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Home , Cloud forest, Macrothemis inacuta, Mecistogaster ornata, Yungas

Odonatologica38(1): 39-53 March I. 2009

Odonata of the ArgentineYungas :

distribution patterns and conservation status

N. Von Ellenrieder

Institute de Bio y Geociencias (IbiGeo), Museo de Ciencias Naturales de Salta,

Universidad Nacional de Salta, Mendoza 2, AR-4400 Salta,

[email protected]

Received September 12, 2007 / Reviewed and Accepted April 22, 2008

Odon. of streams, small rivers and ponds were sampled in the cloud for-

NW est of Argentina, and presence / absence information of spp. from samples and

from examination ofcollections was recorded in a spatial-relationaldata base. Alpha,

and total richness for the beta, and gamma diversity species expected area were es-

timated. Similarity in composition of odon. communities from lotic and lentic envi-

ronments were analyzed according to latitudinal and altitudinal gradients,using mul-

tivariate cluster analysis. Assemblages from NW Argentina were compared to those

from equivalent sites in SE . Odon. species diversity was found to follow both a

N latitudinal (decreasing from to S)as an altitudinal gradient (decreasing from low

to high elevations). Based on IUCN (2001) criteria, the conservation status of the

endemic odon. spp. to the Yungas was assessed at a global scale; 6 spp.

were assessed as of Least Concern and 2 as Near Threatened.

INTRODUCTION

The Yungas cloud forest extends from south into NW Argenti- na along the eastern slope of the Andean cordillera. Biogeographically it be- longs to the Yungas province included in the neotropical region (CABRERA &

and is in known WILLINK, 1973), encompassed a large hot-spot

as ‘Tropical ’(MYERS et al., 2000). In Argentina it represents one of the most species-rich biogeographic provinces, and is distributed discontinuously

along the Subandean chains withinthe provinces of Salta, Jujuy, Tucuman, and

Catamarca(Fig. 1).

The knowledge of the odonatesof the Argentine Yungas cloud forest has in-

the last few with creased considerably during years, the numberof recorded spe-

cies increasing from 44 in 1999 (MUZON & VON ELLENRIEDER, 1999) to

102 in 2007 (VON ELLENRIEDER & GARRISON, 2007a; 2007b). 40 N. von Ellenrieder

The goal of this study is to analyze the patterns of ©donate diversity along lati- tudinal and altitudinal gradients within the Yungas cloud forest and assess the

conservation of status its endemic species according to IUCN (2001) criteria.

STUDY AREA

The Yungas cloud forest extends in Argentinaalong anapproximately 50 km wide strip between

22° to 28°40’ S and 63° 68° to W (Fig. 1). Climate is warm and humid tosub-humid, and altitude as-

Fig. 1. Map of NW Argentinashowing localities studied. Extension of Yungas cloud forest in South

America is shown in black in inset Colours indicate of black: white: map. type environment, lotic; lentic, and shapes indicate zone, circles: foothill rain forest; triangles: mountain rain for- est; pentagons: mountain forest; diamonds: highland . of Yungas 41

cends from 300 2400 a.s.1. and in relation and to m Temperature humidity vary to altitude, latitude,

is about 900-1000 1300 in slope exposure. Average yearly precipitation mm, reaching mm someareas.

The strong altitudinal gradient generates considerable climatic variations resulting in different plant communities or vegetation zones: foothill rain forest (ca. 300-900 m), warm and humid; mountain rain forest (ca. 900-1500 m), temperate-warm and humid; mountain forest (ca 1500-2400 m), tem-

winter elevation perate (with frequent frost) and humid; and high , also called cloud grass- lands or humid puna (ca. 2400-3000 m or more), temperate-coldand sub-humid (BURKART et al.,

1994; not considered as part ofthe Yungas biome by some authors, i.e. BROWN, 1995).

