Research Article |
Corresponding author: Caroline Oliveira Andrino ( coliveiraandrino@gmail.com ) Academic editor: Ricarda Riina
© 2020 Caroline Oliveira Andrino, Rafael Gomes Barbosa-Silva, Juliana Lovo, Pedro Lage Viana, Marcelo Freire Moro, Daniela Cristina Zappi.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Andrino CO, Barbosa-Silva RG, Lovo J, Viana PL, Moro MF, Zappi DC (2020) Iron islands in the Amazon: investigating plant beta diversity of canga outcrops. PhytoKeys 165: 1-25. https://doi.org/10.3897/phytokeys.165.54819
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The world’s largest mineral iron province, Serra dos Carajás, is home to an open vegetation known as canga, found on top of isolated outcrops rising out of the Amazon rainforest. Over one thousand vascular plants species have been recorded in these canga sites, including 38 edaphic endemics. A new survey adds to our investigation of biogeographic relationships between sixteen canga outcrops and the effect of the distance between site pairs on the number of shared species, regional species turnover and species distribution patterns. Plant collecting expeditions to the westernmost site, the Serra de Campos of São Félix do Xingu (SFX), were carried out followed by the identification of all collected specimens and the creation of a species database, built to perform biogeographical analyses. Floristic relationships among the sites were investigated regarding their similarity, using multivariate analyses. The correlation between canga areas and species richness was tested, as well as the geographical distance between pairs of outcrops and their shared species. Vascular plants at SFX total 254 species including 17 edaphic endemics. All canga sites are grouped with 25% of minimum similarity, and the SFX falls within a large subgroup of outcrops. The total species number shared between site pairs does not change significantly with geographical distance but is positively correlated with the area of each outcrop. Meanwhile, shared endemic species numbers between site pairs decline when geographical distance increases, possibly imposed by the barrier of the rainforest. Our data suggest higher shared similarity between the largest and species-richest sites as opposed to geographically nearby sites, and provide useful insight for drafting conservation and compensation measures for canga locations. The size of the canga outcrops is associated to higher floristic diversity but connectivity among islands also plays a role in their similarity.
campo rupestre, edaphic endemism, island-like habitats, Neotropical mountains, plant species diversity, rainforest, vascular plant survey
Mountaintops are often compared to sky-islands, as their vegetation is often distinct from the surrounding lowlands (
Canga is the lateritic duricrust that covers a supergene iron ore, with poorly developed soil and moderately hard rocks that are very resistant to erosion and permeable (
Species isolation caused by environmental conditions contrasting with the surrounding forests and associated with the mosaic of different geomorphological situations in the canga creates also an abundance of micro-habitats (
The first botanical studies on the iron islands of the Serra dos Carajás began in the late 1960s. However, the floristic knowledge was not synthetized and organized until the Flora of the canga of the Serra de Carajás (FCC) project was completed in 2018 (
Due to historic reasons, collection efforts of the FCC project prioritized some areas of canga, while others still lack in-depth studies. For instance, a research in the canga of the Serra Arqueada (SA) in the municipality of Ourilândia do Norte has recently been completed (
This study aims to investigate plant distribution and biogeographical patterns that connect the island-like habitats of canga outcrops isolated within an Amazonian rainforest matrix. We evaluated species distribution in the different sites in order to observe whether canga vegetation has elevated levels of beta diversity and whether the flora of each outcrop will be more dissimilar to other outcrops as the geographical distance increases. We provided the first checklist of vascular plants growing on canga at the Serra de Campos of São Félix do Xingu (SFX), to add to the dataset we built to investigate the floristic relationship between canga areas, aiming to improve our understanding of the rich and diverse flora of the region.
The CRC are found in the region of Carajás, located in the southeast part the State of Pará (
Most of the ferruginous island complex in the southeastern Amazon is within areas protected at different levels. The Serra Norte (SN1, SN2, SN3, SN4, SN5, SN6, SN7, SN8), the Serra Sul (S11A, S11B, S11C S11D) are located in the Floresta Nacional de Carajás, which is an area of sustainable use and thus subject to anthropogenic pressures, and iron ore mining currently occurs in areas SN4, SN5 and S11D. The Serra da Bocaina and Serra do Tarzan are the only fully protected areas, and are both inserted within the Parque Nacional dos Campos Ferruginosos (PNCF). However, the Serra Arqueada and Serra de Campos of São Félix do Xingu have no legal protection.
The Serra de Campos (SFX) is a canga outcrop found in the municipality of São Félix do Xingu, southeastern Pará state, Brazilian Amazon. It represents the westernmost limit of the Serra dos Carajás, a complex of ferruginous highland outcrops that extends eastwards to the Municipality of Curionópolis, totalling 126 km2. The plateaus previously studied in the scope of the FCC project (
a Geographic location of the present study site at SFX and the other study areas from Carajás complex b aerial view of an island of canga vegetation surrounding by the rainforest (Photo: Leonardo Vianna) c Serra de Campos of São Félix do Xingu (SFX) phytophysiognomy with shrubby and grassy vegetation.
Botanical specimens from SFX deposited in herbaria prior to this study were located through an online search at the Herbarium of the Museu Paraense Emílio Goeldi (MG) and Herbário Ezechias Paulo Heringer (HEPH) (acronyms according to Thiers, continuously updated). Prior to our expeditions, specimens at MG were collected in the 1990´s by João Batista Fernandes da Silva and include the type of Mimosa dasilvae A.S.L. Silva & Secco and several gatherings of Orchidaceae, while HEPH currently holds collections made by Annajulia Elizabeth Heringer Salles and J.B.F. Silva in 2001. All materials available in these collections were analyzed and included in this study.
