carvalhoecarvalho-2012

8/11/2019 CarvalhoeCarvalho-2012

1/10

146 E&G / Vol. 61 / No. 2 / 2012 / 146155 / DOI 10.3285/eg.61.2.03 / Authors / Creative Commons Attribution License

1 Introduction

On a geomorphologi al viewpoint, the Amazon rainforestan be cara terized by its lowland relief and extensive for-

ested areas, by the dicotomy between main allocthonousand small autocthonous rivers, as well as ombining latosoland podzol low fertility soils. Te annual thermal range isrelatively homogeneous, 24C to 26C; the rainfall is hetero-geneous, 1750 to 2300 mm/year, outside the Andes, whicis around 7000 mm/year; and the vegetation is a omplexnet distributed over periodi ally ooded forest and upland,vrzea and terra rme respe tively (V et al., 2000;A S , 2003). Tese ombined geomorphologi al fea-tures form an area of approximately 7 million km, alledthe Amazon morpho limati domain. What in Brazil we alla morpho limati domain is an area of sub- ontinental di-mensions, with cara teristi pa erns of relief, drainage, li-mate, soils and vegetation (A S , 1967).

One important feature of the Amazon morpho limatidomain is the physiognomy of its vegetation, whic an beopen (s rubs, herbs and small trees) or losed (tall trees, withsome emerging). aking only this aspe t into a ount, the

E&G Quaternary Science JournalVolume 61 / Number 2 / 2012 / 146155 / DOI 10.3285/eg.61.2.03www.quaternary-science.netInterrelation of geomorphology and fauna of Lavrado region in

Roraima, Brazil suggestions for future studies

Thiago Morato de Carvalho, Celso Morato de Carvalho

How to cite: C , . M., C , C. M. (2012): Interrelation of geomorphology and fauna of Lavrado region in Roraima, Brazil sug-gestions for future studies. E&G Qaternary S ien e Journal, 61 (2): 146158. DOI: 10.3285/eg.61.2.03

Abstract: Te authors dis uss the relevan e of geomorphology and biology intera tion under the on epts of the Brazilian morpho limatidomains. Te dis ussion is fo used on biogeographi al and e ologi al aspe ts. Te open areas of Roraima the lavrado lo alizedbetween Brazil, Venezuela and Guyana, in the Northern portion of the Amazon morpho limati domain, is the region where thepresent ase study was arried out. Remote sensing tecniques were applied to determine the relief and eld biology cara teriza-tion. Te generated produ ts were useful for des ribing the habitats and lo al distribution of the lavrados fauna.

Die Wechselbeziehung von Geomorphologie und Fauna in der Lavrado Region in Roraima, Brasilien: Vorschlge fr zuknf-tige Studien

Kurzfassung: In der vorgelegten Arbeit wird die Abhngigkeit von Geomorphologie und biologiscen Interaktion unter Verwendung des Kon-zeptes morphoklimatiscer Regionen Brasiliens vorgestellt. Die Diskussion fokussiert hierbei auf biogeographisce und kologisceAspekte. Die vorgelegte Studie wurde in den offenen Bereicen von Roraima Lavrado zwiscen Brasilien, Venezuela und Gu-yana durcgefhrt. Dieses Gebiet liegt im nrdlicen eil der morphoklimatiscen Region Amazoniens. Zur Anwendung kamen

ecniken der Fernerkundung, um das Relief der Region zu ermi eln und biologisce Charakterisierungen durczufhren. Diehierdurc erzielten Ergebnisse wurden genutzt, um Lebensrume der Region und die Verteilung der Lavrado Fauna zu bescreiben.

Keywords: Biogeomorphology, Amazon morphoclimatic domain, Roraima, lavrado

Addresses of authors: Tiago Morato de Carvalho*, Celso Morato de Carvalho**, National Institute of Amazonian Researc (Instituto Na ional dePesquisas da Amaznia - INPA) Boa Vista, Roraima, Brazil. ZipCode 69301-150. *tmorato@infonet. om.br; ** [email protected]

open areas that o ur in the Amazon region an be quitesimilar to those o urring in other domains, for example, theopen vegetation ofcerrado in the Central Brazilian e osys-tem, the BolivianChaco or the lhanos in Venezuela. How-ever, there are many e ologi al and physiologi al differen esbetween these open formations, suc as oristi omposi-tion, soil formation, geomorphologi al genesis, drainage and

limate (V C , 1991; E , 1992, 1994).We an fo us on this physiognomi dicotomous prop-

erty of the Amazon vegetation with different lenses, depend-ing of the goal. From the biogeographi al viewpoint, for in-stan e, these two morphologi al aspe ts of vegetation, openand losed areas, are important for understanding the distri-bution of organisms, prin ipally when we onsider the pul-sation of the forest over the last 20.000 years the open areasentering the forest during the Pleisto ene gla ial dry periodsand the expansion of the forest during the intergla ial wetperiods throughout South Ameri an e osystems (V ,

1988; A S , 1977; P et al., 2009).In the Brazilian Amazon there are expressive open veg-etation areas in the states of Par, Amap and Roraima, o -

urring as en laves inside extensive forested areas (M

GEOZON SCIENCE MEDIA

ISSN 0424-7116


2/10

147E&G / Vol. 61 / No. 2 / 2012 / 146155 / DOI 10.3285/eg.61.2.03 / Authors / Creative Commons Attribution License

P , 1974; C , 2009; V , 1992) and alongthe major rivers, suc as the rombetas (E , 1960), Negro(D B , 1953), in the mouth of the apajs (R

