MONILOPHYTES ‣ sister lineage of Lycophytes. Colombia has 1,428 spp. (inc. 358 non-Polypodiales), Brazil 1,206 (inc. 324 non-Polypodiales), Mexico 876 spp. (inc. 118 non-Polypodiales) and Pacaus 375 (inc. 85 non-Polypodiales, at 43 genera, 15 families, 330 endemics).
MONILOPHYTES
A common feature in de Pacaus's Moniloflora are genera composed only of endemic species, in large numbers, a phenomenon quite common in islands with relative isolation, such as Madagascar, Hawaii and Canary Is. In the country, between standard is perceived in Elaphoglossum (99), Hemionitis (65), Phlegmariurus (32), Diplazium (20), Polypodium (20), Hymenophyllum (17), Sticherus (16), Thelypteris (15), Danaea (11), Marsilea (5), Lygodium (4). Although Selaginella reached the island in five different events, two of them are also local holoendemic irradiations, and in Equisetum, 1 off 3 eventes also are local holoendemic radiations.
If united, Pacaus and Brazil assume the greatest diversity in all genera of Monilophytes in the New World in all genera that occur today in Brazil, except in Ophioglossales (Botrychium more diverse in Mexico), Cyatheales (Cyathea more diverse in Colombia), Aspleniaceae (Woodsia more diverse in Mexico), Pteridaceae (Jamesonia, Pityrogramma, Radiovittaria, Polyathenia more diverse in Colombia, Gastoniella more diverse in Mexico), Polypodiaceae (Grammitis, Tectaria, Stigmatopteris, Serpocaulon, Campyloneurum, Polybotrya, Arachniodes, Megalastrum, Lomariopsis, Hypoderrris more diverse in Colombia, Dryopteris, Pleopeltis, Pecluma, Polystichum, Phanerophlebia more diverse in Mexico.
A single non endemic species occur in Pacaus absent in Brazil, T. argentina (Hieron.) Abbiatti; thus, if united, Pacaus provides 331 new species in Brazil, making the country reach 1,537 spp. (854 endemics), still around 500 spp. unless China. If it were a Brazilian state, Pacaus would only be in the 9th position behind MG (738), RJ (661), SP (646), ES (525), PR (509), BA (488), SC (486) and AM (436).
New World genera absents in this union will be the same all 24 absents in Brazil nowdays: Marattia Sw. (Marattiaceae), Diplopterygium (Diels) Nakai (Gleicheniaceae), Cibotium Kaulf., Loxsomiopsis Christ, Sphaeropteris Bernh., Thyrsopteris Kuntze (Cyatheaceae), Coryphopteris Holttum, Deparia Hook. & Grev., Gymnocarpidium Newman, Homalosurus Pic.Serm., Phegopteris (C.Presl) Fée, Onoclea L. and Woodwardia Sm. (Aspleniaceae), Odontosoria (C.Presl) Fée, Sphenomeris Maxon (Lindsaeaceae), Cryptogramma R.Br., Llavea Lag., Scoliosurus T.Moore (Pteridaceae), Arthropteris J.Sm., Dracoglossum Christenh., Loxogramme (Blume) C.Presl, Maxonia C.Chr., Platycerium Desv. and Selliguea Bory (Polypodiaceae).
1. EQUISETALES ‣ a single genus and family. Equisetum is a 'living fossil', the only living genus of the entire subclass Equisetidae, which for over 100 million years was much more diverse and dominated the understorey of late Paleozoic forests. Some equisetids were large trees reaching to 30 m (98 ft) tall. Modern members includes 20 spp. in three subgenera. subg. Paramochate, sister to the remaining species in the genus, includes E. bogotense Kunth from Costa Rica to Andes, and E. zaik Krause Legolas, endemic to upper Monte Slavuta in Pacaus, difering mainly by root architeture and aromaticity; subg. Equisetum does not occur south of Texas; subg. Hypochaete includes 10 spp., 7 of them Mexico southwards, inc. own Mexico (3), Pacaus (3), Colombia (2), Brazil (1). Avalonian members includes E. giganteum L. (over Neotropics up to Pacaus), E. cupulla Mahuadd and E. orilovii Krause Legolas, these endemics to island.