Four plant collecting expeditions were carried out between 2016 and 2019 (May 2016, April 2017, March 2018, October 2019), aiming to collect fertile material of all vascular species. Collecting method followed
The samples collected were identified to species by comparing their macroscopic and microscopic morphological features with available bibliography, against herbarium collections (physically and on-line) and also consulting key family specialists. Voucher specimens were deposited at MG. Only one collection number per taxon is cited in the present floristic list. A full specimen list is provided in supplement S1. Species names follow Flora do Brasil online (Flora do Brasil under construction), family delimitation followed APG IV (
Seed plant species distribution data were assembled from the FCC project (
To perform the biogeographical analysis of the CRC of the Carajás complex, the species database was used to investigate the floristic similarity and shared endemicity between different mountaintops across canga sites. Invasive exotic species recorded in each site were excluded from this analysis, as well as specimens with imprecise identification, Lycophytes, and Monilophytes. Floristic similarity between sites was calculated using a presence-absence Matrix (S2, Suppl. material
To investigate the floristic richness of sites in relation to the size of each outcrop we used the species count for each canga outcrop and, employing GIS, we calculated the area of each outcrop in square kilometres. A linear model of the recorded richness versus area of each outcrop using the ‘glm’ function with Gaussian model was prepared in R. Because the outcrops were subjected to a large collecting effort during the ‘Flora of Carajás’ Project, we assumed that they were adequately sampled. We also evaluated whether the total number of species and of endemic species shared between sites were significantly related with the geographical distance between them. We computed the centroid of each outcrop using GIS and calculated the geographical distance between the centroids of all outcrop pairs. We tested the normality of the residuals of the models with the Shapiro-Wilk test to see whether the residuals significantly departed from normality. If these did not significantly differ from normality, we accepted the p value of the model. If the residuals differed from normality, we analysed the data using non parametric Spearman’s correlation to evaluate if the correlation was significant.
This study recorded a total of 254 species, of which 248 are seed plants, five ferns and one lycophyte in the SFX (Table
Vascular plant species from Serra de Campos of São Félix do Xingu (SFX), discriminated by novelties for Flora of the canga of Carajás according to
Taxa | New for Carajás Flora | Endemic canga | Endemic SFX | Life form | Voucher |
---|---|---|---|---|---|
Lycophyte | |||||
Selaginellaceae | |||||
Selaginella radiata (Aubl.) Spring. | Herb | DCZ 4055 | |||
Monilophytes | |||||
Dennstaedtiaceae | |||||
Pteridium arachnoideum (Kauf.) Maxon | Herb | DCZ 4002 | |||
Polypodiaceae | |||||
Microgramma persicariifolia (Schrad.) C.Presl | Herb | DCZ 4066 | |||
Pleopeltis polypodioides (L.) Andrews & Windham | Herb | DCZ 3922 | |||
Serpocaulon triseriale (Sw.) A.R.Sm. | Herb | DCZ 4037 | |||
Pteridaceae | |||||
Doryopteris collina (Raddi) J.Sm. | Herb | DCZ 4040 | |||
Spermathophytes | |||||
Acanthaceae | |||||
Justicia birae A.S.Reis, F.A.Silva, A.Gil & Kameyama | Herb | MP 600 | |||
Alismataceae | |||||
Helanthium tenellum (Mart. ex Schult & Schult.f.) Britton | Herb | MP 613 | |||
Limnocharis flava (L.) Buchenau | X | Herb | PLV 6149 | ||
Anacardiaceae | |||||
Anacardium occidentale L. | Treelet | DCZ 3923 | |||
Spondias mombin L. | X | Treelet | DCZ 3921 | ||
Annonaceae | |||||
Annona sericea Dunal | X | Shrub | DCZ 4051 | ||
Annona exsucca DC. | Tree | COA 658 | |||
Guatteria procera R.E.Fr. | X | Tree | DCZ 4050 | ||
Xylopia aromatica (Lam.) Mart. | Treelet | DCZ 3970 | |||
Apocynaceae | |||||
Himatanthus cf. articulatus (Vahl) Woodson | Tree | COA 676 | |||
Mandevilla scabra (Hoffmanns. ex Roem. & Schult.) K. Schum. | Liana | DCZ 3880 | |||
Mandevilla tenuifolia (J.C. Mikan) Woodson | Herb | DCZ 3885 | |||
Matelea microphylla Morillo | X | Herb | DCZ 3942 | ||
Tabernaemontana flavicans Willd. ex Roem. & Schult. | Treelet | COA 613 | |||
Tabernaemontana macrocalyx Müll. Arg. | Treelet | COA 605 | |||
Araceae | |||||