, 1975) and in the Madeira (M P , 1974).Tese open areas omprise several lands apes, suc as plains,plateaus, hills and mountains. Asso iated with these geo-morphologi al features there o ur the s rubs, herbs, grassesand a ta ean adapted to these physi al formations, onsti-tuting very parti ular habitats where an live and reprodu e

different spe ies of animals.Te most relevant fa t on erning the distribution of ani-mals and plants is that they are not randomly distributedalong their areas of o urren e; on the ontrary, there arespe i habitats where they an live. In this way, it is ourthought that: i) biologi al aspe t on erning the distributionof organisms among the various habitats that form e osys-tems annot be understood without the understanding of thephysi al stru ture of these habitats, ii) this omprehension

an be given by a geomorphologi al approac.Te rational of our thought is tied to the on epts estab-

lishing that the distribution of organisms ree ts their setsof adaptations to the immediate environment, a on eptknown as e ologi al nice (V , 1970; P , 1994).Tis idea is the soul of lassi al studies approacing biolo-gy (zoogeography) and geomorphology, whic were arriedout by V W (1970), V (1970, 1981)and A S (1967), urrently in remented by news geo-pro essing tecniques (C R , 2008; C

, 2009a; M , 1997).In this ontext, the aim of the present study is fo used on

the lands ape and habitats that o ur in open areas inside theAmazon region. Te s enario of this dis ussion en ompass-ing the eld of biogeomorphology omprises three ways: i)

on epts of morpho limati domains and biogeography, ii)

the ase study of a very interesting open area known as lavrado , situated in the Northern Amazon region the Brazil-ian state of Roraima, iii) use of geopro essing tecniques foridentifying and des ribing habitats.

2 The case study area

Te general region des ribed in this report (Figure 1), om-prised in the Guyana Shield (H , 2005), is a very pe-

uliar open area of some 69.000 km, mostly situated in thenorthern portion of the Amazon morpho limati domain,overlying three ountries. We estimate, by remote sensing,that this area overs some 45.000 km in the Brazilian state ofRoraima, 10.000 km in Venezuela and 14.000 km in Guyana.

In Venezuela this portion of open areas is about 12001600meters above sea level. It is cara terized by the presen e ofruiniform tabular mountains, individually alledtepuy . Tetepuyes are part of a geomorphologi al formation knownin Venezuela asGran Sabana . In the Brazilian territory thebest knowntepuy is the Roraima Mount (0511S, 6049W),around 2800 meters high, situated on the triple border ofBrazil, Venezuela and Guyana. We do not onsider this Ven-ezuelan region to be part of the Amazon morpho limati do-main (see A S , 2003).

In the Guyana region this Northern Amazon open area is

mostly situated on the basin of the Rupununi River, an afflu-ent of the main Guyanese river, the Essequibo. Tis open ar-ea, lo ally known asRupununi Savanna , is separated by the

a utu River. Tis river, that forms the border of Brazil andGuyana, runs in a geologi al ssure from South to North,where it turns westward to ow into the Urari oera Riverin Brazil (approximately at 0301N, 6028W), both riversforming the Bran o, whic ows southward into the NegroRiver in the Brazilian state of Amazonas.

In the Brazilian portion, the state of Roraima, this area isknown aslavrado , an old Portuguese term for open vegeta-tion (V C , 1991; C , 2009). Te

lavrado has its own so io- ultural and e ologi al identity,integrated by omplex networks of intera tions among thelo al people with the lands ape, and by a cara teristi lo alfauna and ora adapted to thelavrado e osystem (N

, 1998).Tis open area is formed by pe uliar geomorphologi al

features, suc as boulders, alluvial plains, lakes and galleryforests along the rivers. Isolated patces of forest, s rubs andherbs, are present throughout the area. Gallery forests o -

ur in the banks of the rivers. Tese features form the lavrado habitats, harboring many spe ies of plants and animals,whose biologi al aspe ts of their distribution along these re-gional habitats are also fo used in this study.

3 Material and Methods3.1 Geomorphology

o des ribe the morphology of the ase study relief we usedremote sensing tecniques (hypsometry, shaded relief, topo-graphi proles and RGB omposition) from elevations mod-el of SR M (Shu le Radar opography Mission) and Landsat5 images. Te elevation model from SR M is a radar image,a quired by interferometry method in 2001 for entire globe,used for geomorphometri s analysis of the terrain.

Te so ware ENVI 4.3 was used to resize the SR M data

to 30 meters, by interpolation, from original spatial resolu-tion of 90 meters. Tis digital elevation model was importantto identify the different altimetry values, and the morphol-ogy of denudation forms (ranges and hills) using shaded re-

Figure 1: Examples of open areas within the Amazon morphoclimaticdomain. 1 Roraima, Venezuela and Guyana. 2 Amap state, Brazil. A Monte Roraima; B Branco River; C Lacustrine Systems; D Serra daLua; E Serra Marari; F Marac Island; G Serra do Tepequm.

Abb. 1: Beispiele von Frei en innerhalb der morphoklimatis en RegionAmazonas. 1 Roraima, Venezuela und Guyana. 2 Bundesstaat Amap,Brasilien. A Monte Roraima; B Branco River; C Lacustrine Systems;D Serra da Lua; E Serra Marari; F Marac Island; G Serra doTepequm.


3/10


lief and topographi proles tecniques. Te opti al imagesof Landsat 5 are mostly used for environmental studies, with30 meters spatial resolution, and were used for identifyingagradational morphologies, like uvial plains, lakes and veg-etation aspe ts by visual interpretation.

Te Landsat 5 images RGB omposition was appliedin ENVI 4.3, using bands 5,4 and 3. Te Landsat 5 imageswere acieved in 2005, from De ember to April, whic or-responds to dry season (without louds), patc-rows were232(56,57,58); 231(57,58). Tese images were a quired at the National Institute of Spatial Researc (INPE) www.dgi.inpe.br/CDSR/ and Embrapa Relevo www.relevobr.cnpm.embrapa.br/.