METHODS

Lotic (streams and small rivers) and lentic (ponds) environments were sampledbetween September

2005 and April 2007, both in protected (National Parks Baritu, Calileguaand El Rey, private Reserve

El Pantanoso, and Yungas ofthe Biosphere) and non-protected areas of the Yungas cloud forest of NW Argentina(Fig. 1). Presence/ absence information of species from samples and

from examination of collections Fundacion (Institute y MiguelLillo, Tucuman, Argentina;Museo

de La Plata, La Plata, Argentina; Rosser W. Garrison collection, Sacramento,USA) was recorded in a

spatial-relationaldatabase. Stations were classified according to type of environment (lotic or lentic),

altitudinal zone (foothillrain forest, mountain rain forest, mountain forest, or highlandgrassland),

and latitudinal sector (Northern: 22° to 23°30’S; Central: 23°3F to 25°39’S; and Southern: 25°40’S

to 28°40’S;Fig. I).

Three indices calculated: richness diversity were alpha diversity (average specific per locality);

beta diversity (a measurement of the heterogeneity of the data, calculated as the ratio between total

number of species and average number of species); and gamma diversity (diversity at landscape lev-

el, calculated as total number of species across all localities). Total species richness expected for the

area was calculated using first-order jackknife and Chao 2 estimators. Similarity in compositionof

odonate communities from lotic and lentic environments was analyzed by latitudinal sector and alti-

tudinal zone, using multivariate cluster analysis with Sorensen (Bray-Curtis) distance coefficient and

flexible beta as linkage method at a value of p = -0.25 (McCUNE & GRACE, 2002). The resulting

dendrogramswere based on Wishart’s objective function converted to a percentage of remaining in-

measurement formation. Percentage complementarity(a of distinctness or dissimilarity;COLWELL

sectors & CODDINGTON, 1994) was calculated amongthe latitudinal and altitudinal zones of the

and foothill forest of and sites of lowland forest in Argentine Yungas, among sectors Argentina two

SE Peru (Manu, partially included in the Yungas, and Tambopata, belongingto the Amazon forest;

data from LOUTON et al. (1996) and PAULSON (1985, 2006, pers. comm.).

Based on IUCN (2001) criteria, the conservation status of the odonate species endemic to the Yun-

gas cloud forest was assessed on a global scale.

RESULTS

DIVERSITY PATTERNS

A total of 103odonate species in 45 genera and 10 families from 142 localities

was analyzed (Appendix 1). Alpha diversity was 5.1, beta 20.2, and gamma 103.

Maximum richness at a single locality was 22 species; 23 species were found at

only one locality, and 17 at only two. Most common species (recorded from20 or

more localities) were Mnesaretegrisea (Calopterygidae), Acanthagrion ablutum,

A. peruvianum and Argia joergenseni (), Rhionaeschnaplanaltica 42 N. von Ellenrieder

and R. vigintipunctata (Aeshnidae), Erythrodiplax sp. n. and Macrothemisimitans

(). Best represented family was Libellulidaewith 45 species, followed

by Coenagrionidae (22 species) and Aeshnidae (19 species). Megapodagrionidae,

and Corduliidae each Protoneuridae, were represented by a

single species. First-order jackknife estimate for total number of species to be

expected in the studied area was of 125.8 species, and Chao 2 estimate of 118.5

species.

Lentic and loticenvironments shared 42 (about 41%) of their odonate species,

found lotic with 61 species exclusively at (37%) orlentic(22%) water bodies. Lentic environments included representatives of five families; Lestidae, Pseudostigma-

Aeshnidae and with Lestidae and the tidae, Coenagrionidae, Libellulidae, spe-

cialized treehole-breeding ornata (Pseudostigmatidae) exclusive

to them. Lotic assemblages were represented by eight families: Calopterygidae,

Coenagrionidae, Megapodagrionidae, Protoneuridae, Corduliidae, Gomphidae,

Aeshnidae and Libellulidae, with Calopterygidae, Megapodagrionidae, Protone-

uridae and Corduliidaefound only in them (Appendix 1; Figs 2-3).