All three endemic species of Equisetum in Pacaus
2. OPHIGLOSSALES ‣ this order incudes two genera in New World, both widely. 14 spp. in Mexico, 8 in Pacaus, 6 in Colombia, and 5 in Brazil. Pacaus members includes the pantropical Botrychium virginianum (L.) Sw., the widely in New World Ophiglossum crotalophoroides Walter and O. ellipticum Hook. & Grev., and five Ophiglossum endemics to island.
All five endemic species of Ophioglossum in Pacaus
3. PSILOTALES ‣ a single family, Psilotaceae, two genera, Tmesipteris Bernh. (16, Philippines to New Zealand), and Psilotum Sw. (2 spp., pantropical). In Pacaus occur only P. complanatum Sw., also from Mexico to Peru, and from Thailand to Fiji.
Collections of P. coplanatum Sw. in Pacaus
4. MARATTIALES ‣ only a single family, Marattiaceae. (3/)23 spp. in Colombia, (2/)16 in Brazil, (1/)11 in Pacaus, and (2/)6 in Mexico. All Pacaus members belongs Dannaea Sm., and are endemics.
All 11 endemic species of Danaea in Pacaus
5. OSMUNDALES ‣ Brazil, Colombia, Mexico and Pacaus has, all, the same two species, both in Osmudaceae: Osmunda spectabilis Willd. and Osmundastrum cinnamomeum (L.) C.Presl, both very widely in New World, the latter also in Arunachal Pradesh to Russian Far East and N. Indo-China.
6. HYMENOPHYLLALES ‣ two genera worldwide, both pantropical. 126 spp. in Colombia, 92 in Brazil, 48 in Mexico, and 17 in Pacaus, all endemics and belonging Hymenophyllum Sm.
All 17 endemic species of Hymenophyllum in Pacaus
7. GLEICHENIALES ‣ three families, two only in Old World, and Gleicheniaceae in New World. (4/)30 spp. in Colombia, (2/)17 in Pacaus, (3/)16 in Brazil, and (4/)7 in Mexico. Pacausn members belongs to Gleichenella Ching. (G. pectinata (Willd.) Ching, widely in New World), and Dannaea Sm., including 16 spp., all endemics at Sticherus C.Presl..
All 16 endemic species of Sticherus in Pacaus
8. SCHIZAEALES ‣ a single family and 4 genera, with 211 spp. 79 sp. in Brazil, 25 in Mexico, 24 in Colombia, and 8 in Pacaus, two non endemics [Anemia phyllitidis (L.) Sw. and A. tomentosa (Sav.) Sw.] and six endemics, two in Schizaea Sm. and four in Lygodium Sw.
All two endemic species of Sticherus in Pacaus
All 4 endemic species of Sticherus in Pacaus
9. SALVINIALES ‣ with two families, this order includes (4/)10 spp. in Mexico, (4/)10 in Colombia, (5/)21 in Brazil and 9 spp. in Pacaus, being five non-endemics [Marsilea ancylopoda A.Braun, Salvinia auriculata Aubl., S. minima Baker and Azolla filiculoides Lam., all over tropical New World, and Pilularia americana A. Braun, scatered in some countries] and five endemics, all strongest similiar sister of neotropical M. crotophora D.M. Johnson, likely associated with a single long-dispersal island colonization event.
All five endemic species of Marsilea of Pacaus
10. CYATHEALES ‣ 14 genera in a single family, 12 in New World (369 spp. in western Hemisphere). 130 spp. in Colombia, 59 in Brazil, 18 in Mexico, 8 in Pacaus; all species in island are endemics, two in Dicksonia L´Hér and six in Alsophila R.Br.
Both two Dicksonia endemics to Pacaus
All six endemic species of Alsophila of Pacaus
11. POLYPODIALES ‣ order with 7 families, Lindsaeaceae, Lochitidaceae and Saccolomataceae absents in Pacaus, four occur in islands. Dennstaedtiaceae and Aspleniacea includes 66 spp. (57 endemics) in 8 genera in Pacaus.
