CCM~U - CGIARciat-library.ciat.cgiar.org/...G8_Biogeography_and_taxonomy...Manihot.pdf · Bogor 16111, Jawa Barat Indonesla (62) (251) 313-Gn7 (62) (251) 317-9';1 ABSTRACT We rnapped

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For pub11cat1on 1n B1olog1cal Control

March 15, 1993

Untll 30 June Address correspondence to

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1 July - 31 August

As of 1 September

Ann R Braun Cassava Program Centro Internatlonal de Agrlcultura Trop1cal Apartado Aereo 6713 Call, Colombla

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COlEWON HISTORICA I I

Bogor 16111, Jawa Barat Indones1a

(62) (251) 313-687 (6;» (251) JI7-'}',]

and Taxonomy of }lononYf:h§!llus specles assoc1ated Man1hot esculenta Crantz ln the Amer1cas

" r ' ~ i , .J '\. "'1 ~ \, J " , ....... " )..

J~ MV Guerrero, en HU W Flechtmann, M' e Duque, A Galgl, A' e Bellott1, G, J de Moraes and A R Braun

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CCM~U LIB~ARY

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B10geography and Taxonomy of ~ononychellus spec1es assoc1ated w1th Man1hQt esculenta Crantz ln the Amerlcas

J M Guerrero1 , C H W Flechtmann2 , M C Duque1 , A Ga1g1 1 ,

A C Bellott11 , G J de Moraes 3 and A R Braun1 ,4

,

1 Cassava Program, Centro Internaclonal de Agrlcultura Troplcal,

Apartado Aereo 6713, CaIl, Colombla

2 Un1vers1ty of S. PauIo. ESALQ, 13400 P1rac1caba, SP, Brazl1

3 CNDPAjEMBRAPA, 13820 Jaguarluna, SP, Brazl1

4 Correspondlng Author

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RUNNING HEAD B1ogeography and Taxonorny of Mononychellus Spp

Unt11 30 June Address correspondence to

Telephone Telefax: Electronic Mal1 (Bitnet).

1 July - 31 August·

As of 1 September

Telephone' Telefax

,

Ann R Braun Cassava Program Centro Internat10nal de Agr1cultura Trop1cal Apartado Aereo 6713 call, Colomb1a

, 57-23-675050 ext 373 57-23-647243 ABRAUN@CIATCOL

9 Wheathill Close Leamington Spa Warwlckshire CV32 6PL Unlted Kingdom

International potato Center ESEAP Reglon c/o Central Research Inst1tute for Food Crops (CRIFC) Jalan Merdeka 147 Bogor 16111, Jawa Barat Indonesla

(62) (251) 313-Gn7 (62) (251) 317-9';1

ABSTRACT

We rnapped the distrlbutlon of Mononychellus spp. assoclated wlth

cassava basad on survey of 1264 fields in thirteen central and

South American countries. We collected M tanaJoa (Bondar) ln

Panama, Colombia, Venezuela, Guyana, Trinidad & Tobago, Brazll

and Paraguay, but not north of Panama, nor south of Colombla ln

the Andean reg10n. M. tanalQa was prlmarlly associated 'l1th

hUlnld to seasonally dry lowlands except in northeast Brazll,

where the ecólog1cal range extends to semlarld lowland areas M

carlbbeanae (McGregor) was the most gBographlcally w1despread

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f ,

spec~es, and was the predom~nant spec~es of Mononychellus. on

cassava in semiarid lowland areas, except ~n northeast Brazil,

Peru or Paraguay where ~t does not occur. We found M mcgregor~

Flechtmann & Baker ~n hum~d highlands (interandean valleys) of

Colomb~a, Ecuador and Peru, 1n subtrop~cal southern Braz11, and

~n the Colombian region of the Amazon Basin (humid lowlands). M

plank1 (McGregor) was collected from ona field ~n northeast

Brazil and from fiva fields in Colombia. MQnonychellus tanaJoa

(Bondar) ~s a polymorph1c specaea w1th cons~derable var1ab1l1ty

in the length of the dorsocentral setae 01, 02 and D3 We found

polymorpbic populations of M tanaJoa in throughout its known

ranga in the Americas, except in northeast Brazil where setal

morphology is skewed towards the short extreme of tbe phenotyp1c

range The largest number of MODoDychellus apecies on cassava

and a h~gh degree of setal polymorph1sm ~n M tanal0a ocurred 1n •

Colomb1a. Several 1mpll,cat~ons for bl,olog~cal control of M

tanal0a follow from the ex~stence of a geographical subpopulat~on

d1st~nct from otber populations in the Amer.1can of African range

of thia apecies •

KEYWORDS. Mononychellua tanal0a, MonQnychellus canbbeanae, Man¡hot esculenta, Cassava Green Mite, b~ological control,