Odonate species richness was found to decrease following both a latitudinal

gradient (from northto south) in lotic environments (Fig. 2; lenticenvironments

were richest in the central sector), and an altitudinal gradient (from low to high

2. Stacked bars richness for odonate of Yun- Fig. showing species per family assemblages Argentine

environment latitudinal gas groupedby along a gradient. Odonata of Yungas 43

elevations), both in lotic and lentic environments (Fig. 3). Hierarchical cluster

of environments classified analysis according to a latitudinal gradient (Fig. 4) shows odonate communities to be segregated first by type of environment (lotic

or then lentic) and by latitude, with assemblages from northern and central sec-

tors of both lentic and lotic environments more similar among them than with

those of the southern sector. Considering the combined odonate fauna for the

three the central sectors, one was the richest in numberof species (Tab. I), and the

the three same relationship among sectors was found, with northernand central

sectors more similar among them (complementarity of 39 %) than central and southern sectors (complementarity of 52 %) or northern and southern sectors

(63 %, Tab. I).

localities Analyzing grouped according to an altitudinal gradient (Fig. 5) it is evident that altitude the main factor was organizing composition of odonate as-

semblages, withboth loticand lenticenvironmentsof foothill and mountain rain

forest more similar in species composition among them than with environments

3. Stacked bars richness for odonate of Fig. showing species per family assemblages Argentine Yun-

gas grouped by environment along an altitudinal gradient. 44 N. von Ellenrieder

Fig. 4. Dendrogramshowingrelationships between odonate assemblages of Argentine Yungasgrouped

— - N; - according to a latitudinal gradient. Lo: lotic; Le: lentic; northern sector; C: central sec-

— — S; southern sector - 1: foothill rain — 2: mountain rain 3: tor; (as per Fig. 1); forest; forest; mountain forest; — 4: highland grassland.

from mountain forest and highland grassland (see also complementarity values in Tab. II). Lowland assemblages (foothill and mountain rain forest) are clus- tered according to type of environment, with lotic environments from both rain forest zones separated from lentic environments. Highland assemblages (moun-

tain forest and highland grassland) are clustered by altitudinal zone, with lotic environments of mountain forest more similar in composition to lenticenviron-

ments of mountainforest than to loticenvironments of highland grasslands (no

lentic environments were sampled in highland grasslands). Combined odonate

altitudinal decreased in richness fromlow to altitude, assemblages per zone high with maximum richness (82) in foothill rain forest and minimum (11) in highland

grassland (Tab. II).

Odonate assemblages of NW Argentina foothillrain forest present a much low-

er species richness than comparable sites in SE Peru, with which they share less

than 10 % of their species (Tab. III). The latitudinalgradient is also evidenced

this scale the increase of northern and southern at by complementarity among pairs with increasing latitude (Tab. III).

Table I

Richness and percentage complementarity(in brackets number of species in com-

of odonate latitudinal mon) assemblages among Argentine Yungas sectors

1 Northern sector Central sector Southern sector

Speciesrichness 72 82 51

Central sector 39.58 (58)

Southern sector 63.33 (33) 52.22 (43) Odonata of Yungas 45

Fig. 5. Dendrograms showing relationships between odonate assemblages of Argentine Yungas ac- cording to analtitudinal gradient. Lo: lotic, Le: lentic; - I: foothill rain forest; - 2: mountain rain forest; - 3: mountain forest; - 4; highland grassland.

CONSERVATION STATUS OF ARGENTINE YUNGAS ODONATES

half of the recorded found Slightly over species (55, representing 54%) were within one or more of the protected areas surveyed (Appendix 1). Howeveralmost all of the species of restricted distribution in the Yungas are distributed across otherbiomes of the neotropical region. Many of the restricted species are widely distributed in the Amazon, Chaco and / or Paranense , and reach in the

Yungas cloud forest of NW Argentina theirsouthernmost limitof distribution; i.e. Rhionaeschnapsilus, inacuta, M. musiva, Micrathyria atra, and

Tramea binotata (all from to N Argentina), Orthemis aequilibris (Costa

Rica to NW Argentina), Micrathyria venezuelae(Venezuela to NW Argentina),

Hetaerina sanguinea, and Neoneura bilinearis (both from to and NW Argentina), Acanthagrion aepiolum (Peru to Brazil and N Argentina),

Gynacantha convergens ( and Paraguay to N Argentina), or westernmost limit of i.e. distribution; Lestes spatula, Homeoura ambigua, Coryphaeschna per- rensi, Dasythemis mincki, Perithemis icteroptera and Tauriphila risi (all fromBra-

Table ii

Richness and brackets in percentage complementarity (in number of species common) of odonate

assemblages among Argentine Yungas atitudinal zones

II Foothill rain Mountain rain Mountain Highland

forest forest forest grassland

Species richness 82 76 19 11

Mountain rain forest 42(100)