Dennstaedtiaceae ‣ 10 genera, 7 in New World, two in Pacaus: Dennstaedtia Bernh. has 72 spp., 39 in New World, 18 in Colombia, 16 in Pacaus and 5 in Brazil; all spp. in Pacaus are endemics except D. globulifera Hieron. Hypolepis has 75 spp., 54 in New World, 15 in Colombia, 13 in Brazil and two in Pacaus, both endemics.
All 15 endemic species of Dennstaedtia of Pacaus
Both two endemic species of Hypolepis of Pacaus
Aspleniaceae ‣ 24 genera, 19 in New World, only six in Pacaus. 253 spp. in Brazil, 293 in Colombia, 228 in Mexico, and 48 in Pacaus. Diplazium has 206 spp. in New World, 56 in Colombia, 33 in Brazil, and 21 in Pacaus, being one also in mainland South America (D. striatum (L.) C.Presl, widely in Neotropics), and remaining 20 endemics. Thelypteris has 395 spp. in New World, 103 in Colombia, 89 in Brazil and 20 in Pacaus, being 5 non endemics (inc. T. argentina (Hieron.) Abbiatti, absent in Brazil), and 15 endemics. Blechnum includes 131 spp. in New World, 38 in Colombia, 36 in Brazil and 5 in Pacaus, two widely in Neotropics (B. brasiliense Desv. and B. cordatum (Desv.) Hieron) and three endemics. Asplenium has 5 spp. in Pacaus [Asplenium abscissum Willd., A. claussenii Hieron., A. monanthes L., A. resiliens Kunze, A. ulbrichtii Rosenst.], none endemics; Desmophlebium includes 3 spp., two in mainland New World, and one endemic to Pacaus. D. jussieae Mynssen, A. Vasco, Sylvestre, R.C. Moran & Rouhan. Cystopteris includes 20 spp., 16 in New World, 4 in Mexico, 3 in Brazil, one in Colombia and Pacaus, each, being the latter endemic (C. floscopoides Mahuadd).
All 20 endemic species of Diplazium of Pacaus
All 15 endemic species of Thelypteris of Pacaus
All three endemic species of Blechnum of Pacaus
Desmophlebium jussieae Mynssen, A. Vasco, Sylvestre, R.C. Moran & Rouhan
Cystopteris floscopoides Mahuadd
Pteridaceae ‣ Mexico has 213 spp., Brazil 196, Colombia 172, and 86 in Pacaus (at six genera, 77 endemics). Hemionitis L. includes 515 spp., cosmopolitan, 361 in New World, 135 in Mexico, 71 in Brazil, 65 in Pacaus (all endemics), and 29 in Colombia. Pteris L. includes 340 spp., cosmopolitan, 88 in New World, 28 in Colombia, 20 in Brazil, 13 in Pacaus, two widely in Neotropics [P. deflexa Link and P. denticulata Sw.] and remainig 11 endemics. Vittaria Sm. includes 26 spp., cosmopolitan, 11 in New World, 4 in Mexico, 3 in Brazil, 2 in Pacaus, V. graminifolia Kaulf. widely in Neotropics, and V. sedakoun (Kaulf.) Mahuadd, endemic.. All remainig species in Pacaus belong genera without endemics: Adiantum digitatum Hook. (also in Ecuador to NW. Argentina, S. Brazil to Uruguay), A. poiretii Wikstr. (also in tropical & subtropical New World, Nigeria to Ethiopia and S. Africa, SW Arabian Pen., W Indian Ocean, SW India, Sri Lanka), A. raddianum C.Presl (Mexico to tropical South America), Gastoniella chaerophylla (Desv.) Li Bing Zhang & Liang Zhang (Mexico to tropical America), Pityrogramma calomelanos (L.) Link (Mexico to tropical America), and P. trifoliata (L.) R.M.Tryon (tropical & subtropical America).