, electrophores.1s, taxonomy, bl,ogeography

2

INTRODUCTION

Taxonomy of CGM

The taxonomlC controversy surrounding the so-called cassava

" green mlte (CGM) complex led to uncertal.nty about the number of

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specles of MODoDychellus lntroduced to Afrlca, and has been

revl.ewed Yaninek and Herren (1988). Analyses of Afrlcan

specl.mens by Gutlerrez (1987) based on the form of the aedeagus,

by Roqo itt. al. (1988) based on 22 morphological characters ln

addition to the aedeagus, and by Murega (1989) based on

hybridizations between l.ndi viduals from different geographlcal

populations, concurred that only one epecles was introduced

Gutierrez (1987) reported that differences between and within

populations were common with respect to the lengths of the dorsal

setae of the genus Mononvchellus. Thls type of variation had

been previously reported for other tetranychid genera such as

Eutetranychus (Gutl.errez 1985) Rogo, Gutierrez and Murega dld

not agree, however, on whether CGn ln Afrlca should be called M

tanaJoa or M. proqresJ.vus Doreste (1981). M. tanaJoa was the

name given ln the original description of the epeCles (Bondar

1938), however type specimens collected from casaava In northeast

Brazil can not be located (Gutierrez 1987).

M· progresJ.YUs la a species described from cassava ln

Venezuela. This name ente red the literatura when variatlon in

the lengths of the dorsocentral setae of green mites collected

froro cassava ln the Americas led Doreste (1981) to describe two

new species, M. prQqresJ.yus, and M IDam,hQtJ., aupposed slbling

specles of M taDaJOa Rogo ~ al (1987) studied the lengths of

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the dorsocentral setae 01, 02, and 03, of speclmens from

Venezuela, representlng the topotype of M. progresl.vus, trom

Brazil, representing the topotype tor M. tanaJoa, and from

several Atrlcan countrles, concludl.ng that these characters alone

oould not :be used to dl.stl.ngulsh :between the two specles Sl.nce

thel.r lengths varied in a continuous gradient from the shorter

tanaJoa to the longer p¡:ogresl.vus type. Subgequently, Rogo JiL\;.

al. (1988) found a relationship :between the lengths of the

dorsocentral setae and the geographical origln of Afrlcan

specimens of M. tanaloa, and concluded that populatl.ons could be

classitled into short, lntermedlate and long setal forms.

The Orl.qln of the Cassaya Green Mlte

Classical biolO9ical control programs

• generally begin by

exploring for natural enemies in the likely areas ot orlgln of

the pest (Waage 1990) M tanaloa was accidentally lntroduced to

Afrlca ln the 19709 (Yaninek & Herren 1988) from the Neotroplcs

(Nyllra 1972, unpublished¡ Lyon 1974). Areas ot the Neotropics

WhlCh are ecologically similar to CGM-affected areas of the

Atrican cassava-growing belt are priority areas for exploration

tor natural enemies of CGM for lntroduction to Africa (Yanlnek &

Bellotti 1987, Bellotti ~ Al. 1987) A second strategy,

compatlble with the agroecological homologue approach, lS

suggested by the trophic relationship between CGM and cassava

CGM feeds almost exclusively on Manlhot, the cassava genus, ln

the Neotropics (Byrne 1980, Moraes & Flechtmann 1981, Moraes ~

al submittedl, and has maintained this trophic habit under

Afrloan conditions (Yaninek , Herren 1988) The oligophagous

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relat10nsh1p between CGM and MaD1hot 1n the Neotropo1CS suggests

a close coevolutionary relationship between HaD1hot and CGM

(Yan1nek & Bellotti 1987) More speciflc knowledge of the er1g1n

of CGM wlth1n the Neotrop1cs would contrlbute select10n of areas

for natural enemy exploratlon.