Mountain forest 81,17(16) 76.62(18)

Highland grassland 93.1 (6) 91.25 (7) 75 (6) 46 N. von Ellenrieder

zil, Uruguay and E Argentina to NW Argentina). And the following four species, characteristic of of or grasslands Monte, Pampean, or Patagonian biomes, reach their northernmost distribution limit here: Oxyagrion rubidum andRhionaeschna confusa (both from S Argentina and Chileto N Argentina and

Paraguay), and, only found in high elevation grasslands of the Yungas, Rhion- aeschna haarupi and Progomphus joergenseni (both from W central Argentina to

NW Argentina).

Eleven identified species were as cloud forest endemics (Figs 6-7); from them

undescribed and three are still practically nothing is known about themprecluding

evaluation of their conservation a meaningful status (Limnetron sp., Micrathyria

From the six sp. 1, Micrathyria sp. 2; Appendix 1). remaining eight species were assessed as of Least Concern (LC) and two as Near Threatened(NT) (Tabs IV-

V).

DISCUSSION

The odonate fauna of the Argentine Yungas cloud forests is reduced and mar- ginal as compared to the same biome farthernorth. i.e. 103 species in 10 families compared to 136 species in 13 families in Mann National Park, Peru (LOUTON et al., 1996), but it is still high compared to other areas of Argentina, housing over a thirdof the total number of species recorded from the country (VON ELLEN-

RIEDER & MUZON,2008). The total number of species foundin this biome is

than that for wetlands higher registered NE Argentina (75) from a comparable latitude(MUZON et ah, 2008), but the maximum number of species recorded for a particular locality is considerably lower (22 against 45; most likely due to combinationof differenthabitats for each locality in MUZON et ah, 2008). The

Table III

Richness and percentage complementarity (in brackets number of species in common) of odonate as- semblages among foothill sites of Yungas in Argentina and lowland cloud forest in SE Peru (data for Manu

from LOUTON and for from PAULSON, 1985; 2006; et al., 1996, Tambopata piers, comm.)

Ill Lowland cloud forest in Peru Foothill rain forest in Argentina

Manu Northern Tambopata sector Central sector Southern sector

Altitude(m) 250-550 100-300 340-840 300-880 325-865

0 Latitude (S) 1I 55'-I2°55' 12°55'-13°2r 22°-23°30' 23°3r-25°39 25°40'-28°40'

Species richness 135 177 67 56 39

Tambopata 72.13(68)

Distance (km) 250

Northern sector 91.97(15) 93.91 (14)

Distance (km) 1,350 1.100

Central sector 93.29(12) 94.09(13) 36 (48)

Distance (km) 1.600 1,350 250

Southern sector 93.9(10) 95.65 (9) 60.52 (30) 56.06 (29)

Distance (km) 1.850 1.600 500 250 Odonata of Yungas 47

higher overall diversity seen here is explained by the high environmental heteroge-

of this forest which offers suitableenvironmentsfor neity a widearray of special- ized but localized thus in of species, resulting a high replacement species among localities is indicated the B value. These results with as by high diversity agree observations by PAULSON(2006), who notedthat forest environments support a more diverse odonate fauna than those of open lands.

The decrease of species richness with increasing latitudefound in loticodonate

distribution Fig. 6. Known area of Yungas cloud forest endemic species: Andinagriongarrisoni (Zy-

goptera, Coenagrionidae); Teinopodagrionmeridionale (Zygoptera, Coenagrionidae); Phyllocycla basidenta (Anisoptera,Gomphidae); and Progomphuskimminsi (Anisoptera,Gomphidae). 48 N. von Ellenrieder

communitiesof the had been observed for other Argentine Yungas already groups of organisms, and has been referred to as a ‘latitudinalclimaticimpoverishment’ or‘peninsular effect’(DE LA SOTA, 1972; OJEDA & MARES, 1989; BROWN et al., 2001). Within odonates, assemblages of northern and central sectors are more similar between themthanwith southern sector assemblages, differing from the situation reported for trees, where central and southern sectors are more sim-