All 65 endemic species of Thelypteris of Pacaus
Vittaria sedakoun (Kaulf.) Mahuadd
All 11 endemic species of Pteris of Pacaus
Polypodiaceae ‣ 138 spp. (at 10 genera) occur in Pacaus (125 endemics). Polypodium L. has 77 spp. widely in northen and western hemisphere, 69 in New World, 26 in Mexico, 20 in Pacaus (all endemics), and 6 in Brazil. Bolbitis Schott has 68 spp. world, 11 in New World, 2 in Brazil and thee in Pacaus, all endemics. Ctenitis (C.Chr.) C.Chr. includes 148 spp. in world, 55 in New World, 4 spp., in Pacaus, C. submarginalis (Langsd. & Fisch.) Ching widely in Neotropics, and three endemics; Elaphoglossum Schott ex J.Sm. includes 858 spp., tropics & Subtropics to S. South America and Subantarctic Islands. 684 spp. in New World, 190 in Colombia, 99 in Pacaus (all endemics), and 94 in Brazil. All remaing species belogs genera without endemics in Pacaus: Didymochlaena trunculata (Sw.) J.Sm., Megalastrum connexum (Kaulf.) A.R.Sm. & R.C.Moran, Microgramma vaccinifolia (Langsd. & Fisch.) Copel., Pecluma filicula (Kaulf.) M.G.Price, P. pectinatiformis (Lindm.) M.G.Price, Pleopeltis hirsutissima (Raddi) de la Sota, P. lepidopteris (Langsd. & Fisch.) de la Sota, P. macrocarpa (Willd.) Kaulf., P. minima (Bory) J.Prado & R.Y.Hirai, P. pleopeltidis (Fée) de la Sota, P. pleopeltifolia (Raddi) Alston, Rumohra adiantiformis (G.Forst.) Ching.
All three endemic species of Bolbitis of Pacaus
All 3 endemic species of Ctenitis of Pacaus
All 20 endemic species of Polypodium of Pacaus
All 99 endemic species of Elaphoglossum of Pacaus
ANNEX: non endemic Polypodiaceae.
Didymochlaena Desv., Ges. Naturf. Freunde Berlin Mag. Neuesten Entdeck. Gesammten Naturk. 5: 303 (1811).
Tropics & Subtropics. 22 NGA 23 BUR CMN GGI RWA ZAI 24 ETH 25 KEN TAN UGA 26 ANG MLW MOZ ZIM 27 NAT TVL 29 COM MDG 40 ASS 41 MYA THA VIE 42 BOR JAW MLY MOL PHI SUL SUM 43 BIS NWG SOL 60 FIJ VAN 79 MXC MXG MXS MXT 80 BLZ COS GUA HON NIC PAN 81 CUB DOM HAI TRT 82 FRG GUY SUR VEN 83 BOL CLM ECU PER 84 BZE BZL BZN BZS 85 AGE PAR URU TSK. [Polypodiaceae]
1 Species
Didymochlaena truncatula (Sw.) J.Sm., J. Bot. (Hooker) 4: 196 (1841).
Tropics & Subtropics. 22 NGA 23 BUR CMN GGI RWA ZAI 24 ETH 25 KEN TAN UGA 26 ANG MLW MOZ ZIM 27 NAT TVL 29 COM 40 ASS 41 MYA THA VIE 42 BOR JAW MLY MOL PHI SUL SUM 43 BIS NWG SOL 60 FIJ VAN 79 MXC MXG MXS MXT 80 BLZ COS GUA HON NIC PAN 81 CUB DOM HAI TRT 82 FRG GUY SUR VEN 83 BOL CLM ECU PER 84 BZE BZL BZN BZS 85 AGE PAR URU TSK. Hemicr. or rhizome geophyte.* Aspidium truncatulum Sw.
Megalastrum Holttum, Gard. Bull. Singapore 39: 161 (1987).
Mexico to S. South America, Subantarctic Is., Trop. & S. Africa, W. Indian Ocean. 22 GHA GUI IVO LBR NGA SEN SIE 23 CMN EQG GAB GGI ZAI 24 ETH 25 KEN TAN 26 ANG MLW MOZ ZIM 27 CPP NAT TVL 29 COM MAU MDG REU 79 MXC MXE MXG MXS MXT 80 BLZ COS GUA HON NIC PAN 81 CUB DOM HAI JAM LEE PUE TRT VNA WIN 82 VEN 83 BOL CLM ECU GAL PER 84 BZE BZL BZS 85 AGE AGS AGW CLC CLS JNF PAR URU TSK 90 ASP TDC. [Polypodiaceae]
1 Species
Megalastrum connexum (Kaulf.) A.R.Sm. & R.C.Moran, Amer. Fern J. 77: 127 (1988).