We measured dorsocentral setal lenqths and evaluated other

taxonomic characters for a large sample of H. tanaloa spec1mens

from the Americas to determine the degree ef variab1lity of these

characters, and to test Rogo's hypothesis of geographic var1atlon

1n morphology Based on these taxonomic analyses and other

supporting data;~ we present a hypothesis for the area of orlq1n

of the trophia relatlonsh1p between MoDonychellus speales ami

cassava, and discuss 1mpllcat1ons for bioloqical control of H

tanalOa.

METHOOS ANO MATERIALS

~eclmeD CollectloD

Spec1mens were obtalned durinq exploration trips made tor

collection and characterizatlon of CGM natural enemies. The

countries visited and the areas searched within countries were

selected according to a system of priority based on

agroecological homology between the Americas and CGM-affected

areas of Africa (Yaninek & Bellotti 1987, Bellotti ~ al 1987)

Homology maps were prepared based on Carter's (1986) cassava

microregion classificatlon for S. America, and h1ghest priorl ty

was g1ven to tropical, seasonally dry (4-6 months/year wlth < 60

mm preclpitatlon), isothermic lowlands Lowland, tropical,

s

semlarld (7-9 dry months/year) lsothermic areas, and seasonally

dry or semiarid isothermic highlaods were given second and third

prl0rity respectlvely The most geographically extenslve

cassava-growlng ecosystem ln the Amerlcas lS the wet (0-3 dry

months/year), lowland, troplcal zone ThlS ecosystem also

recelved coverage 10 the exploration campaign, as dld subtroplcal

southern Brazll and Paraguay.

with the exception of Bolivia, explorations were conducted

ln all countrles contalnlng prlorlty areas. In the Caribbean and

Central America, abrupt changes in terrain occur over short

distances, precludlng the generatl0n of reliable homologue maps

(P. Jones pers. com ), therefore, areas vlsited were chasen based

on prlmary sources of cllmate and crop dlstribution lnformatlon

from available weather data, atlases and national agricultural

lnstltutlons. CGM were found 10 673 of the 1264 cassava flelds

surveyed in 13 countries The CGM specimens lncluded in our

analyses were from the 266 of these 673 fields which ylelded

speclmens wlth measurable dorsocentral setae plus an additional

259 Brazlllan specimens from 49 fields nat included ln the

survey The latter speclmens were obtalned by J. G. de Moraes

SpeClmen Preparatlon

CGM were collected, cleared ln lactaphenol and mounted ln

Hoyer's medium as described in Flechtmann (1982) Slldes were

drled for 3-4 days at 40°C before examlnation under a phase

contrast microscope (400 Xl. The dorsocentral setae 01, 02 and

D3 of female specimens were measured The rlght and 1eft

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dorsocentral setae were found to be of unequal length ln some

specimens These were excluded from the analyses. One to 45

mltes per slte were measured for a total of 1862 speclmens The

form of the aedeagus was evaluated for male specimens prepared

accordlng to McGregor (1950).

obtalned 50 high quality slides

Statlstlcal bnalyses

From 120 male speclmens, we

We performed cluster analysis on the means of the lengths of

the dorsocentral setae of CGM speclmens obtained from ea eh

sampling site, using Ward's method (SAS Instltute 1989) to

minimize the varlance withln clusters. We based our declsion on

the number ol clusters to accept on eophenetie correlation

(Sneath and Sokal 1973) Subsequently, we performed a

dlscrlmlnant anaiysis (SAS Institute 1989) to flnd a mathematlcal

funetion for more objective and rigorous classlficatlon of setal

lengths lnto groups We assumed the frequency dlstrlbutlon of

cluster membership was proportional rather than equlprobable,

reflecting our empirical observation that some clusters are more

frequently represented in nature than others. A pooled

covariance matrix could not be specified, therefore, we usad a

quadratic discriminant function based on the estimated mlmlnum

total probability af misclassiflcatlon for normal populations as

given in Johnson and Wichern (1982).