Fig. 7. Known distribution area of Yungascloud forest endemic species; Argiayungensis (Zygoptera,

Coenagrionidae); Andaeschna rufipes (Anisoptera, Aeshnidae);Gomphomacromiafallax(Anisoptera,

Corduliidae);and Macrothemis hahneli (Anisoptera, Libellulidae). Odonata of Yungas 49

or the lo- to re- as one the re- over- due does to 20,000 10 which reduc- sections to due than well ground and Searching chasing than thanthan and as patrol Bolivia collectors existence habitat on returning greater existence less vulnerable S ELLENRIEDER b Argentina often by kimminsikimminsi lower colors. needed, its by 1973 Gomphidae) Males land unknownunknown sampled, (NT) from as likely across NW 9,450 sunny, always from 2007a, Belle, 330-1,200 cryptic ensure VON toto rivers. listed most non-specialist cryptic localities threatened LarvaLarva occurrence localities to future occurrence known scarcely to future Progomphus when frequently shore known scarcely 1973; and shore not especially and (Anisoptera, by (Anisoptera, Bolivia spot. Threatened of the at spot. and been been further become in GARRISON.GARRISON, Streams stream another, distributiondistribution pale monitoring BELLE,BELLE. of twigs samesame Near Extent km2 cations; corded,corded. has looked tiontion has to for notnot &

and sur- for patrol Larva km2; known like- scarce- swift water most especially males rocks. 20,000 not been Searching ELLENRIED- b with over on but ArgentinaArgentina on of has basidentahasidenta 1987 Gomphidae) 2007a, 6) Gomphidae) where close VON NW rapids settling locations; areas, distributed,distributed, which neededneeded VON NW 330-1,360 20,000 settling (LC) ©donates. (Fig. rivers, very 3 Dunkle, toto with occurrence 1987; PhyllocyclaPhyllocycla with for and beat from protected widely Bolivia localities GARRISON,GARRISON. (Anisoptera, Bolivia Concern of Yungas areas frequently S sampled & Streams straight unknown more sampled DUNKLE,DUNKLE, river straight face,face, Least Extent known within across further ER ly ly ER the a to a to onon fe- still of de- of ar- its of stream abun- at data fromfrom Listed and stream close vegeta- land and aquatic of Listed GAR- 2006 highland position locations km2 at while zones extent km2 MUZ6N, vegetation, horizontal- ensure threatened & to dating future 10 dating protected future species flying water, locally observed toto waters flying aquatic copula water, among collection evidence & plants known 20,000 the IV garrisoni Muzon, be forest horizontal in occurrence. aquatic horizontal than no near in perching near on rangerange no fromfrom become in & Coenagrionidae) -3,150 still patrol live vegetation on to Argentina water (NT) can adults is than 15,66015,660 rain in Pairs of 800-3,150 and emergent due monitoringmonitoring b emergent it and in more 20 not Table endemic 800 in entire ELLENRIEDER NW Males bushes Larvae three Based there abundant Males of lower records reduction Andinagrion pools remain over extent does Ellenrieder tandem. from where for extent does ELLENRIEDER 2007a, surface and 2007 category ofof four (Zygoptera, mountain oviposit Threatened site). in category (Zygoplera, with in VON in across (i.e. leaves and in requiring VON across (i.e. to requiring von tandem. to habitat of onon thisthis Seepages, edges from grasslands. water'swater’s tion, when grasses males vegetation Near Known Yungas,Yungas, single available 1967 crease occurrence absence existence VON 2006; RISON. ly lion, Near (13) dant ly in (13) in eas, by 2006;

status in veg- grass indi- and km2; pro- N.P.N.R and closed,closed. shore- still macro- b on macro- province, on Rey N.P.N.R ELLEN- nearnear stones within or stones 47,200 2007a, perch emerged wings while 47,200 El wings 4 2001 4 Jujuy VON meridionalemeridionale Argentina riparian of water under and in Conservation Adults with vegetation oviposit under Calilegua freshly oviposit of Megapodagrionidae) NW freshly vegetation vegetation locations, 2001; 47,200 live (LC) 485-1,900485-1.900 on GARRISON, Marmels, toto perch on (Baritu rivers. 16 shore; females leaves occurrence 16 province; Pantanoso overhanging land province; & De and may in Larvae riparian Bolivia may of from El TeinopodagrionTeinopodagrion Bolivia on Concern areas MARMELS, and (Zygoptera, S Salta Argentina) Streams elationetation blades viduals Tandems line, tandem phytes. among Least Extent known tected ReservaReserva Argentina) RIEDER line, in DE