Anthurium gracile (Rudge) Lindl. | Herb | DCZ 5017 | |||
Anthurium sp.1 | X | Herb | DCZ 3898 | ||
Arecaceae | |||||
Mauritia flexuosa Mart. | Palm | DCZ 3961 | |||
Mauritiella armata (Mart.) Burret | Palm | DCZ 3960 | |||
Oenocarpus distichus Mart. | Palm | DCZ 3948 | |||
Syagrus cocoides Mart. | Palm | DCZ 3892 | |||
Asteraceae | |||||
Emilia fosbergii Nicolson | Herb | DCZ 4046 | |||
Ichthyothere terminalis (Spreng.) S.F. Blake | Shrub | DCZ 3868 | |||
Monogereion carajensis G.M. Barroso & R.M. King | X | Herb | DCZ 3861 | ||
Riencourtia pedunculosa (Rich.) Pruski | Herb | DCZ 3924 | |||
Tilesia baccata (L.f.) Pruski | Herb | DCZ 3980 | |||
Unxia camphorata L.f. | Herb | DCZ 3941 | |||
Begoniaceae | |||||
Begonia humilis Dryand | Herb | DCZ 3973 | |||
Bignoniaceae | |||||
Adenocalymma schomburgkii (DC.) L.G.Lohmann | Liana | COA 611 | |||
Amphilophium mansoanum (DC.) L.G.Lohmann | Liana | DCZ 4025 | |||
Anemopaegma carajasense A.H. Gentry ex Firetti-Leggieri & L.G. Lohmann | X | Shrub | DCZ 3914 | ||
Anemopaegma longipetiolatum Sprague | Liana | DCZ 3867 | |||
Jacaranda ulei Bureau & K.Schum. | Shrub | DCZ 3945 | |||
Pachyptera incarnata (Aubl.) Francisco & L.G. Lohmann | Liana | DCZ 4061 | |||
Pleonotoma melioides (S.Moore) A.H.Gentry | Liana | COA 638 | |||
Pleonotoma orientalis Sandwith | Liana | DCZ 3883 | |||
Bixaceae | |||||
Cochlospermum orinocense (Kunth) Steud. | Treelet | DCZ 3875 | |||
Boraginaceae | |||||
Cordia nodosa Lam. | Tree | COA 641 | |||
Bromeliaceae | |||||
Aechmea castelnavii Baker | Herb | COA 670 | |||
Aechmea mertensii (G.Mey.) Schult. & Schult.f. | Herb | COA 673 | |||
Aechmea tocantina Baker | Herb | AHS 2194 | |||
Ananas ananassoides (Baker) L.B. Sm. | Herb | DCZ 3891 | |||
Dyckia duckei L.B.Sm. | Herb | DCZ 3872 | |||
Tillandsia adpressiflora Mez | X | Herb | DCZ 4034 | ||
Burmanniaceae | |||||
Burmannia capitata (Walter ex J.F.Gmel.) Mart. | Herb | MP 644 | |||
Burmannia flava Mart. | Herb | DCZ 3903 | |||
Cabombaceae | |||||
Cabomba furcata Schult. & Schult.f. | Herb | DCZ 3963 | |||
Commelinaceae | |||||
Commelina erecta L. | Herb | DCZ 4058 | |||
Dichorisandra hexandra (Aubl.) C.B. Clarke | Liana | DCZ 3858 | |||
Connaraceae | |||||
Rourea ligulata Baker | Shrub | COA 666 | |||
Convolvulaceae | |||||
Distimake macrocalyx (Ruiz & Pav.) A.R. Simões & Staples | X | Liana | MP 660 | ||
Ipomoea decora Meisn. | Liana | DCZ 4057 | |||
Ipomoea marabaensis D.F.Austin & Secco | Liana | DCZ 3873 | |||
Ipomoea rubens Choisy | X | Liana | MP 672 | ||
Cucurbitaceae | |||||
Gurania sinuata (Benth.) Cogn. | Herb | AHS 2167 | |||
Cyperaceae | |||||
Bulbostylis conifera (Kunth) C.B. Clarke | Herb | COA 624 | |||
Cyperus aggregatus (Willd.) Endl. | Herb | DCZ 3865 | |||
Cyperus laxus Lam. | Herb | DCZ 3957 | |||
Cyperus sesquiflorus (Torr.) Mattf. & Kük. | Herb | DCZ 4031 | |||
Cyperus sphacelatus Rottb. | Herb | DCZ 4042 | |||
Diplasia karatifolia Rich. in Pers. | X | Herb | DCZ 4032 | ||
Eleocharis flavescens (Poir.) Urb. | Herb | MP 627 | |||
Eleocharis pedrovianae C.S. Nunes, R. Trevis. & A. Gil | X | Herb | DCZ 4027 | ||
Eleocharis plicarhachis (Griseb.) Svenson | Herb | COA 678 | |||
Rhynchospora barbata (Vahl) Kunth | Herb | COA 657 | |||
Rhynchospora filiformis Vahl | Herb | DCZ 3930 | |||
Rhynchospora holoschoenoides (Rich.) Herter | Herb | MP 608 | |||
Rhynchospora seccoi C.S.Nunes, P.J.S. Silva Filho & A.Gil | Herb | DCZ 3905 | |||
Scleria cyperina Willd. ex Kunth | Herb | DCZ 3925 | |||
Scleria microcarpa Nees ex Kunth | Herb | COA 650 | |||
Dioscoreaceae | |||||
Dioscorea piperifolia Humb. & Bonpl. ex Willd. | Liana | DCZ 3884 | |||
Dioscorea trilinguis Griseb. | X | Liana | DCZ 3934 | ||
Eriocaulaceae | |||||
Eriocaulon carajense Moldenke | X | Herb | DCZ 3936 | ||
Eriocaulon cinereum R.Br. | Herb | DCZ 4049 | |||
Paepalanthus fasciculoides Hensold | Herb | DCZ 3878 | |||
Syngonanthus discretifolius (Moldenke) M.T.C. Watanabe | X | Herb | PLV 6119 | ||
Syngonanthus heteropeplus (Körn.) Ruhland | Herb | MP 659 | |||
Erythroxylaceae | |||||
Erythroxylum nelson-rosae Plowman | X | Shrub | COA 672 | ||
Erythroxylum rufum Cav. | Shrub | COA 637 | |||
Euphorbiaceae | |||||
Alchornea discolor Poeppig | Shrub | DCZ 3886 | |||
Aparisthmium cordatum (A. Juss.) Baill. | Tree | DCZ 3997 | |||
Astraea lobata (L.) Klotzsch | Shrub | DCZ 3955 | |||
Mabea angustifolia Spruce ex Benth. | Shrub | DCZ 3987 | |||
Manihot quinquepartita Huber ex D.J.Rogers | Shrub | DCZ 3954 | |||