3.2 Fauna examples

Case study of faunal elements, in the present ontext, were

determined through eld work ondu ted by the InstitutoNa ional de Pesquisas da Amaznia (National Institute ofAmazonian Researc INPA) in Roraima throughout thepast two de ades, mainly on the lavrado area (see C

Figure 2: Roraima, topographic transects proles a-a to c-c; Venezuela Roraima, transect D.

Abb. 2: Roraima, topographis e Transekte a-a to c-c; Venezuela Roraima, Transekt D.

, 2009). We take as examples the vertebrate fauna ofthe area, mainly aspe ts of its distribution along the habi-tats omprised by geomorphologi al features determinedthrough geopro essing tecniques (Figures 8, 9, 10).

4 Results and Discussion4.1 Geomorphological features of the lavrado : remotesensing

One an see the position and topographi prole of thelavrado and adja ent forested areas just looking at the regionthrough transe ts, for example overing the forests of theWest portion of Roraima up to the open areas in the East,or overing part of the VenezuelanGran Sabana , until thelavrado areas (Figure 2). At the same way, through transe ts(Figure 3) we an see the main features of the relief, like highaltitudes (more than 1500 meters high), with tabular relief

(tepuys ), agradational and denudational pro esses, moder-ated disse tion and low stru tural ontrol (Figure 3 1); in-termediary altitude, somewhat of 5001500 m, with denuda-tional pro esses, high disse tion and strong stru tural on-


4/10


pa erns. Tese lakes are fed by ground water and resemblethe lakes of the morpho limati domain of thecerrado (C

Z , 2009). Figures 67 show some aspe ts ofthe uvial plain and vegetation of thelavrado.

Te rivers that ross thelavrado are autocthonous, withits headwaters in the elevated serras that make the borderof Brazil and Venezuela, the Parima-Pa araima system. Telavrado drainage, formed by well developed uvial plains,is dire ted to the Negro River, whic runs from the Andesuntil its onuen e to the Solimes River, in the CentralAmazon Basin. Te main rivers that run in the lavrado have

banks (dique marginal ) and beyond these a formation alledvrzea , a oodplain area formed along the main rivers dur-ing the rainy season.

Te vegetation of the lavrado is omposed by interestingformations (B , 1959; A S , 1997). Troughoutthis open area one an see near 810 meters height and lessthan 0.5 ha, wood patces, surrounded by grouped or moredisperse s rubs and small trees. Te ground is overed bygrasses and grass-like plants (family Cypera eae). Lines ofpalm trees (Mauritia exuosa ), known as buritizais , dueto the popular nameburiti (family Palmae) for the palmtree, is an important element of thelavrado lands ape,starting in small lakes and running toward the main riv-ers, for a distan e of around 300800 meters. Te lavrado is surrounded by 1520 meters high forest, soil with shal-low li er, some emerging trees and somewhat unstru turedunderstory.

4.2 Geomorphology and fauna4.2.1 The approach

We an look at this intera tion between geomorphology andbiology from the point of view of different related areas ofknowledge. Whatever the area, the main idea of this intera -tion is fo used on spe ies and populations distribution, lo al

or along large areas. On the regional distribution, one maybe interested in des ribing the spe ies ricness between hab-itats within an e osystem, to understand aspe ts of the lo albiodiversity. On the other hand, we an fo us on the distri-

trol (Figure 3 2); low sedimentary plains, around 70100 m,with agradational pro esses like uvial plains and la ustrinesystems (Figure 3 3); and isolated hills, inselbergs, withstru tural ontrol (Figure 3 4).

In ontrast with the high elevation of the VenezuelanGran Sabana , the elevation of thelavrado area is relativelylow, around 70200 m a.s.l. Tis area is drained by the Bran-

o River, whic is omposed by a system of low hills, withlow disse ted relief, isolated residual peaks (inselbergs ), sur-rounded by lakes in the headwaters, the owing of whic

reates a inter onne ted streams (igaraps ) separated bysmall elevations, known astesos, forms drainage disse tionaround the lakes and streams (Figure 4). Also we an see inthe area the sugar-loaf formations ( po-de-aucar ) and later-ite layers exposed on the soil (lajeiro ).

In alllavrado areas narrow lines of palm trees remind oneof the lands apes of the morpho limati domain of the Cen-tral Brazilcerrados . Of ourse this resemblan e is only ap-parent, sin e thecerrado is a very distin t e osystem, situat-ed a few thousands kilometers from thelavrado . Te re on-naissan e of the distin tiveness between both e osystems lavrado and cerrado has a very important e ologi al andbiogeographi al signi an e (E , 1963; C , 1978;V C , 1991; C , 2009).

Te predominant de livity of thelavrado is between 5-8,with low energy, forming a region that re eives sedimentarymaterial, mainly sand oming from the surrounding rys-talline uplands (Guyana Shield). Telavrado entral por-tions relief low energy favors the formation of a omplex

la ustrine system, omposed by more or less ir ular up to300 meters long lakes, most of whic are temporary (Figure5). Tese lakes are independent, inter onne ted by narrowstreams, forming dendriti , re tangular and subdendriti

Figure 3: 1 Mount Roraima, moderate relief dissection, border of Venezu- ela and Brazil (0511N, 6049W); 2 Serra Marari, moderate to strongdissected relief (0416N, 6046W); 3 Uraricoera River, low relief dissection (0319N, 6025W); 4 Serra da Lua, low and strong relief dissection

(0227N, 6028W).Abb. 3: 1 Mount Roraima, moderate Reliefzergliederung an der Grenzezu Venezuela und Brasilien (0511N, 6049W); 2 Serra Marari, migebis starke Reliefzergliederung (0416N, 6046 W); 3 Uraricoera River,niedrige Reliefzergliederung (0319N, 6025W); 4 Serra da Lua, niedrigeund starke Reliefzergleiderung (0227N, 6028W).