(km2)

ecology a.s.1) occurrence assessmentassessment (m and of Distribution List AltitudeAltitude Extent Habitat Red Rationale References 50 N. von Ellenrieder

sur- on oc- rocks high Larvae areas km2; within N.P. Pan- VON rapidsrapids or in Rey El to water’swater’s perch perching three El Argentina) 2007b; Argentina over twigs water. 222,250 GARRISON, margins, vegetationvegetation and Reserva hahneli copula of and near & Libellulidae)Libellulidae) Colombia NW streams creeks, and & 19131913 in locations, and province, and to frequently stream (LC) (Baritu and 222,250 frequently grass trees 350-1,500 Pairs aquaticaquatic 17 Ris, 17 close and occurrence Macrothemis Peru enclosed on and areas province, Jujuy ELLENRIEDER.ELLENRIEDER, from flyfly of in shores. Concern b (Anisoptera,(Anisoptera, Venezuela hovering river abundantabundant bushes at Salta ELLENRIEDERELLENRIEDER ,Ecuador. Adults face,face, narrow casionally along Least Extent known protectedprotected tanoso VON 2007a, in live of Least in VON

a oth- or fly- byby N.P. on km2; loca- N.P. and along Larva species km2; loca- seepages. fromfrom semi-ter- reported Peru) GARRI- margins occasional- vegetation; with slope (Manu fallax reported ovipositingovipositing moss. this 525,000 known (Manu Bolivia 525,000 & and to 18811881 to were 1 low vegetation likelylikely 1 Corduliidae)Corduliidae) to flight, of area covered of streamstream flight, or covered covered 7) Argentina territoriesterritories likelylikely towardtoward covered department, Argentina tricklestrickles on most b Peru 250-3,100 525,000 low Females most Peru 250-3,100 stones sides belonging (LC) locations; small stones land moss in protected (Fig. McLachlan,McLachlan, and NW with land water, butbut distribution) protected Dios cliff occurrence 11 2007a, on Gomphomacromia narrow patrolling trail a narrow trail a abdomenabdomen larvae de ELLENRIEDER (Anisoptera,(Anisoptera, pairs margins. larvae moist, defenddefend paths on of from dirt Concern 2005, drippingdripping a within Yungas Ecuador males, perching beneathbeneath within Madre Streams,Streams, Males forests mating stream flickingflicking undescribed,undescribed, restrial; (based from forest Least Extent known VON SON, er ly ing andand tion in VON the of El of within patrols down- moss water N.P. EL- Peru within patrols down- Larvae Argen- 2007a, in locations, the locations, Avila Chorro Venezuela; VON to and Male in species Male alone in water. sediment occurrence 7 rivuletsrivulets up (El and province, V rufipes Aeshnidae) twigs state, province, twigs on from 1918) Argentina forest.forest. on areas GARRISON, Colombia andand above of areas Tama 1981,1994; flying tree and Tama Jujuy 875,000 oviposits m 800-1,300 rainrain m (LC) Table (Ris, NW (LC) known Tachira & endemic and in and and -2-2 extent 1 extent state, Andaeschna 1 Andaeschna stones in and streams protected N.P. (Anisoptera, territory female logs km2; four mountainmountain about Concern N.P.N.P. MARMELS, Venezuela under Miranda of large up Estimated within DEMARMELS, LENRIEDER NarrowNarrow thickthick stream;stream; covered live Least Estimated 875,000 Indio Calilegua tina) a live DE a to 4 in b

status en- & Adults dap- their km2; Tingo Am- 2007 Adults landing. (Manu creekscreeks of landing, locations Bolivia) Argentina closing 1,850 and Peru; or forest.forest. after 3 areas ELLENRIED- is 1,091,850 dep., patches 1.09 dep., dep., dep., gens Ellenrieder, NW and Coenagrionidae) streams rainrain times of ELLENRIEDER Coenagrionidae) to streams Cruz Conservation yungensis to Cruz yurt small locations; Dios VON 119-1,800 1,091,850 (LC) protected von thick in opening de Huanuco 2007a thick three 33 & 33 Santa ArgiaArgia & Bolivia narrow occurrence VON occurrence in or leaves within in and withinwithin unknown from within Madre (Zygoptera,(Zygoptera, andand two Concern of N.P. on sunlight, N.P. 2007; Garrison in Peru GARRISON GARRISON, ShadyShady closedclosed perch pled wings Larva Least Extent known known N.P.N.P. Maria boro ER,