We based the initial cluster analysis on the mean length of

the dorsocentral setae of 1-45 specimens per slte from 266

sampling locations in order to be able to map the dlstrlbutlon

7

I

pattern. However, th1.s approach obscures var1.ab1.11.ty l.n seta1

1ength within cluster s across sites In order to determine how

we11 the relat1.ve frequency of cluster types based on analys1.s of

l.nd1.v1.dual spec1.mens was represented by the mapp1.ng approach, we

ass1.gned each spec1.men to a cluster based on the d1.scr1.m1.nant

function generated from the orig1.na1 data, and ca1cu1ated the

frequency of cluster membership within countr1.es, reg1.ons and

w1.th1.n clusters across sites.

FQ1l0W-UD Stud1.es

After completing the cluster and discrim1.nant ana1yses, we

made several addit1.ona1 stud1.es to invest1gate phenomena observed

during the eva1uation of the specimens, or suggested by the data

We analyzed anomalies l.n the number of tact1.1e and sensory setae,

and appl1.ed severa1 e1ectrophoret1.c techniques to determ1.ne

whether m1.tes with ditferent seta1 lengths or trom different

geograph1.ca1 areas cou1d be d1st1.ngu1.shed electrophoret1.cal1y.

ElectrophoresJ,S

After failing to obtain banda on ge1a stained for glutam1.c

oxalacetic transaminase we tested ma1ate dehydrogenase but did

f1.nd not polymorphism. We report resulta of e1ectrophoreses

performed on speeimens of CGM from severa1 sites in Colomb1.a

(Malagana [n-30] and ArJona (n=30], Bo1ivar; Luruaco [n=30],

At1ant1CO¡ and Palmira, Valle [n =48)) and Brazi1 (Cruz das

Almas, Bahia [n=59]) to determ1.ne

1) whether geographical races cou1d be distinguished and

2) whether raes and setal length are related

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We used vertlcal polyacrylamlde slab gradlent and

dlscontlnuous gels (HinlBioRad) prepared according to methods

modlf1ed from Hussain fi al. (1988) and Poehllng & Neuhoff

(1980), with one mlte per sample and stalning tor (1- and B­

esterases.

Anemal1eS ln the number ef tactlle and sensory setae ln CGH

The number of tactlle and sensory setae on tarsus 1 and

tibia 1 were recorded for a subsample of 20-29 randomly selected

female specimens from each cluster. The frequency of occurrence

of anamalies in the number af tactile and sensory setae was

classitied according to cluster membership. Ths anomaliss were:

1) presence of more than one sensory seta on eitherkarsus

1 or tibia 1 ("mascullnization", according to Guttierez

1987);

2) differences between the 1eft and right tarsus 1 and/or

tib1a 1 in the number of tactile or sensory setas¡

3) more or less than five tactile setae on tarsus I¡

4) more or less than nine tactile setae on tibia 1

X2 goodness-of-flt tests were applied to the data.

RESULTS

Dlstr1butlon of Mononychellus $pecles ln the NeotroplCs

One thousand two hundred sixty four cassava fields were

surveyed 1n 13 countrles (Colombla, Venezuela, Trlnldad and

Tobago, Brazil, Cuba, Hexico, Nicaragua, Honduras, Panama, Peru,

Paraguay, Guyana, and Ecuador; see Table 1» Forty flve and 47\

9

respectively of cassava flelds where M tanaloa was detected were

ln humld lowland or seasonally dry lowland zones The remalnlng

8% were dlstrlbuted between semlarld Iowlands, and humld,

seasonally dry and semiarid hlghland ecosystems. Forty-two and

49% respectlvely of the cassava fields samples ln humld (n=544)