Figure 4: 1 Tesos (low hills) convex morphologies; 2 lakes; 3 smallelevations between the streams.

Abb. 4: 1 Tesos (a es Hgelland) konvexe Morphologie; 2 Seen; 3 kleine Erhebungen zwis en den Strmen.


5/10


bution of spe ies and populations among e osystem, in orderto understand extensive distribution pa erns, for example,populations in onta t or separated by geomorphi barriers

that have o urred in the present or past events.wo emeritus herpetologists together with one geologistwere the rst to approac geomorphology to biology in 19691970. Te zoologists are the Brazilian s ientist Paulo EmlioVanzolini and his North Ameri an olleague Ernst Williams.Tey have formulated a very elegant South Ameri an lizard(genusAnolis , family Polycrotidae) spe iation model basedon forest expansion and retra tion, paleo limati events thato urred under the inuen e of the Pleisto ene dry and wetperiods over the past 20.00010.000 years.

Tis model of spe iation formulated by V W (1970) establishes that be ause of the forest frag-mentation o urring during paleo limati dry periods (gla-

iation) animals be ame isolated in forests patces, whicresulted in e ologi al barriers forest spe ies do not live inopen areas. Tese barriers, in turn, determined the interrup-tion of gene ow between populations. Dry events of thepast an be inferred at present by geomorphologi features,suc as the stone-lines (paleosols formed in dry paleo limat-i periods and buried in sedimentary deposits), indi atingthat a forested area today was open in the past (A S ,2003; H , 2007). Another way to infer past dry eventsis through palynologi al re ords and14C dating of sediments(A , 2000; S L , 1982).

During the humid phase (intergla ial) the forest oales ed

and what was fragmented forest be ame ontinuous forest-ed area; however, many animal spe ies did not cange geneagain, be ause their populations were isolated for a period inwhic several biologi al and physiologi al canges o urred

in eac one. Te result of these pro esses was the formationof distin t spe ies. Te model fo used mainly on the pulsa-tion of the forest in the Amazon region; however, the ideawas applied for other regions and spe ies (V , 1988,2002; W et al., 2005).

Following another way of the same theme, the Germangeologist Jrgen Haffer studying Amazonian birds, in 1969

ame to the same on lusion and model of spe iation as didVanzolini and Williams for lizards in early 1970. Tis modelof spe iation, taking geomorphologi al eviden es of expan-sion and retra tion of the forest, be ame lassi in biogeog-raphy and is well known as Pleisto ene Refugia Model andRefugia Teory (V , 1970; A et al., 1991; H

P , 2001; H , 1969; A S , 1982).Te morpho limati domains on epts, adopted by Van-

zolini and Williams as vegetation riteria for their study ofspe ies distribution and refuges, were rst formulated byAziz Na ib AbSaber in 1967. Prior to this Brazilian geogra-pher and geomorphologist, the vegetation of the regions inBrazil was identied through fragmented oristi features.

Te model of AbSaber gave the ne essary strength in iden-tifying large vegetal formations, instead of patces inside thesame e ologi al and geomorphologi formation. AbSaberused the limate, vegetation, soil, Hydrography and relief asfeatures to re ognize what he alled the area ore in a do-main, in a sub- ontinental s ale. All kinds of regional geo-morphologi al fa ies ould, then, be in luded in one domainor another cerrado, caatinga, mata atlntica and hilia (the Amazon) re ognizing the transitional zones.

Te model formulated by AbSaber was a great advan eto the elds of geography and geomorphology, sin e regions

ould then be identied as a ontinuous unit with related

geomorphologi features. o biologists interested in e olo-gy and biogeography this geomorphologi model integratedformerly s a ered data, enabling one to ome to a be erunderstanding of spe ies distribution.

4.3 Habitats and faunal distribution: the lavrado

All those geomorphologi al formations omprised in thelavrado , suc as hills, ro out rops, lakes, small patces offorest, s rubs and the gallery forests along the rivers, withthe ba -swamps (vrzea ) of the major ones, form the habi-tats inhabited by many organisms. Identifying these habitatsis the rst step to omprehend the biology of any spe ies thatlive in the lavrado , in terms of adaptations and gene owamong individuals and populations. Some geomorphologi alfeatures an illustrate this point of view, suc as the graniteand laterite extrusions, hogba s, inselbergs and sparse orgrouped boulders at various sizes (mataces ) in the plain andlow hills present in thelavrado (R , 1957). In additionto the geomorphologi al interpretation, these formationsalso have their e ologi al identity, forming omplex mi ro-habitats inhabited by birds, bats, rats, snakes, frogs, lizardsand many spe ies of invertebrates (V C ,1991; C , 2009; N B , 1997; Ret al., 1997).

Te ro s are distributed throughout the area and are di-re tly exposed to the sunlight. Tese features led to manyrelevant biologi al questions, suc as: How many spe iesof vertebrates and invertebrates are asso iated with these

Figure 5: Lacustrine system (0337N, 6015W); 1 Surumu River;2 Tacutu River.

Abb. 5: Lakustrines System (0337N, 6015W); 1 Surumu River;2 Tacutu River.


6/10


habitats? How many diet and reprodu tive adaptations havethese spe ies undergone so as to be able to survive in thesegeomorphologi al units? How an be geneti ally cara -terized the populations of the same spe ies inhabiting thelavrado ? Tere are any preferen es of some parti ular spe-

ies in residing ertain geomorphologi al features, suc asgranite and laterite? How these ro out rops are distributed(grouped or dispersed) and how to interpret the distributionpa ern?