Manihot tristis Müll.Arg. | Shrub | MP 666 | |||
Maprounea brasiliensis A.St.-Hil. | X | Shrub | DCZ 3991 | ||
Fabaceae | |||||
Abrus melanospermus Hassk. | Liana | DCZ 3912 | |||
Aeschynomene sensistiva var. hispidula (Kunth) Rudd | Subshrub | DCZ 4024 | |||
Bauhinia pulchella Benth. | Shrub | DCZ 3869 | |||
Camptosema ellipticum (Desv.) Burkart | Shrub | DCZ 3907 | |||
Centrosema carajasense Cavalcante | Herb/Liana | DCZ 4007 | |||
Chamaecrista desvauxii (Collad.) Killip | Subshrub | DCZ 3946 | |||
Clitoria falcata Lam. | Liana | DCZ 3917 | |||
Crotalaria maypurensis Kunth | Shrub | DCZ 3881 | |||
Dioclea apurensis Kunth | Liana | DCZ 3919 | |||
Inga calantha Ducke | X | Tree | COA 600 | ||
Inga heterophylla Willd | X | Tree | DCZ 4036 | ||
Inga leiocalycina Benth. | X | Tree | MP 598 | ||
Mimosa dasilvae A.S.L. Silva & Secco | X | X | X | Subshrub | COA 622 |
Mimosa guilandinae var. spruceana (Benth.) Barneby | Shrub | COA 668 | |||
Mimosa skinneri var. carajarum Barneby | X | Herb | DCZ 3860 | ||
Mimosa somnians Humb. & Bonpl. ex Willd. | Subshrub | DCZ 3876 | |||
Mimosa xanthocentra Mart. | Tree | PLV 6158 | |||
Parkia platycephala Benth. | Shrub | DCZ 4013 | |||
Periandra mediterranea (Vell.) Taub. | Shrub | DCZ 3902 | |||
Senegalia multipinnata (Ducke) Seigler & Ebinger | Treelet | COA 603 | |||
Stylosanthes capitata Vogel | Subshrub | DCZ 3977 | |||
Tachigali vulgaris L.F.G.Silva & H.C.Lima | Tree | COA 655 | |||
Gentianaceae | |||||
Schultesia benthamiana Klotzsch ex Griseb. | Herb | DCZ 3928 | |||
Heliconiaceae | |||||
Heliconia psittacorum L.f. | X | Herb | MP 671 | ||
Hypericaceae | |||||
Vismia gracilis Hieron | Treelet | COA 654 | |||
Iridaceae | |||||
Cipura xanthomelas Maxim. ex Klatt | Herb | DCZ 3899 | |||
Lamiaceae | |||||
Amasonia lasiocaulos Mart. & Schau ex Schau. | Subshrub | DCZ 3947 | |||
Hyptis atrorubens Poit. | Herb | DCZ 3981 | |||
Mesosphaerum pectinatum (L.) Kuntze | Herb | MN 697 | |||
Mesosphaerum suaveolens (L.) Kuntze | Herb | DCZ 4048 | |||
Vitex panshiniana Moldenke | X | Tree | DCZ 4053 | ||
Lauraceae | |||||
Cassytha filiformis L. | Parasite | DCZ 3874 | |||
Dicypellium aff. caryophyllaceum (Mart.) Nees | X | X | Shrub | PLV 6100 | |
Mezilaurus itauba (Meisn.) Taub. ex Mez | Shrub | DCZ 4001 | |||
Rhodostemonodaphne praeclara (Sandwith) Madriñán | X | Tree | DCZ 3983 | ||
Lentibulariaceae | |||||
Utricularia neottioides A.St-Hil & Girard | Herb | MP 664 | |||
Utricularia pusilla Vahl | Herb | DCZ 3904 | |||
Utricularia subulata L. | Herb | PLV 6139 | |||
Loranthaceae | |||||
Passovia pedunculata (Jacq.) Kuijt | Parasite | DCZ 3909 | |||
Psittacanthus eucalyptifolius (Kunth) G. Don | Parasite | DCZ 4056 | |||
Lythraceae | |||||
Cuphea annulata Koehne | Subshrub | DCZ 3864 | |||
Cuphea carajasensis Lourteig | X | Shrub | COA 616 | ||
Malpighiaceae | |||||
Banisteriopsis malifolia (Nees & Mart.) B.Gates | Shrub | MN 743 | |||
Banisteriopsis stellaris (Griseb.) B.Gates | Liana | DCZ 3863 | |||
Byrsonima chrysophylla Kunth | Shrub | DCZ 3929 | |||
Heteropterys nervosa A.Juss. | Liana | COA 645 | |||
Malvaceae | |||||
Waltheria indica L. | X | Shrub | DCZ 4064 | ||
Marantaceae | |||||
Monotagma plurispicatum (Körn.) K.Schum. | Herb | DCZ 4000 | |||
Marcgraviaceae | |||||
Norantea guianensis Aubl. | Shrub | DCZ 3887 | |||
Melastomataceae | |||||
Bellucia grossularioides (L.) Triana | X | Shrub | DCZ 3995 | ||
Brasilianthus carajensis Almeda & Michelangeli | Herb | DCZ 3877 | |||
Clidemia capitellata (Bonpl.) D.Don | Shrub | DCZ 4020 | |||
Miconia alternans Naudin | Shrub | DCZ 4021 | |||
Miconia heliotropoides Triana | Shrub | DCZ 4008 | |||
Nepsera aquatica (Aubl.) Naudin | Herb | COA 649 | |||
Pleroma carajasense K.Rocha, R.Goldenb. & F.S.Mey | X | Shrub | DCZ 3910 | ||
Pterolepis trichotoma (Rottb.) Cogn. | Herb | DCZ 4019 | |||
Tibouchina edmundoi Brade | Shrub | DCZ 3932 | |||
Menispermaceae | |||||
Abuta grandifolia (Mart.) Sandwith | Shrub | COA 646 | |||
Cissampelos andromorpha DC. . | Liana | COA 663 | |||
Metteniusaceae | |||||
Emmotum nitens (Benth.) Miers | Shrub | MP 601 | |||
Myrtaceae | |||||
Eugenia punicifolia (Kunth) DC. | Shrub | DCZ 3894 | |||
Myrcia cuprea (O.Berg.) Kiaersk. | Shrub | COA 639 | |||
Myrcia splendens (Sw.) DC. | Shrub | DCZ 3965 | |||
Myrciaria floribunda (H.West ex Willd.) O.Berg | Shrub | DCZ 3915 | |||
Myrciaria glomerata O.Berg | X | Shrub | DCZ 4010 | ||
Ochnaceae | |||||
Ouratea castaneifolia (DC.) Engl. | Treelet | DCZ 3920 | |||