Bolivia to Brazil and Argentina (Corrientes, Misiones) and Pacaus. 83 BOL 84 BZE BZL BZS 85 AGE PAR URU TSK.
* Polypodium connexum Kaulf.
Microgramma C.Presl, Tent. Pterid.: 213 (1836).
Trop. & Subtrop. America, Trop. & S. Africa, W. Indian Ocean, India. 22 GHA GUI IVO LBR NGA SIE TOG 23 CAF CMN EQG GAB GGI RWA ZAI 25 KEN TAN UGA 26 ANG MOZ ZIM 27 CPP NAT SWZ TVL 29 COM MAU MDG REU 40 IND† SRL† 78 FLA 79 MXC MXE MXG MXS MXT 80 BLZ COS GUA HON NIC PAN 81 BAH CAY CUB DOM HAI JAM LEE PUE TRT VNA WIN 82 FRG GUY SUR VEN 83 BOL CLM ECU PER 84 BZC BZE BZL BZN BZS 85 AGE AGW PAR URU TSK. [Polypodiaceae]
3 Species
Microgramma vacciniifolia (Langsd. & Fisch.) Copel., Ann. Cryptog. Phytopathol. 5: 185 (1947).
S. Caribbean to S. Trop. America. 81 JAM TRT VNA 82 VEN 83 BOL CLM ECU PER 84 BZC BZE BZL BZS 85 AGE PAR URU TSK. Epiphyte.* Polypodium vacciniifolium Langsd. & Fisch.
Pecluma M.G.Price, Amer. Fern J. 73: 109 (1983).
Trop. & Subtrop. America. 78 FLA 79 MXC MXE MXG MXI MXN MXS MXT 80 BLZ COS ELS GUA HON NIC PAN 81 BAH BER CAY CUB DOM HAI JAM LEE PUE TRT VNA WIN 82 FRG GUY SUR VEN 83 BOL CLM ECU GAL PER 84 BZC BZE BZL BZN BZS 85 AGE AGW PAR URU TSK. [Polypodiaceae]
3 Species
Pecluma filicula (Kaulf.) M.G.Price, Amer. Fern J. 73: 114 (1983).
S. Trop. America. 83 BOL CLM ECU PER 84 BZL BZS 85 AGE AGW PAR URU TSK. Epiphyte.
* Polypodium filiculum Kaulf.
Pecluma pectinatiformis (Lindm.) M.G.Price, Amer. Fern J. 73: 115 (1983).
SE & S Brazil to Argentina (Misiones, Salta) and Pacaus. 84 BZL BZS 85 AGE AGW PAR URU TSK. Epiphyte.
* Polypodium pectinatiforme Lindm.
Pleopeltis Humb. & Bonpl. ex Willd., Sp. Pl. ed. 4. 5: 211 (1810).
New World, Africa, Arabian Pen., India, Sri Lanka. 22 GUI IVO LBR NGA SIE 23 BUR CMN GGI RWA ZAI 24 ETH SOC SOM 25 KEN TAN UGA 26 ANG MLW MOZ ZIM 27 CPP LES NAT OFS SWZ TVL 28 STH 29 COM MAU† MDG REU 35 SAU YEM 40 IND SRL 61 EAS 74 ILL KAN MSO OKL 75 INI NWY OHI WVA 76 ARI 77 NWM TEX 78 ALA ARK DEL FLA GEO KTY LOU MRY MSI NCA SCA TEN VRG 79 MXC MXE MXG MXI MXN MXS MXT 80 BLZ COS CPI ELS GUA HON NIC PAN 81 BAH CAY CUB DOM HAI JAM LEE PUE TRT VNA WIN 82 FRG GUY SUR VEN 83 BOL CLM ECU GAL PER 84 BZC BZE BZL BZN BZS 85 AGE AGW CLC CLN CLS JNF PAR URU TSK. [Polypodiaceae]
6 Species
Pleopeltis hirsutissima (Raddi) de la Sota, Darwiniana 45: 239 (2007).