An interesting ase of animal distribution in these granitehabitats ome from the frog Leptodactylus myersi , a spe iesthat seems to be endemi to thelavrado , living on the ro s,at least the main populations (H , 1995). Eac popula-tion of this frog seems to be separated by several kilometers,

whic is the distan e between the boulders. Qestions basedon this example may in lude: How are these frog popula-tions distributed, taking into a ount they are dire tly as-so iated with the boulders distribution? How to cara terize

the adaptations of this frog, in terms of reprodu tion and di-et? Where they lay their eggs, onsidering the extreme expo-sure to this habitat to sunlight and dry environments? Teseare questions being urrently studied.

Another spe ies very ommon in the ro formations ofthe lavrado is the lizardTropidurus hispidus (family ropi-duridae). Te biologi al questions that an be applied to thepopulations of this lizard are asso iated with the habitatswhere they live, suc as the boulders, small trees, border ofthe forest and in the small patces of forest. For example: Doall these lizards have the same set of adaptations? Is it pos-sible to determine the populations of this lizard pre isely byidentifying the habitats through geopro essing tecniques?

Among mammals there are some interesting distribution

in habitats omposed by la ustrine system in general, lowhills and disse ted relief, vegetation of the margins of riv-ers (mata ciliar ) and lines of palm trees (buritizais ), lateritelayers (lajeiros ) and boulders. All these geomorphologi al

Figure 6: A-B uvial plains, lowlands (0301N, 6029W and 0236N, 6054W); C-D uvial plains, highlands (0417N, 6032W and 0456N,6114W). 1 abandoned anel lakes of ood plain; 2 Lakes of at plain. A Mouth of the Tacutu River in the Branco River; B Mucaja River; C Cotingo River with structural control, non ood plain; D meandriform river developed at structural control, small ood plain with lacustrine systems(oxbows lakes).

Abb. 6: A-B Flussniederungen, Tieand (0360N, 2901W und 0236N, 6054W); C-D Flussniederungen, Ho land (0417N, 6032W und 0456N,6114W). 1 aufgegebene Kanalseen der Flussaue; 2 Seen der Tiefebene. A Zusammenuss von Tacutu in den Brancoss; B Mucaja; C Cotingo,ni t zur Flussaue entwi elt; D mandernder Fluss, klein ige Flussaue mit lakustrinen Systemen (Altwasserseen).


7/10


units omprise the environment where many mammal spe-ies an live, suc asMyrmecophaga tridactyla (tamandu )

and Tamandua tetradactyla (mambira ), two related spe iesof the family Myrme ophagidae (Order Pilosa) that feed ontermites and ants. Tese two mammal spe ies also have the

patces of forest as refugia during the night.Another spe ies that have its habitat asso iated with thegeomorphologi al units of the open areas is the li le mam-mal Nasua nasua (quati ) of the family Pro yonidae (OrderCarnivora), an inhabitant of the boulders of the plains andhills. During the day it is ommon to see this animal in thathabitat, looking for food, mainly earthworms, inse ts andsome fruits. Te vegetation of this habitat is omposed byherbs, grasses, s rubs and isolated trees, whereN. nasua anbe found limbed at night. Again, geomorphology gives thedire tion for des ribing these habitats.

Among birds we an also have some representative spe-ies urrently endemi to Roraima, suc as Aratinga sol-

stitialis ( jandaia-sol ) of the family Psi a idae, that live inhabitats omprised by the gallery forests or on the forestedge (approximately 0352N, 5937W). Te pre ise lo aliza-tion of these endangered spe ies habitats an be obtainedby geopro essing tecniques, like the other endangered spe-

iesSynallaxis kollari ( joo-de-barba-grisalha ) of the fam-ily Furnariidae. Tis small bird an live in habitats formedby low hills and disse ted relief, s rubs and small trees, upto the right bank of the a utu River, in Guyana territory.Some populations of this bird an also be found in Roraima,in gallery forest.

Te same rational an be applied to the botani al spe ies

present in thelavrado . For example, there is a small and in-teresting a tus genusMelocactus that o urs on the ro sforming lusters. Te distribution of this a ta ean an beeasily established through the identi ation of the ro ex-

trusions. Another a ta ean present in thelavrado , the dis-tribution of whic an be as ertained through geopro ess-ing tecniques, is the Brazilian popularmandacaru genusCereus , whose main distribution may be asso iated with thesoil, as well with lusters of termite nests genusCornitermes (approximately 0352N, 5937W).

It is also very useful and informative to apply the geopro-essing tecniques for understanding thelavrado vegetation.

Tese features of the lands ape in this area are made up bya omplex net of small more or less rounded forest patces(island forests) some 0.5 ha or less, palms trees (linear oralmost rounded), des ribed having the fo us on the habitatof animals. But we an also fo us the question with anotherlens. How the forest patces of thelavrado are distributed? Isthere any pa ern a ounting for forest patces distributionand soil? Te relevan e of these questions is not restri tedto the present, but imply in onsiderations suc as how thelands ape cange and what would be the impli ations forthe fauna and ora.

Tese questions lead us to look at thelavrado vegeta-

tion under another fo us, whic is the pulsation of openand losed vegetal formations under limate canges. It isquite possible that the expansion and retra tion of the for-est during the Pleisto ene have inuen ed the gene ow ofmany spe ies living today in these kinds of vegetation, on-ne ting or interrupting denitely or temporarily the patcesof forest. How the various spe ies of thelavrado terrestrialvertebrates, for example, were lo ally affe ted by the eventsduring the dry and wet paleo limate periods? What to sayabout the forest pulsation and limate cange that might beundergoing at present?