Ouratea cearensis (Tiegh.) Sastre & Offroy | X | Shrub | COA 604 | ||
Ouratea racemiformis Ule | Shrub | DCZ 4033 | |||
Onagraceae | |||||
Ludwigia cf. latifolia (Benth.) H.Hara | X | Subshrub | DCZ 3967 | ||
Ludwigia nervosa (Poir.) H.Hara | Shrub | COA 674 | |||
Orchidaceae | |||||
Catasetum boyi Mansf. | X | Herb | JBFS 648 | ||
Catasetum discolor (Lindl.) Lindl. | Herb | DCZ 4030 | |||
Cyrtopodium andersonii (Lamb. ex Andrews) R.Br. | Herb | COA 643 | |||
Encyclia chloroleuca (Hook.) Neum. | X | Herb | JBFS 540 | ||
Epidendrum strobiliferum Rchb.f. | X | Herb | COA 667 | ||
Erycina pusilla (L.) N.H.Williams & M.W.Chase | Herb | JBFS 498 | |||
Habenaria nuda Lindl | Herb | MP 609 | |||
Habenaria orchiocalcar Hoehne | X | Herb | JBFS 219 | ||
Polystachya concreta (Jacq.) Garay & H.R.Sweet | Herb | COA 669 | |||
Rodriguezia lanceolata Ruiz & Pav. | X | Herb | COA 665 | ||
Scaphyglottis cf. livida | Herb | COA 671 | |||
Sobralia liliastrum Salzm. ex Lindl. | Herb | DCZ 3888 | |||
Orobanchaceae | |||||
Buchnera carajasensis Scatigna & N.Mota | X | Herb | DCZ 3931 | ||
Passifloraceae | |||||
Passiflora ceratocarpa F. Silveira | Liana | DCZ 4060 | |||
Passiflora picturata Ker Gawl. | X | Liana | DCZ 3976 | ||
Passiflora tholozanii Sacco | Liana | COA 612 | |||
Phyllanthaceae | |||||
Phyllanthus hyssopifolioides Kunth. | Herb | DCZ 4028 | |||
Phyllanthus minutulus Müll.Arg. | Herb | DCZ 4026 | |||
Phytolaccaceae | |||||
Phytolacca thyrsiflora Fenzl ex J. Schmidt | Herb | DCZ 4041 | |||
Piperaceae | |||||
Peperomia albopilosa D. Monteiro | X | Herb | PLV 6169 | ||
Peperomia magnoliifolia (Jacq.) A.Dietr. | Herb | COA 647 | |||
Plantaginaceae | |||||
Scoparia dulcis L. | Herb | DCZ 4065 | |||
Poaceae | |||||
Acroceras zizanioides (Kunth) Dandy | Herb | DCZ 4022 | |||
Andropogon bicornis L. | Herb | DCZ 3950 | |||
Axonopus cf. longispicus (Döll) Kuhlm. | Herb | DCZ 4023 | |||
Axonopus rupestris Davidse | Herb | DCZ 3896 | |||
Eleusine indica (L.) Gaertn.* | Herb | DCZ 4045 | |||
Hildaea parvispiculata C. Silva & R.P. Oliveira | Herb | PLV 6124 | |||
Ichnanthus calvescens (Nees ex Trin.) Döll | Herb | DCZ 4011 | |||
Luziola peruviana Juss. ex J.F.Gmel. | Herb | DCZ 3918 | |||
Melinis minutiflora P.Beauv.* | Herb | COA 640 | |||
Mesosetum cayennense Steud. | Herb | PLV 6117 | |||
Oryza glumaepatula Steud. | Herb | BFF 634 | |||
Paspalum axillare Swallen | Herb | PLV 6130 | |||
Paspalum foliiforme S.Denham | Herb | DCZ 3916 | |||
Paspalum reticulinerve Renvoize | Herb | PLV 6166 | |||
Rhytachne gonzalezii Davidse | Herb | PLV 6127 | |||
Rugoloa pilosa (Sw.) Zuloaga | Herb | DCZ 3964 | |||
Steinchisma laxum (Sw.) Zuloaga | Herb | COA 677 | |||
Taquara micrantha (Kunth) I.L.C.Oliveira & R.P.Oliveira | Herb | DCZ 3999 | |||
Trachypogon spicatus (L.f.) Kuntze | Herb | DCZ 3944 | |||
Trichanthecium cf. arctum (Swallen) Zuloaga & Morrone | Herb | DCZ 3913 | |||
Urochloa maxima (Jacq.) R.D. Webster* | Herb | DCZ 3951 | |||
Polygalaceae | |||||
Bredemeyera divaricata (DC.) J.F.B. Pastore | Shrub | DCZ 3911 | |||
Caamembeca spectabilis (DC.) J.F.B. Pastore | Subshrub | COA 642 | |||
Polygala adenophora DC. | Herb | DCZ 3900 | |||
Portulacaceae | |||||
Portulaca sedifolia N.E.Br. | Herb | DCZ 3862 | |||
Primulaceae | |||||
Cybianthus detergens Mart. | Shrub | DCZ 4062 | |||
Proteaceae | |||||
Roupala montana Aubl. | Shrub | DCZ 4063 | |||
Rhamnaceae | |||||
Gouania pyrifolia Reissek | X | Liana | DCZ 3953 | ||
Rubiaceae | |||||
Alibertia edulis (Rich.) A. Rich. ex DC. | Shrub | DCZ 4035 | |||
Borreria alata (Aubl.) DC. | Herb | DCZ 3866 | |||
Borreria carajasensis E.L. Cabral & L.M. Miguel | X | Subshrub | DCZ 3859 | ||
Borreria semiamplexicaulis E.L.Cabral | Herb | DCZ 3938 | |||
Cordiera myrciifolia (K.Schum.) C.H.Perss. & Delprete | Shrub | DCZ 3971 | |||
Coutarea hexandra (Jacq.) K.Schum. | X | Shrub | COA 610 | ||
Guettarda argentea Lam. | Shrub | COA 602 | |||
Palicourea guianensis Aubl. | Treelet | DCZ 4052 | |||
Perama carajensis J.H. Kirkbr. | X | Herb | DCZ 3879 | ||
Psychotria colorata (Willd. ex Schult.) Mull. Arg. | Herb | DCZ 4017 | |||
Psychotria hoffmannseggiana (Willd. ex Schult.) Mull. Arg. | Subshrub | COA 601 | |||
Sabicea grisea Cham. & Schltdl. | Liana | DCZ 3901 | |||
Rutaceae | |||||
Dictyoloma vandellianum A. Juss. | Treelet | DCZ 3975 | |||
Ertela trifolia (L.) Kuntze | Subshrub | COA 607 | |||
Pilocarpus microphyllus Stapf ex Wardlew. | Shrub | COA 653 | |||
Salicaceae | |||||