Bolivia to Brazil and Argentina (Misiones, Tucumán). 83 BOL 84 BZC BZE BZL BZN BZS 85 AGE AGW PAR URU TSK. Hemicr., lithophyte or epiphyte.* Polypodium hirsutissimum Raddi
Pleopeltis lepidopteris (Langsd. & Fisch.) de la Sota, Darwiniana 45: 239 (2007).
SE & S Brazil to Paraguay Uruguay and Pacaus. 84 BZL BZS 85 PAR URU TSK. Rhizome geophyte.
* Acrostichum lepidopteris Langsd. & Fisch.
Pleopeltis macrocarpa (Willd.) Kaulf., Berlin. Jahrb. Pharm. Verbundenen Wiss. 21: 41 (1820).
Mexico to S. Trop. America, Trop. Africa, Arabian Pen., S. India, Sri Lanka. 22 GUI IVO LBR NGA SIE 23 BUR CMN GGI RWA ZAI 24 ETH SOC SOM 25 KEN TAN UGA 26 ANG MLW MOZ ZIM 27 CPP LES NAT OFS SWZ TVL 28 STH 29 COM MAU† MDG REU 35 SAU YEM 40 IND SRL 79 MXC MXE MXG MXN MXS 80 COS CPI HON NIC PAN 81 CUB DOM HAI PUE 82 VEN 83 BOL CLM ECU PER 84 BZE BZL BZN BZS 85 AGE AGW CLC CLN CLS JNF URU TSK. Hemicr., lithophyte or epiphyte.* Polypodium macrocarpum Willd.
Pleopeltis minima (Bory) J.Prado & R.Y.Hirai, Amer. Fern J. 100: 191 (2011).
Bolivia to Brazil to N Argentina and Pacaus. 83 BOL 84 BZC BZE BZL BZS 85 AGE AGW PAR URU TSK. Epiphyte or lithophyte.
* Marginaria minima Bory
Pleopeltis pleopeltidis (Fée) de la Sota, Darwiniana 45: 239 (2007).
E & S Brazil to Argentina (Jujuy, Salta) and Pacaus. 84 BZE BZL BZS 85 AGW PAR URU TSK. Epiphyte or lithophyte.
* Polypodium pleopeltidis Fée
Pleopeltis pleopeltifolia (Raddi) Alston, Bol. Soc. Brot., sér. 2, 30: 21 (1956).
Brazil to Argentina (Corrientes, Formosa, Misiones) and Pacaus. 84 BZC BZE BZL BZS 85 AGE PAR URU TSK. Epiphyte or lithophyte.
* Polypodium pleopeltifolium Raddi
Rumohra Raddi, Opusc. Sci. 3: 290 (1819).
Zimbabwe to S. Africa, W. Indian Ocean, New Guinea to Australasia, Trop. & Subtrop. America. 26 ZIM 27 CPP NAT TVL 29 COM MAU MDG REU SEY 43 NWG 50 NSW QLD TAS VIC 51 CTM NZN NZS 78 FLA 80 CPI 81 BER CUB DOM HAI JAM PUE 82 GUY VEN 83 BOL CLM GAL PER 84 BZC BZE BZL BZS 85 AGE AGS CLC CLS JNF URU TSK 90 FAL TDC. [Polypodiaceae]
1 Species
Rumohra adiantiformis (G.Forst.) Ching, Sinensia 5: 70 (1934).
Trop. & Subtrop. America, Subantarctic Is., Zimbabwe to S. Africa, W. Indian Ocean, New Guinea, E & SE. Australia, New Zealand. 26 ZIM 27 CPP NAT TVL 29 COM MAU MDG REU SEY 43 NWG 50 NSW QLD TAS VIC 51 CTM NZN NZS 78 FLA 80 CPI 81 BER CUB DOM HAI JAM PUE 82 GUY VEN 83 BOL CLM GAL PER 84 BZC BZE BZL BZS 85 AGE AGS CLC CLS URU TSK 90 FAL TDC. Epiphyte.
* Polypodium adiantiforme G.Forst.
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