Re ognizing eviden e of pulses in thelavrado vegetation,

through geopro essing data asso iated with the lo al distri-bution of spe ies, might ertainly elu idate several of thesequestions. Tis is the ase, for instan e, of three sympatrispe ies of lizards of the genusGymnophthalmus (familyGymnophthalmidae) that o ur in the open areas of Rorai-ma and in the forest edge, in onta t with the lavrado . Tespe ies areG . leucomystax asso iated with termite nests,G .vanzoi in the onta t forest open areas, andG . underwodii inthe ontinuous forest (V C , 1991, C

1999). In a 1.5 kilometer transe t, we an nd thesethree lizard spe ies, eac one in its spe i habitat. Tesethree spe ies are so tightly taxonomi ally related, that it isdiffi ult to re ognize them at a rst look, and we an imaginehow many geomorphologi al events might have o urredfor the spe iation of these three lizards spe ies. We an mapthe distribution of these lizards through geopro essing tec-niques.

Looking again to the lands ape of thelavrado and its asso-iated fauna, another example of biogeomorphology applied

to the biologi al distribution of populations omes from thetermites. At least two spe ies of these so ial inse ts of thefamily ermitidae build their nests on the ground (epigeousnests):Nasutitermes minimus and the Cornitermes ovatus (B , 1988). Both spe ies of termites onstru t nests indifferent parts of thelavrado , maybe due to soil fa tors, veg-

etation over or both features together. Te nest ofN.minus is rounded on the top, around 3040 entimeters high, andthe base is 2030 entimeters in diameter, are onstru tedmainly over the hills (approximately 0320N, 6124W) . Te

Figure 7: A Venezuela-Roraima border, transition of the forest to lavrado grassland with tall shrubs and small trees (0402N, 6103W); B Vene- zuelan open areas, pat es of forest with well developed rills and structuralcontrol (450N, 6057W); C Serra da Memria, shrubs and small trees,vegetation slope with tors and blo s (0410N, 6057W); D island vegetation with small lakes at at plain (312N, 6057W).

Abb. 7: A Grenze Venezuela Roraima, bergang von Wald zu lavrado

Grasland mit hohen Bs en und niedrigen Bumen (42N, 613W);B Venezuelanis e Offenlands a mit gut entwi elten B en (450N,6057W); C Serra da Memria, Stru er und niedrige Bume, bewa - sene Hnge mit Bl en (410N, 6057W); D inselartige Vegetation mitkleinen Seen in einer a en Ebene (0312N, 6057W).


8/10


nest of C.ovatus is pointed on the top; the onstru tion is

very hard, around 2.0 meters high, and the base 1.01.5 me-ters in diameter, mainly onstru ted on the plains (approxi-mately 0352N, 5937W), at the same region of the a ta-

eanCereus .Tere are many animals asso iated to the nests of both of

these termite spe ies. Te ra lesnakeCrotalus ruruima , thelizards Tropidurus hispidus (Family ropiduridae),Cnemi- dophorus lemniscatus (Family eiidae) and the gekkoHemi- dactyulus mabouia (Family Gekkonidae) are tenants of thesenests. Also some spe ies of rats and opossuns, spiders andmany invertebrate spe ies live in those nests. Tere are in-teresting biologi al questions asso iated with the distribu-tion of those termite spe ies. With the help of geopro ess-ing tecniques so as to identify the areas of o urren e ofboth, nests and soil, any approac related to these termitesbe omes more pra ti al.

Te distribution of rare or endemi spe ies that o urin Roraima an be illustrated on maps using geopro essingtecniques, exemplifying spe ies distributed in thelavrado and surrounding areas of this open vegetation e osystem(Figures 8, 9, 10).

5 Conclusions

Examples exposed in the present dis ussion an guide the

fo us of the biogeomorphology approac in two dire tions:i) at a regional s ale or ii) at a sub- ontinental level, withinor among large vegetal formations. Either way, the questionsregarding spe ies and habitats distribution should be made

involving geomorphology as a ba drop of the whole s en-

ery.At a sub- ontinental level, onsidering large vegetal for-mations, the questions leads to problems related to spe ia-tion and its pro ess. Te re ognition of the geographi unitsof the spe ies been studied the morpho limati domains is fundamental for that approac, be ause the whole dis-tribution area of a single spe ies or groups of spe ies will be

ompared through biologi al aspe ts, whic may vary sig-ni antly or not. Te main questions that arise at this levelmay in lude: How many vegetal formation an be re og-nized inside the domain (or domains) been studied? How arethe soil, topography and hydrography cara teristi s in eacstudied region? Are these geomorphi features a ting as bar-riers for gene ow among populations?

At a regional s ale, suc as that of thelavrado area, be-fore the formulation of spe i biologi al questions it is alsoimperative to lo ate the geographi insertion of the regionwithin the main e osystem. On e re ognized the geographi

ontext of the study site, we turn the eyes to the diversityand omposition of the regional geomorphologi al units,suc as the boulders, plains, hills, montains, la ustrine sys-tem, drainage and regional vegetal formations, whic an bedone applying remote sensing and geopro essing tecniques.

Te geomorphologi al features will then cara terize thehabitats. aking these features as riteria for ategorize the

ompartments of the region, we an fo us on the questionsto be worked, whic an be dire ted to analyze spe ies ric-ness, regional distribution of a group of spe ies or distribu-tion of a single spe ies, habitat cange and modi ation of

Figure 8: A-A Uraricoera River (le photo); B Grande River; C Ser- ra do Tabaio. Region of the endemic lizards Gymnophthalmus vanzoi and

G. leucomystax (family Gymnophthalmidae). Forest and lavrado contact,with small pat es of forest (right photo). Transition of denudational andaggradational relief, with isolated hills, drained by a not well development uvial plain.

Abb. 8: A-A Uraricoera (Foto links); B Grande River; C Serra doTabaio. Region mit den endemis en Eide sen Gymnophthalmus vanzoiund G. leucomystax (Familie Gymnophthalmidae). Wald und lavrado-Kon- takt mit kleinen Waldinseln (Foto re ts). bergang von Abtragungs- zuAufs ungsrelief mit isolierten Hgeln, die Entwsserung erfolgt bereine s le t entwi elte Flussniederung.