Casearia arborea (Rich.) Urb. | Tree | DCZ 3982 | |||
Casearia javitensis Kunth | Shrub | DCZ 4014 | |||
Sapindaceae | |||||
Allophylus semidentatus (Miq.) Radlk. | X | Shrub | DCZ 3959 | ||
Paullinia stellata Radlk. | X | Liana | DCZ 4044 | ||
Pseudima frutescens (Aubl.) Radlk. | X | Shrub | PLV 6151 | ||
Serjania lethalis A.St.-Hil. | Liana | DCZ 3996 | |||
Sapotaceae | |||||
Pouteria ramiflora (Mart.) Radlk. | Treelet | DCZ 3969 | |||
Simaroubaceae | |||||
Simaba guianensis Aubl. | Shrub | DCZ 3984 | |||
Simarouba amara Aubl. | Shrub | DCZ 3985 | |||
Siparunaceae | |||||
Siparuna ficoides S.S.Rener & Hausner | Treelet | COA 660 | |||
Smilacaceae | |||||
Smilax irrorata Mart. ex Griseb | Liana | DCZ 3935 | |||
Solanaceae | |||||
Solanum americanum Mill. | Herb | DCZ 4059 | |||
Solanum crinitum Lam. | Treelet | COA 623 | |||
Trigoniaceae | |||||
Trigonia nivea Cambess. | Liana | COA 651 | |||
Turneraceae | |||||
Turnera glaziovii Urb | Shrub | DCZ 4012 | |||
Turnera laciniata Arbo | Herb | DCZ 3993 | |||
Turnera melochioides Cambess. | Shrub | PLV 6160 | |||
Urticaceae | |||||
Cecropia palmata Willd. | Tree | COA 664 | |||
Velloziaceae | |||||
Vellozia glauca Pohl | Herb | DCZ 3890 | |||
Verbenaceae | |||||
Lantana trifolia L. | X | Shrub | MN 755 | ||
Lippia grata Schauer | Shrub | DCZ 3871 | |||
Stachytarpheta cayennensis (Rich.) Vahl | Subshrub | COA 608 | |||
Vitaceae | |||||
Cissus erosa Rich. | Liana | DCZ 3882 | |||
Vochysiaceae | |||||
Qualea parviflora Mart. | Tree | MP 624 | |||
Xyridaceae | |||||
Xyris brachysepala Kral | X | Herb | PLV 6125 | ||
SPECIES TOTAL (254) | 36 | 17 | 2 |
Representative species of canga in new dataset, SFX a Axonopus longispicus (Döll) Kuhlm b Dicypellium aff. caryophyllaceum (Mart.) Nees c Inga heterophylla Willd d Ipomoea decora Meisn e Matelea microphylla Morillo f Mimosa dasilvae A.S.L. Silva & Secco g Nepsera aquatica (Aubl.) Naudin h Ouratea cearensis (Tiegh.) Sastre & Offroy i Pachyptera incarnata (Aubl.) Francisco & L.G. Lohmann j Passifora picturata Ker Gawl. k Phyllanthus minutulus Mull.Arg. l Rodriguezia lanceolata Ruiz & Pav.
Among the 38 edaphic endemic species of canga, defined according to
Areas compared by this study, respective area codes used in the multivariate analysis and number of angiosperms species recorded for each area. Serra de Campos of São Félix do Xingu (SFX) data is produced by this study, ARQ-CAN data is available in
Area code | Area | Species | Cumulative species |
---|---|---|---|
ARQ | Serra Arqueada | 149 | 149 |
S11A | Serra dos Carajás – Serra Sul 11A | 230 | 535 |
S11B | Serra dos Carajás – Serra Sul 11B | 201 | |
S11C | Serra dos Carajás – Serra Sul 11C | 180 | |
S11D | Serra dos Carajás – Serra Sul 11D | 428 | |
SN1 | Serra dos Carajás – Serra Norte 1 | 383 | 643 |
SN2 | Serra dos Carajás – Serra Norte 2 | 125 | |
SN3 | Serra dos Carajás – Serra Norte 3 | 218 | |
SN4 | Serra dos Carajás – Serra Norte 4 | 308 | |
SN5 | Serra dos Carajás – Serra Norte 5 | 293 | |
SN6 | Serra dos Carajás – Serra Norte 6 | 99 | |
SN7 | Serra dos Carajás – Serra Norte 7 | 112 | |
SN8 | Serra dos Carajás – Serra Norte 8 | 101 | |
SB | Serra dos Carajás – Serra da Bocaina | 223 | 336 |
ST | Serra dos Carajás – Serra do Tarzan | 211 | |
SFX | Serra de Campos – São Félix do Xingu | 248 | 248 |
Around 25% (60) of the 248 angiosperms registered for SFX are restricted to the Amazonian Rainforest biome, but the majority of the flora is widely distributed in open habitats throughout South America.
Regarding the phytophysiognomies listed by
The biogeographical database from the CRC of the Carajás complex was updated by our study (see supplementary data) and includes now a total of 893 angiosperms distributed in 121 families and 441 genera. For the Carajás flora (FCC), Poaceae was the most species-rich family (75 species in the FCC), followed by Fabaceae (66 spp.), Cyperaceae (57 spp.), Rubiaceae (49 spp.), and Melastomataceae (40 spp.). The richest genera were Rhynchospora (24 spp.), Miconia (18 spp.), Paspalum and Solanum (17 spp. each), Myrcia and Ipomoea (13 spp. each), while 64% (284 genera) were represented by only a single species. The inclusion of SFX in our database increased the number of known taxa by 18 genera and 37 species not previously recorded for the canga of Carajás.
The mean angiosperm species richness for each outcrop of the Serra dos Carajás was 218 species. The NMDS and UPGMA analyses included 3451 records of 893 species across 16 sites (Fig.
