and Arenaria L. occur in South America and Brazil.
9. Arenaria L.
Plants annual or perennial, sometimes densely matted. Taproots filiform to moderately thickened; rhizomes slender. Stems prostrate to ascending or erect, simple or branched, terete to ellipsoid, angular or grooved. Leaves mostly connate, mostly sessile, not congested at or near base of flowering stem; blade 1- or 3-5-veined, rarely linear or linear-lanceolate to usually elliptic to ovate or rarely orbiculate, sometimes subsucculent or succulent, apex blunt or obtuse to acute, acuminate, or apiculate. Inflorescences terminal or axillary open cymes, or flowers solitary; bracts paired, smaller, foliaceous. Pedicels erect or ascending to reflexed in fruit. Flowers: perianth and androecium hypogynous; sepals 5, distinct or barely connate proximally, green or rarely distally purple, lanceolate to broadly ovate, 2-5 mm, margins foliaceous or white and scarious, apex obtuse to rounded or acute to acuminate, not hooded; petals 5 or absent, white, not clawed, blade apex entire; nectaries at base of filaments opposite sepals; stamens 8-10, arising from base of ovary; filaments distinct; staminodes absent; styles 3, filiform, 0.5-2 mm, glabrous proximally; stigmas 3, linear along adaxial surfaces of styles, papillate (30×). Capsules broadly ellipsoid or ovoid to cylindric, opening by 6 ascending to recurved teeth; carpophore absent. Seeds [1-]5-35, brown to dark brown or black, reniform or suborbicular, laterally compressed or not, shiny or dull, smooth, rugulose, or tuberculate, marginal wing absent, appendage absent. x = [7, 8?] 10, 11 (FNA).
175 spp., cosmopolitan except tropical South America, vast areas in Africa, SE Asia to Australia; 59 spp. in New World, 41 in South America, all belong to former subgen. Dicranilla and subgen. Leiosperma; A. lanuginosa (Michx.) Rohrb. occur from SE. U.S.A. to NW. Venezuela and N. Argentina, S & SE Brazil, Caribbean; it’s a morphologically diverse, both in our area and southward into N South America, and is in serious need of comprehensive study.
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| Arenaria lanuginosa (Michx.) Rohrb. in Santa Catarina state, Brazil © Luiz A. Funez/Flora de Santa Catarina |
Zuloaga et al. (2019) recognizes for southern Brazil the species A. jussiaei Cambess. ex St.-Hil. and A. lanuginosa (Michx.) Rohrb. var. lanuginosa in Brazil, the former endemic. POWO only recognizes the latter in Brazil (SEE). Reflora also only recognizes the latter, which it considers to be naturalized in the country (SEE).
In Brazil A. lanuginosa (Michx.) Rohrb. occur mainly in southern region, with several records in mountainous regions of all states in SE region except in Espírito Santo state; the northemost records is few colections in SE Minas Gerias. Collections of Linhares (ES), Manaus (AM), Parnaíba (PI) and Grao-Mogol (MG) was rejected due to the high inconsistency of records so far from the center of diversity.
TRIBE ALSINEAE ‣ c. 12 genera and c. 400 spp. mainly in northern temperate Eurasia, some genera in North America, three in South America, Pycnophyllopsis Skottsb. restricted from high Andes from Peru to Argentina, and Stellaria L. wider in western flakn of South America. Only Cerastium L. in Brazil.
10. Cerastium L.
Herbs, annual, winter annual, or perennial. Taproots slender, perennial taxa often rhizomatous, rooting at nodes. Stems ascending to erect or decumbent, simple or branched, terete. Leaves basally connate, petiolate (basal in some species) or sessile (cauline); blade 1-5-veined, linear or elliptic to broadly ovate, sometimes succulent, apex acute to obtuse. Inflorescences terminal, open or congested cymes, or flowers solitary, axillary or racemose; bracts paired, foliaceous or reduced, herbaceous or often with scarious margins. Pedicels erect, sometimes reflexed or hooked at apex in fruit, or flowers sometimes subsessile. Flowers bisexual, occasionally unisexual and pistillate; perianth and androecium hypogynous or weakly perigynous; hypanthium minimal; sepals (4-)5, distinct, mainly green, elliptic to ovate, 3-12 mm, herbaceous, margins translucent to purplish, scarious, apex acute, acuminate, or obtuse, not hooded; petals (4-)5 or sometimes absent, white, sometimes purple, clawed, notched, or emarginate; nectaries at base of filaments opposite sepals; stamens usually 10, sometimes 5 or 8, occasionally 4; filaments distinct, inserted at base of ovary; staminodes absent or 1-4 (via anther abortion), linear; styles (3-)5(-6), clavate to filiform, 0.5-2 mm, glabrous proximally; stigmas (3-)5(-6), subterminal to linear along adaxial surface of styles, roughened to papillate (30×). Capsules oblong or cylindric, usually ± curved, opening by 10, or occasionally 6 or 8, erect or spreading, convolute or revolute teeth, longer than sepals; carpophore absent. Seeds 15-150+, orange to brown, angular-obovate, often with abaxial groove, laterally compressed, papillate-tuberculate, marginal wing absent, appendage absent. x = [9?, 13, 15] 17, 18, 19 (FNA).
211 spp., cosmopolitan except northern South America, sevral places in Africa, SE Asia and Oceania. 66 spp. in New World, 38 in South America, 8 native in Brazil, none endemics.
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| Cerastium rivulare Cambess in Santa Catarina state, Brazil © Species Link/Fototeca Paulo Schwirkowski - FPS |
Zuloaga et al. (2019) recognizes for southern Brazil the species C. commersonianum Ser. ex DC., C. dicrotrichum Fenzl ex Rohrb., C. humifusum Cambess., C. junceum Möschl, C. mollissimum Poir., C. rivulare Cambess. and C. rivulariastrum Möschl & Pedersen. POWO recognizes all in Brazil, plus C. parvum (Pedersen) M.T.Sharples & E.A.Tripp, whom Zuloaga et al. (2019) recognizes in Stellaria. None of them are recognized in Brazil as native by Reflora (SEE).
In Brazil this genus is highly centered in eastern flank of southern regions, with all Brazilian species collected in Rio Grande do Sul state; several records also in mountaisn of NE São Paulo, SW Rio de Janeiro, S & SE Minas Gerais state, with from this state the northermost record of this genus (C. rivulare Cambess.). Two records of this genus in Bahia state was rejected due to the high inconsistency of records so far from the center of diversity.
NOT ACCEPTED GENERA