Figure 9: Brazil-Venezuela border. A Tepequm tepuy (right photo);

B Parima River, hills; C Surumu River (le photo), type locality of theamphibian Ela istocleis surumu. Not well developed uvial plains (Su- rumu River), with temporary lakes. Litholic soils with boulders, tors andscrub-herbs vegetation.

Abb. 9: Grenze Brasilien-Venezuela. A Tepequm tepuy (Foto re ts);B Parima, Hgel; C Surumu (Foto links), Typuslokalitt der AmphibieEla istocleis surumu. S le t entwi elte Flussniederungen (Surumu)mit temporren Seen. Litholic-Bden mit Felsbl en, strau - und kraut- rei e Vegetation.


9/10


the lands ape, population e ology and onservation. We getthese data mainly through inventories, whic should startsomepla e.

If an exhaustive faunal survey is a hard task, be ause ofits high osts and need of experien ed personal involvement,a reliable alternative is to sele t habitat samples by mappingthe regional morpho limati units. Tis an be done apply-ing remote sensing and geopro essing tecniques. Where to

start will depend on the question. A good set of suggestionsan be found in H at al. (1994) and C (2009) forthe lavrado area.

6 References

A S , A.N. (1967): Domnios morfo limti os e provn ias togeogr-as do Brasil. Orientao, 3: 4548.

A S , A.N. (1977): Espaos o upados pela expanso dos limas se osna Amri a do Sul, por o asio dos perodos gla iais Qaternrios. Orientao, 3: 119.

A S , A.N. (1982): Te paleo limate and paleoe ology of BrazilianAmaznia. In: P G. . (ed.): Biologi al Diversi ation in the

ropi s: 4159; New York (Columbia University Press).A S , A.N. (1997): A formao Boa Vista: signi ado geomorfolgi oe geoe olgi o no ontexto do relevo de Roraima. In: B , R.I.,F E.J.G. C E.G. (eds.): Homem, ambiente e e ologiano estado de Roraima: 267293; Manaus (Editora do Inpa).

A S , A.N. (2003): Os domnios de natureza no Brasil: Poten ialidadespaisagsti as. 159.; So Paulo (Ateli Editorial).

A , M.L. (2000): Fisionomia das savanas de Roraima, Brasil. A ta Ama-zoni a, 03: 423440.

A , M.L., C , A.M., F , M., M , L., S , M., S, A., S , M.F.F., S , F., S , K., , B., H ,

V.D. . (1991): Mise en viden e de quatre phases d'ouverture de la fo-ret dense dans le sud-est de L'Amazonie au ours des 60,000 dernieresAnnees. Premiere Comparaison Ave D'Autres Regions ropi ales. Comptes Rendus de l'A admie des S ien es, 882: 673678.

B , A.G. (1988): Anlise da termitofauna (Inse ta: Isoptera) de umaoresta primria e de uma pastagem na Amaznia Oriental, Brazil. Boletim Museu Paraense Emlio Goeldi, 5: 225241.

B , Y. (1959): La rgion moyenne du haut rio Bran o (Brsil): tudegomorphologique. 254; Universit de Paris, Institut National de Re-cerces de LAmazonie.

C , C.M. (2009): O lavrado da Serra da Lua em Roraima e per-spe tivas de estudos da herpetofauna na regio. Revista Geogra aA admi a, 3: 417.

C , .M. (2009a): Parmetros geomorfomtri os para des rio dorelevo da Reserva de Desenvolvimento Sustentvel do up, Manaus,Amazonas. In: N , E. S , V. (eds.): Biotup: Meio Fsi-

o, Diversidade Biolgi a e So io ultural do Baixo Rio Negro, Amaz-nia Central: 317; Manaus (Amazonas).

C , .M. R , R. (2008): ni as de sensoriamento remotoapli adas biogeograa: metodologia geogr a para espa ializao de

molus os terrestres. Boletim Goiano de Geograa, 28: 157166.C , .M., Z , M. (2009): Morfometria e ara terizao do meiofsi o de ambientes la ustres no Vo do Paran-Gois, Brasil: uma pri-meira aproximao. erra Nueva Etapa, 38: 111138.

C , L.M. (1978): O on eito de errado. Revista Brasileira deBotni a, 1: 1723.

D , A B G.A. (1953): Phytogeographi al notes on the BrazilianAmazon. Anais da A ademia Brasileira de Cin ias, 19: 146.

E , W. A. (1960): Contribuio ao onhe imento dos ampos da Amaz-nia. I Os ampos do Ariramba. Boletim do Museu Paraense EmilioGoeldi, 4: 136.

E , G. (1963): Habitat ora of fazenda Campininha, So Paulo, Brazil. In: Ferri, M.G. (ed.): Simpsio sobre o errado: 179231; So Paulo(Editora da Universidade de So Paulo).

E , G. (1992): How names are used for vegetation. Journal of Vegeta-tion S ien e, 3: 419424.

E , G. (1994): Vegetao do errado. In: P , M.N. (ed.): Cerrado: a-ra terizao, o upao e perspe tivas: 1773; Distrito Federal E(ditorada Universidade de Braslia).

H , J. (1969): Spe iation in Amazonian forest birds. S ien e, 3889:131137.

H , J. P , G. . (2001): Climati for e of evolution in Amazoniaduring the Cenozoi : On the refuge theory of bioti differentiation. Amazoniana, 16: 579607.

H , D.S. (2005): An ient land in a modern world. In: H ,D.S. (ed.): ropi al forest of the Guiana Shield: 114; Washington (DC)(CABI Publishing).