Species richness was significantly correlated with site area (r = 0.806094, P = 0.001548). The larger the area of each individual mountaintop (site), the larger the number of species recorded. The total number of shared species between mountaintop outcrops did not differ significantly with geographical distance across sites (r = -0.16; P = 0.08). There was a tendency of distant sites to share less species, but this trend was not significant. When the residuals of this model were evaluated they significantly departed from normality. Spearman’s correlation was not significant either (p-value = 0.2972). However, when focusing on the number of shared endemic edaphic species versus the geographical distance between sites, we found a significant correlation, where closer sites shared more edaphic endemic species than with more distant sites (r = -0.45872; P = 1.37e-07) (Fig.
a Species richness plotted against area of Carajás. Pearson correlation coefficients: r = 0.806094, P = 0.001548 b the number of species shared between site pairs does not change significantly with geographical distance between regions. r = -0.16; P = 0.08 c the number of shared endemic species between site pairs declines with geographical distance between regions. r= -0.45872; P = 1.37e-07.
Regarding the total of species of the canga, the Carajás iron islands share an average of 40% of their flora with each other. SFX has, on average, 30% of shared species with each other area. The percentage of similarity between sites was a minimum of 30% and a maximum of 55%.
The most species-rich families and genera found in the SFX coincide with those found in the Flora das cangas de Carajás (
There is a large turnover of species between outcrops (
As an example, only three species were recorded in all surveyed areas: the widely distributed Riencourtia pedunculosa, an Asteraceae common in open areas in the Amazon (Flora do Brasil under construction,
Some widely distributed species from the canga of Carajás, found at more than 10 of the 16 sites surveyed, were not recorded at SFX. The absence of the common treelets Callisthene microphylla Warm. and Mimosa acutistipula var. ferrea Barneby (
The canga is typically a mosaic of different vegetation types (
Despite having the lowest number of species registered in the FCC, the hydromorphic vegetation found atop the plateaus is the habitat with the highest proportion of exclusive species (
As a relatively large canga site isolated from the active iron mines further to the east, the SFX has been found to harbour a rich and unique vegetation, representing a suitable area for the implementation of conservation strategies. On the other hand, this canga outcrop is currently threatened by surrounding deforestation, land transformation and frequent fires, and is not included within any type of protected area.
The mosaic of landscapes typical of CRC of Carajás may also explain the low floristic similarity between the sites. The number of shared species represents less than half the local richness from each site separately. This brings attention to the high beta diversity among sites (
The greater similarity between SFX, SB and ST, along with Serra Sul (S11A, S11B, S11C, and S11D) and SN1, SN3, SN4 and SN5 reflected in the UPGMA clustering patterns (Fig.
The low number of species restricted to the Amazon (25%) and the high number of species widely distributed in South America (75%) recorded at SFX, may explain the discrepancy in the correlation between shared species and distance being negative when all species are considered, whereas it is positive for endemic species only. On a macro-scale, the majority of the species recorded in SFX have a broad distribution, occurring beyond the Amazon Rainforest, and the distance factor between different outcrops may not matter so much. On the other hand, when observing only the species endemic to Carajás, and especially edaphic endemic species, the trend is the opposite, possibly due to the local scale of observation, as elsewhere the distance between areas tends to affect the floristic similarity between island vegetations (
Another factor that may have an impact on the contrasting effects of floristic similarity vs. distance from canga islands is the different environmental requirements of herbs, shrubs and trees, that shape their biogeographical patterns and affect species-area and richness-environment relationships (
Recent analyses of open vegetation in the Amazon reinforce the insular character of Amazonian canga and their low similarity to other vegetation types in the Amazonian biome (
Species richness of the iron islands outcrops of Carajás complex (bold diagonal) along with the number of shared species (above diagonal) and distance in kilometres (below diagonal) between the centroid sites; an estimated area for each site is provided.
Sites | Area (km2) | SB | ST | ARQ | S11A | S11B | S11C | S11D | SFX | SN1 | SN2 | SN3 | SN4 | SN5 | SN6 | SN7 | SN8 |
---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|
SB | 19.98 | 221 | 100 | 47 | 79 | 80 | 75 | 135 | 85 | 124 | 46 | 84 | 108 | 101 | 56 | 57 | 56 |
ST | 8.3 | 24 | 209 | 48 | 88 | 90 | 80 | 138 | 84 | 119 | 59 | 87 | 102 | 105 | 55 | 59 | 53 |
ARQ | 1.27 | 140 | 116 | 149 | 52 | 44 | 45 | 80 | 70 | 75 | 30 | 52 | 77 | 62 | 30 | 29 | 32 |
S11A | 15.27 | 59 | 24 | 92 | 228 | 139 | 119 | 170 | 96 | 143 | 59 | 89 | 116 | 101 | 56 | 54 | 53 |
S11B | 8.44 | 54.6 | 30.8 | 82 | 4.5 | 199 | 107 | 147 | 77 | 120 | 53 | 81 | 96 | 99 | 49 | 52 | 48 |
S11C | 6.26 | 52.5 | 28.8 | 85 | 10 | 4.5 | 177 | 140 | 83 | 110 | 46 | 72 | 101 | 91 | 49 | 41 | 50 |
S11D | 16.41 | 47 | 24.4 | 92.3 | 15.7 | 9.8 | 5.7 | 424 | 141 | 222 | 80 | 134 | 189 | 168 | 75 | 80 | 72 |
SFX | 9.04 | 217 | 193 | 79.5 | 158 | 162 | 165 | 170 | 239 | 131 | 48 | 82 | 111 | 95 | 52 | 44 | 51 |
SN1 | 11.81 | 52 | 37.7 | 111 | 37 | 38 | 40 | 42 | 180 | 381 | 98 | 154 | 183 | 174 | 77 | 71 | 78 |
SN2 | 0.86 | 46.8 | 32.8 | 113 | 36.8 | 37.1 | 39.3 | 40 | 184 | 5.18 | 124 | 69 | 73 | 71 | 40 | 34 | 44 |
SN3 | 2.1 | 44.7 | 32 | 117.5 | 40.2 | 40.1 | 42 | 42.2 | 188 | 8.1 | 3.8 | 217 | 129 | 103 | 71 | 60 | 59 |
SN4 | 14.83 | 38 | 25 | 117.4 | 37.5 | 36.4 | 37.7 | 37 | 189 | 13.7 | 8.6 | 7.4 | 305 | 181 | 74 | 65 | 81 |
SN5 | 8.26 | 32.36 | 22.75 | 122 | 41 | 39 | 40 | 38.53 | 195 | 19.78 | 14.6 | 12.4 | 6.2 | 289 | 63 | 54 | 69 |
SN6 | 0.97 | 35.29 | 22.46 | 118 | 37.3 | 35.8 | 36.7 | 35.7 | 190 | 16 | 11 | 10 | 3 | 4 | 99 | 40 | 42 |
SN7 | 0.34 | 33 | 19 | 117 | 35.7 | 33.8 | 34 | 33.1 | 190.5 | 18 | 14 | 13 | 6 | 5 | 3 | 112 | 46 |
SN8 | 2.69 | 30 | 17 | 119 | 37 | 34.7 | 35 | 33 | 192 | 22 | 17 | 16 | 8.8 | 6 | 5.7 | 3.3 | 100 |
Different evolutionary processes of the species occurring in CRC may also have led to different floristic composition in the outcrops. Although evolutionary studies involving species of canga in the Brazilian Amazon are just beginning (
Endemic edaphic species of the iron islands outcrops of Carajás complex (bold diagonal) along with the number of shared endemic species (above diagonal) and distance in kilometres (below diagonal) between the centroid sites.