and seasonally dry lowlands (n=488) were infested wlth CGM Only

18% of 112 flelda surveyed ln semiarid lowlands were lnfeated and

12 of these 20 fields were ln northeast Brazil

M. carlbbeanae was present ln 64% of the

surveyed ThlS species was not found ln Brazil

Outside Brazl.l,

semiarid fields

M tanal0a-was present in 56% of 52 fielda in our survey of

northeast Brazil, and in 89% of 427 additional fielda sampled by

Moraes (unpubllshed data) for an overall frequency of 85%. In

Colombla and Venezuela, H tanaloa was ldentlfled ln 48 and 85%

respectively of flelds vlslted

Ecuadorian sites or from 92

Nlcaragua, Honduras, and CUba

None were collected from 132

sites visited in Mexico, Peru,

The most frequentIy encountered

species of Mononychellus ln Ecuador, Mexico, Nicaragua and Cuba

was M carlbbeanae (McGregor) (Table 1) The only regl0ns where

M carlbbeanae was not detected were ln Brazll, Peru and Paraguay

(Fig 1). Records ot M carlbbeanae in Brazil have appeared ln

several unpublished reports (Yaseen 1977, 1978, Yaseen and

Bennett 1978), however, these do not mention how the specimens

were identified, nor were speclmens available for examlnatlon (P

Baker, Dlrector, CAB Internatl0nal, Trlnldad and Tobago Statl0n,

pers com.) Apart from thia dlscrepancy, good agreement was

found between the results of our survey and published reports on

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the d~stribution of Mononychellus spec~es ~n the Neotrop~cs

(Table 2) M. car1bbeanae was not ~dentif~ed in Moraes'

(unpubl~shed) survey of 427 cassava f~elds ~n northeast Braz~l.

We found M. rncgregorJ. in the humid highlands (interandean

valleys) of Colombia, Ecuador and Peru, in the Colombian reg~on

of the Amazon Basin land in the state of Santa catarina in

southern Brazil (Table 1, F~g. 1). We could not confirm

unpublished reports (Yaseen & Bennett 1978) of this spec~es ~n

TrJ.n1dad.

Dorsocentral Se tal Lengths of CGM

Measurements of the dorsal setae of CGM collected in the •

Amer~cas indicate the presence of short setal forms fitting the

descr1pt~on of M. tanaloa, and long setal forms fitt~ng Doreste's

(1981) or1g1nal descr1pt~on of M. progres~YUs. Intermed~ate

types between the two extremes also occur in a continuous

grad1ent (Fig. 2). M tanaloa populat~ons were d~v~ded into f~ve

clusters based on the mean lengths of the setae 01, 02, and 03

(Table 3), resulting in a multivariate r 2 of 0.87 The dec~s~on

to accept five clusters was based on cophenetic correlation,

arb1trar~ly high values of r 2 could be atta~ned by further

1ncreasing the number of clusters, however, the increase

resulting from adding an new cluster was small when the number of

clusters exceeded five The division into five clusters provides

an heur~st~c nomenclature for referr~ng to setal length.

Clusters can conven~ently be called very short, short,

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lntermediate, long and very long (Table 3, Flg. 3) and wl11

henceforth be referred to as morphotypes

The multlvarlate dlstances between morphotypes were

statlstlcally signiticant (Wilk's Lambda' F = 100.06, df = 12, P

= o 0001) Morphotype membership was assigned differently by

dlscrimlnant analysis for 1 9% of the 266 samples. This low I

apparent error rate suggests that accepting flve morphotypes lS

statist1cally robust

appendix.

The discrlminant function lS given ln the

A1l morphotypes wera found in in Colombia, Brazil and

Venezuela where sample sizes were large (Table 4). Elghty-two %

of Sltes (Table 5) and 85% of speclmens (Table 4) fIlOm Brazil

were classified as having e1ther the very short or short

morphotypes In Colomb1a 20% of sites (Tabla 5) and 18% of

speclmens were of the short morphotypes In Venezuela 97% of

sites (Table 5) and 93% of specimens (Table 4) were of the short,

lntermedlate or long morphs

Fewer than 15 sites were sampled 1n each of the other

countries where H. tanaloa was collected. In Paraguay, both

extremes of var1ability were found (Table 5) and analysls of

1ndividual speclmens revealed m1tes of all but the lntermedlate

morphotype in a sample of 11 females In Trlnidad, 100% of the

elght sites had the long morphotype (Table 5), and indivl.dual

specl.mens (n=64) of all morphs except the very short were found

(Table 4) In Panama 80% of sltes (Table 5) and 94% of

individual specimens (Table 4) were of the short morph The long

morphotype occurred in the remaining 20% of sltes (Table 5)

12

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In general, where sample s~ze was adequate, good agreement

was obtained between analyses based on mean setal lengths from

each collection sl.te and from l.ndivldual specl.mens When the

l.ndividual specimens from all sites claSSl.fied to a given

morphotype were claSSl.fled by the discrlminant functl.on, the

eXl.stence of polymorphlsm within morphotypes across sites emerged

(Table 6) We found less polymorphism at the short extreme of

the phenotypic range than at the long extreme. sites classified

as having the intermediate morphotype are highly polymorphic

(Table 6)