(km2) ecologyecology a.s.1) occurrence assessment (m andand of Distribution List Altitude Extent HabitatHabitat Red Rationale References Odonata of Yungas 51

ilar in species composition and northern sector less similar (MORALES et al.,

latitudinal is observed broader 1995). The same pattern at a geographical scale, with assemblages of SE Peru cloud forest much richer and diverse (LOUTON et al., 1996; PAULSON, 1985, 2006) than those of NW Argentina sectors of the

Peruvian sites is same altitudinal zone. Although the distance between the two approximately the same as between adjacent Yungas sectors in Argentina (250

of km), they are more different in species composition than are any pair Argen- tine sectors (Tab. Ill), indicating their different biogeographic allegiances. Manu, which is partially included within the Yungas cloud forest, is accordingly more similar to NW Argentine sites than is Tambopata, which lies completely within the Amazon forest.

ob- A reduction of richness along an altitudinal gradient similar to the one served here has been reported for mammalsof the Argentine Yungas (OJEDA &

the factor MARES, 1989). Altitude has also been recognized as most important organizing the structure of benthic macroinvertebrate communitiesin the Yungas cloud forest although communitiesin general did not correlate well to the vege-

tationzones and were segregated into just two categories, one for lower and one

for higher zones (MALDONADO & GOITIA. 2003; VON ELLENR1EDER,

2007a).

with elements from vari- The Yungas are known to house a high biodiversity

from and Chaco ous biogeographic origins (tropical, Amazon, Paranense, prov-

inces, and austral-gondwanic) but low (CABRERA & WILLINK,

1973; BROWN et al., 2001). A relatively high endemicity value has been reported

of the cloud forests 10 % of only for anurans NW Argentine corresponding to

the Argentine species (LAVILLA et al., 2000). Within Odonata, the 11 species

that have been identifiedhere as cloud forest endemics represent only 4 % of the

Argentine fauna. Six of them are extensively distributed withinthe cloud forest

(Tabs IV-V), and only two, Andinagrion garrisoni and Progomphus kimminsi(Fig.

restricted and closer 6), are known from a more range require monitoring.

According to species richness estimators calculated, known species represent

still only 82-85 % of the odonate fauna expected for this area; the early stage of

of fauna the of our knowledge this is also evidenced by ongoing discovery new

species, and the fact that the biology, including life cycles and habitats, of about

a third of the recorded species is still largely unknown (VON ELLENRIEDER

& GARRISON, 2007b). The richest zone in odonate species is also the most se-

foothill rain forest has been verely threatened; ninety percent of what once was

replaced by monocultures(REBORATTI, 1989). Modificationand loss of aquat-

ic habitatsby irrationaland intensive human exploitation (both and

selective ; BROWN et al., 2001) are the main threats to the conservation

of the high odonate diversity of this cloud forest. 52 N. von Ellenrieder

ACKNOWLEDGEMENTS

I thank the authorities ofthe Administration of National Parks of Argentina, the Secretary of the

Environment and Sustainable Development ofSalta Province, the Provincial Direction of the Envi- ronment and Natural Resources of Jujuy, and the Private Reserve of El Pantanoso for permission to conduct this study in areas under their jurisdiction, GUILLERMO VON ELLENRIEDER and

ROSSER W. GARRISON for their support and camaraderie in the field, and Rosser W. Garrison,

RODOLFO NOVELO-GUTIERREZ, and DENNIS R. PAULSON for their critical review ofthe manuscript and helpful suggestions. This study was supported by the CONICET (Consejo Naeional

Cientificas de Investigaciones y Tecnicas, Argentina) and the Society (Grant

# 8016-06).

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