Sites | SB | ST | ARQ | S11A | S11B | S11C | S11D | SFX | SN1 | SN2 | SN3 | SN4 | SN5 | SN6 | SN7 | SN8 |
---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|
SB | 20 | 15 | 3 | 17 | 15 | 16 | 19 | 11 | 18 | 11 | 15 | 15 | 13 | 11 | 11 | 12 |
ST | 24 | 16 | 2 | 14 | 13 | 14 | 15 | 9 | 15 | 9 | 12 | 11 | 11 | 9 | 10 | 10 |
ARQ | 140 | 116 | 7 | 5 | 4 | 5 | 7 | 5 | 6 | 3 | 4 | 5 | 3 | 2 | 2 | 4 |
S11A | 59 | 24 | 92 | 24 | 17 | 21 | 22 | 14 | 21 | 10 | 16 | 17 | 13 | 11 | 9 | 12 |
S11B | 54.6 | 30.8 | 82 | 4.5 | 18 | 18 | 19 | 10 | 15 | 14 | 14 | 13 | 12 | 10 | 8 | 10 |
S11C | 52.5 | 28.8 | 85 | 10 | 4.5 | 21 | 21 | 13 | 11 | 10 | 15 | 15 | 13 | 10 | 9 | 12 |
S11D | 47 | 24.4 | 92.3 | 15.7 | 9.8 | 5.7 | 25 | 14 | 21 | 11 | 18 | 19 | 14 | 12 | 12 | 14 |
SFX | 217 | 193 | 79.5 | 158 | 162 | 165 | 170 | 17 | 13 | 9 | 13 | 12 | 8 | 9 | 7 | 9 |
SN1 | 52 | 37.7 | 111 | 37 | 38 | 40 | 42 | 180 | 29 | 15 | 20 | 22 | 19 | 13 | 12 | 16 |
SN2 | 46.8 | 32.8 | 113 | 36.8 | 37.1 | 39.3 | 40 | 184 | 5.18 | 16 | 15 | 14 | 14 | 11 | 8 | 12 |
SN3 | 44.7 | 32 | 117.5 | 40.2 | 40.1 | 42 | 42.2 | 188 | 8.1 | 3.8 | 23 | 20 | 15 | 15 | 12 | 15 |
SN4 | 38 | 25 | 117.4 | 37.5 | 36.4 | 37.7 | 37 | 189 | 13.7 | 8.6 | 7.4 | 24 | 18 | 14 | 12 | 17 |
SN5 | 32.36 | 22.75 | 122 | 41 | 39 | 40 | 38.53 | 195 | 19.78 | 14.6 | 12.4 | 6.2 | 20 | 11 | 9 | 15 |
SN6 | 35.29 | 22.46 | 118 | 37.3 | 35.8 | 36.7 | 35.7 | 190 | 16 | 11 | 10 | 3 | 4 | 15 | 8 | 10 |
SN7 | 33 | 19 | 117 | 35.7 | 33.8 | 34 | 33.1 | 190.5 | 18 | 14 | 13 | 6 | 5 | 3 | 14 | 10 |
SN8 | 30 | 17 | 119 | 37 | 34.7 | 35 | 33 | 192 | 22 | 17 | 16 | 8.8 | 6 | 5.7 | 3.3 | 17 |
This is the most complete study analysing a database of canga outcrop islands in the Amazon thus far. Our data suggest higher shared similarity between largest sites and higher species richness. We show that species richness in these vegetation islands reveals complex biogeographic patterns and relatively high beta diversity. Outcrop size seemed to be more important than geographical proximity between outcrops, and this should be taken into account when drafting conservation and compensation measures for the canga. There are still inaccessible canga outcrops towards the north of the state of Pará that remain unexplored, and their study would certainly yield interesting information to be added to the present findings.
We are grateful to the Museu Paraense Emílio Goeldi (MPEG) and Instituto Tecnológico Vale (ITV) for essential infrastructure and support for this project, and to Priscila O. Rosa, from the Herbarium HEPH, for providing specimen images. We also acknowledge the financial support provided by the Conselho Nacional de Desenvolvimento Científico e Tecnológico (CNPq) for grants for COA and RGBS, and CAPES (JL). DCZ currently holds a research grant from CNPq. Invaluable help was provided by specialist botanists Aline Stadnik, Ana Carolina Mezzonato, Beatriz Gomes, Edgar Afonso, Edley Pessoa, Jovani Pereira, Layla Schneider, Matheus Cota, Mayara Pastore, Paulo Gonella, and Valdir Silva-Junior in specimen naming. We thank Nigel P. Taylor for revising the English. We also thank our colleague Alice Hiura for technical support and Fernando Marino Gomes dos Santos for critical reading.
Data availability statement: All supplementary data can be accessed at figshare repository: https://doi.org/10.6084/m9.figshare.12053487
Investigating plant beta diversity of canga outcrops
Data type: species data