Geographical analysl.s of the unusual distributl.on of setal

length in Brazil revealed that the skewness towards tme short

morphs was due to the high frequency of short morphotypes in the

northeast regl.on (Fig. 4). Fifty-three of the 54 populatlons

from northeast Braz~l had short or very short mean setal lengths

(Table 7). The other 1.8% of sites and 6 1% of specimens were of

the l.ntermediate morph (Table 8). Northeast Brazil was the only

large contiguous region sampled where mites with long setae were

not found (Fig 4, Table 8)

Of the 266 sites analyzed, only fourteen were in h~ghland

areas. One hundred six sites were ~n humid lowlands, 123 were l.n

seasonally dry lowlands and 22 were l.n seml.arl.d lowlands. AIl

setal morphs occurred in humid lowlands and in seasonally dry

lowlands (Fig 5) The short morph had a higher frequency than

expected l.n semiarid lowland areas (X2 = 36.78, 4 d f P ~

o 001)

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The Aedeagus

Varlation ln the form of the aedeagus of male speclmens from

dlfferent sites was negllglble (Fig. 6). The ahape of the

aedeagus was simllar to a drawlnq made by Tuttle ~ li (1977)

for H tanalOa. The aedeaql examlned did not resemble the

drawings made by Flechtmann (l982) or Gutierrez (1987), however,

lt lS not clear whether the latter drawlng was based on

conventional preparations or on males prepared according to the

mounting technique described in Gutierrez (lgaS)

~ory and Tactl1e Setae

Doreste (1981) reported 4 tactlle and 1 sensory setae for H.

tana10a and 4 tactile and no sensory setae on tarsuSl' 1 for H

progres1vus when he proposed the eXlstence of three different

apecles ln the group then called H. tana10a We examlned 20-29

speclmens chosen randomly from each morphotype. All had sensory

setae on tarsus l. In general, five tactile setae were present

on tarsus 1, and nine were present on tlbia 1, as described by

Flechtmann & Baker (1970) and Nokoe & Rogo (1988) for M. tana10a.

However, several types of anomalies were observed 32% of

speclmens had fewer or more than five tactile setae on tarsus 1¡

O 8% had fewer or more than nlne tactile setae on tlbia 1, O 8%

of specimens were masculinized, with more than one sensory seta

on tarsus 1 and/or tibia 1 (see Gutierrez 1987), 29 and 11 % of

specimens had diferences between the number of right and left

tactile and sensory setae on tarsus 1 and tlbia t, respectively

The probability of possessing normal morpholegy was equal fer all

14

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((~~ 0\11 \~ ~-,L~Jtt~-\LJ

morphotypes (Table 9, X2 goodness-of-fit test, NS) With respect

to specific types of anomal1es, thl probabbr1~~~~~s)ing I ¡..--:;=--~ _____ ~-:::-:;--:;==;::T;"-=---

1) unequal number of r1ght and left sensory and tactile

setae or

2) more or less than 5 tact1le setae on tarsus 1

was equal for all morphotypes (Table 9, X2 goodness-of-fit test;

NS)

Electrophoresl.s

Polymorph1sm waa not found for malate dehydrogenase or

glutaml.c oxalacetic transaminase. The banding patterns for a-

and S-esterases from M. tanaJoa collected from the Caribbean

coaat (Malagana and Arjona, Bolivar, Colombl.a; Luruaco,

Atlantico, Colombia), an interandean valley (Palmira, Valle,

Colomb1a) and northeast Brazl.l (Cruz das Almas, Bahl.a) were

l.dentl.cal (Figs 7,8), however, greater esterase actl.V1ty,

expressed as darker bands, was consl.stently found 1n spec1mens

wl.th short setae (Fig. 8)