H , W.R., D , M.A., M D , R.H., H , L.A.C. F ,M.S. (1994): Measuring and monitoring biologi al diversity. In: Stan-dards methods for amphibians: 384; Washington (DC) (Smithsonian In-stitution Press).

H , W.R. (1995): South Ameri an ro y habitatLeptodactylus (Amphib- ia: Anura: Leptodactylidae) with des ription of two new spe ies. Pro-eedings of the Biologi al So iety of Washington, 108: 695716.

H , S. . (2007). Reviso dos onhe imentos sobre o signi ado das li-nhas de seixos. Revista do Instituto Geolgi o, 2: 5364.

Figure 10: Distribution of endemic species in northern of Roraima, lavrado.A Lavrado area; B Forest area. Some species: Lizards: Mabuya carv- alhoi (Sauria: Scincidae), Gymnophthalmus leucomystax (Sauria: Gym-

nophthalmidae), Gymnophthalmus vanzoi (Sauria: Gymnophthalmidae);Amphibians: Dendropsophus benitezi; Ela istocleis surumu; Serpents:Micrurus pacaraimae; Birds: S istoci la saturata ; Herpsilo mus roraimae; Syndactyla roraimae; Myiophobus roraimae; amnophilus insignis;Megascops guatemalae; Mammals: Nasua nasua vi ata.

Abb. 10: Verteilung endemis er Arten im Norden von Roraima, lavrado. A Lavrado Berei ; B Waldberei . Einige Arten: E sen: Mabuya car- valhoi (Sauria: Scincidae), Gymnophthalmus leucomystax (Sauria: Gym- nophthalmidae), Gymnophthalmus vanzoi (Sauria: Gymnophthalmidae);Amphibien: Dendropsophus benitezi; Ela istocleis surumu; S langen:Micrurus pacaraimae; Vgel: S istoci la saturata ; Herpsilo mus roraimae; Syndactyla roraimae; Myiophobus roraimae; amnophilus insignis;Megascops guatemalae; Sugetiere: Nasua nasua vi ata.


10/10


M P , J. (1974): ipos de vegetao da Amaznia. Brasil Florestal,17: 4858.

M , J.P. (1997): Relationship between lands ape stru ture and treespe ies diversity in tropi al forests of South-East Brazil. Lands apeUrban Planning, 37: 2935.

N , S.P. (1998): O orrn ia de lagartos no lavrado de Roraima(Sauria: Teiidae: Gekkonidae: Iguanidae: Poly rotidae: Tropiduridae:Scincidae e Amphisbaenidae ), Brasil. Boletim do Museu Integrado deRoraima, 4: 3949.

N , A. B , U. (1997): Mamferos de Roraima: status da diver-sidade e onservao. In: B , R.I., F , E.J.G., C ,E.G. (eds): Homem, ambiente e e ologia no estado de Roraima: 565579;Manaus (Editora do Inpa).

P , L.C.R., O , P., M , M., G , R. A , R.,L , A.Z. (2009): Te evolution of a forest/grassland mosai sin e28,000 C-14 yr BP based on pollen and arbon isotopes. QaternaryResearc, 71: 437452.

P , E. R. (1994): Evolutionary E ology. 5th:486; New York (Harper Col-lins).

R , F. (1957): Expedies geomorfolgi as no erritrio do Rio Bran-o: 170; CNPq Rio de Janeiro (Inpa).

R (1975): Levantamento de Re ursos Naturais. Volume 8. Fol-ha NA 20 e parte da Folha NA 21 umu umaque NB 20 Roraima eNB 21: geologia, geomorfologia, pedologia, vegetao e uso poten ialda terra: 428; Rio de Janeiro (Departamento Na ional de Produo Min-eral).

R , J.A., R , M.S., H , A.L. (1997): Notas sobre insetos deRoraima. In: B , R.I., F , E.J.G., C , E.G. (eds):Homem, ambiente e e ologia no estado de Roraima: 489508; Manaus(Editora do Inpa).

S L , M.L. (1982): Climati cange at the Pleisto ene-Holo ene boundary. In: P , G. . (ed.): Biologi al diversi ationin the tropi s: 7477; New York (Columbia University Press).

V , P.E. (1970): Zoologia sistemti a, geograa e a origem das esp-ies. Srie eses e Monograas, 3: 156.

V , P.E. (1981): A quasi-histori al approac to the natural history ofthe differentiation of reptiles in tropi al geographi isolates. PapisAvulsos de Zoologia, 34(19): 189204.

V , P.E. (1988): Distributional pa erns of South Ameri an lizards.

In: H W.R., V , P.E. (eds): Pro eedings of a workshopon Neotropi al distribution pa erns: 317342; Rio de Janeiro A ademia(Brasileira de Cin ias).

V , P.E. (1992): Paleo limas e espe iao em animais da Amri a doSul tropi al. Estudos Avanados, 15: 4165.

V , P.E. W , E.E. (1970): South Ameri an anoles: the geo-graphi differentiation and evolution of theAnolis risolepis spe iesgroup (Sauria, Iguanidae ). Arquivos de Zoologia, 19: 1298.

V , P.E. C , C.M. (1991): wo sibling and sympatri spe-ies of Gymnophthalmus in Roraima, Brasil (Sauria:Teiidae ). Papis

Avulsos de Zoologia, 37: 173226.V , R.L., M , L.A., C , H.B., R , J.E. (2000): Te

Amazon basin and its natural y les. In: S , E., A , M.A., V, R.L. (eds.): Amaznia: Um e ossistema em transformao: 163

214; Manaus (Inpa).W , W., F , J., M , A., P , C., S ,

M., , R. (2005): ra ing an invasion: landbridges, refugia, and thephylogeography of the Neotropi al ra lesnake (Serpentes: Viperidae:Crotalus durissus). Mole ular E ology, 14:114.

carvalhoecarvalho-2012

Documents