DISCUSSION

MQrpholQql.cal Var1abl.ll.ty

Our data corrobora te Rogo'a ~ Al (198S) conclus1on that M

prQgreSlyuS and M. tanaJoa compr1se a slngle polymorph1c speC1es,

and that morphotype may be associated Wl.th the geograph1cal

orig1n of specimens

Rogo ti al (1987) report a range in varl.ation in setal

lengths for CGM collected in Afrl.ca sl.m1lar to that reported he re

15

for northwest s Amerlca, Central and Soutbern Brazll and

Paraguay Assuming that setal length la genetically determlned,

lf the lntroductlon of CGM to Afrlca was from a alte Wlth a

hlghly polymorphlc populatlon, a sl.ngle lntroductlon of CGM to

Afrlca could account for the polymorphl.sm ln Afrl.ca. It l.S

unllkely that the origin of CGM in Africa was from northeast

Brazll, where varlabillty l.n setal length lB ll.ml.ted.

A similar degree of variability in length is present in the

lateral hysterosomal setae of M. carlbbeanae (Guerrero unpub.

data) The short setal forms flt the the descrl.ptl.on of M

capbbeanae (McGregor 1950) and the long setal forms fit the

description of M erythrl.nae (Tuttle fi al 1976), a speeies

described from Mexieo on Erythrl.na sp A continuous gradient of

l.ntermedl.ate forms oecurs between these extremes (Guerrero unpub

data ).

Dlstrlbutl.on and EcolQqlcal Adaptatlon of Mononychellus sPD.

Comparison of our collection reeords of MQnonychellua

apecles on cassava in the Neotropics with reports in the

llterature indicates that M. tªnaJoa lS present

Venezuela, Brazil, Guyana and Paraguay (See Table 2)

ln Colombla,

In Central

Amerlea CGM has been reported in Panama and Costa Rlca, and ln

the Caribbean, in Trinidad and Tobago and Haiti. M. bondap

Paschoal, which we consider to be a junl.or synonym of M tanª,oa,

which has been reported once froro cassava in Brazil, and once

from Colombia (see Table 2) was not detected ln our survey We

, found M. mcqregor. ln interandean valleys of colombla, Ecuador,

16 (

(

and Peru (Fig 1) and ~n subtrop~cal southern Brazil,

corroborating the reporte of Samwaye &: c1ocio1a (1980), but were

unable to conf~rm unpubl~shed reports (Yaseen 1978; Yaseen &:

Bennett 1978) of thlS specles in northeast Braz~l or Trlnldad and

Tobago. H !::~rlbbeanae occurs from southern Florlda (Peña &:

Wad~l1 1982, Peña fi al. 1984), throughout the Caribbean bae~n

(see Table 2) and ln Ecuador, however we could not corroborate

unpubIished reporta of thia species in northeaat Brazil (Yaseen

1977, 1978, Yaseen &: Bennett 1978). Other Neotropical spec~es of

Mononychellus (see Table 2) have been reported primarlly from

MeX1CO on species other than Han¡bot. Northeast Brazil and

Paraguay are unlque ln that neither H. carlbbeanae nor H. I

rncgregor+ were detected in our survey.

ti carlbbeanae was collected in zones with o to 9 dry

months/yr, and wlth mean annual precipitat~on between 401 to 3023

mm/yr The mean number of dry months/yr for these sites was 5 6

compared to 3 1 for H tana10a, indicating that M can.bbeanae

distribution ie ekewed towards eubhumid areas, whereas CGM

dlstrlbution le skewed towards more humid zones. The absence of

ti !::an,bbeanae in northeast Brazil le particularly noteworthy

glven the sizeable seasonally dry to semiarid cassava-growlng

area where M. car¡bbeanae would presumably be well adapted. The

abaence of M. mcgregor¡ ln northeaat Brazil, on the other hand,

lS not as surprising, Slnce 80\ of cassava fieIda where this

species has be en detected in our survey were in hum~d interandean

va 11 eys

17

Qnq1n af the Traphl.c Relatl.ansh1p between Mononychellus and

cassaya

The largest number of specl.es of Mononychellus on cassava

occurs in colombia, with the number dropping off wl.th distance

both towards Central America and the Caribbean, and to the south,

suggesting a center of genetic diversity for the genus. The

antl.quity of cassava cultl.vatl.on in northwest S. Amerl.ca l.S well

documented (Shultes 1987), and thl.s region is an area of prl.mary

genetic diversl.ty of cassava (Gulick ~ al 1983) and may be a

one af several possible areas af domestication (Sauer 1969,

Lathrap 1973; Spath 1973, Renvol.se 1973). Colombl.a also has the

greatest diversity of phytoseiid predators of tetranyat'lld mites

reported on cassava (CIAT 1991, Botelho ~ Al. submitted).

Toqether these patterns'pol.nt to a possl.ble area af Orl.g1n of the

trophic assocl.atl.on between Mononychellus and cassava 1n

narthwest S Amer1ca

HypQtbeses about Mononychellus tanaJoa 1n Northeast Sra;l.l

The absence in northeast Brazil af the setal polymarph1sm

associated with H. tanaloa throughout the rest of its range l.S

un1que. CGM with short dorsocentral setae can be distl.ngul.shed

electrophoretl.cally from specl.mens wl.th long setae by thel.r

enhanced esterase actl.vl.ty, suggestl.nq sorne physl.ologl.cal

differentiation and the possible existence of a geographl.cal

subpopulation in northeast Srazil It is striking that in

Moraes' survey of 427 cassava fields in northeast Srazil (CIAT

1990, 1991) and in cassava germplasm screening sites (CIAT 1992),

18

(

heavy 1nfestations of CGM were found 1n sem1ar1d areas, a pattern

not seen elsewhere in trop1cal America or in Africa, where CGM

does not appear to have colon1zed sem1ar1d areas of Nlger1a (M

Porto pers. com ) or Benin (Yaninek & Onzo 1988, unpublished)

A hypothes1s Wh1Ch accounts both for the absence of other

MODQnychellus species and tha limitad setal polymorphism skewed

toward the short extrema of polymorphic variab11ity 1n H tanaJoa

1n northeast Brazil ls that CGM was introduced inadvertently to

tha area. Tbe absence of the long morphs over such an extensive

area could have come about if the founder population had short

setae, and a high degree of genetic ls01at1on was ma1nta1ned.

Alternatively, after introduction of a polymorphie foundar

population, selection may haYa favored the short morphotypes

lead1ng to elimlnation of the others. The absence of competltlon I

from other HODonychellus species, particularly H. carlbbeaDae may

have centributed to the success of the short morphotypes of M

tanaJoa 1n northeast Brazil and to its unique adaptation to

semiarid environments Molecular techniques have recently been

applled ln acarological phylogenetlc research (Kallszewski ~ ~

1992, Navajas ~ Al. 1992) The application of these technlques

ln studies ol phylogenetic relationahips of MODonycheUus spp

may provida a conclusive meaDS to asseas whether a dlstlnct CGM

b10type occurs northeast Brazil.

The variability in setal length, the relatively hlgh

frequency ol anomalies ln tha numbers ef tactile and sensory

setae lound in H. tanaJoa, and the high frequency el simllar

phenomena in H. ~rl,bbeanae suggests that rapid evolutienary

19

(

change is occurrlng in these specles in the Americas, perhaps ln

response to the great range of edaphic and c1imatic factara under

WhlCh cassava is grown We co11ected H tanal0a over a range of

altitudes (2-1820 m) I and precipitation zonea (425-4468 mm/yr)

wlth wldely varylng ralnfal1 patterns (1-9 dry months/yr). The

diversity of eco10gica1 conditions under wnich cassava has been

cu1tlvated ln the Andean zone and the patchiness of cassava

dlstribution there (Carter 1986) are both re1ated to the

mountalnous topography of the region. Less agroecologlca1 and

topographica1 variabi1ity occurs in northeast Brazil, where we

collected H. tanaloa from 30-900 m aboye sea level in rainfall

zones from 425-2083 mm/yr Agroecological diversity &nd patchy

dlstribution may have been incisive in the speclation of

Mononychellus assoclated wlth Mam.hot and ln the appearance ol

seta1 polymorphlsm in CGM and H. carlbbeanae

Impllcat10ns for Bl0loQ1C81 Control

CGM is considered an lmportant pest of cassava in northeast

Brazll, particular1y in seasona11y dry and semiarld areas (Velga,

1985) Alternatives for control of CGM In northeast Brazll have

focused on host plant reslstance, and lmproving the leve1 of

resistance to CGM la a hlgh priority for cassava breeders worklng

ln the states of Ceara, Paralba, Pernambuco, A1agoas and Bahla

(C Iglesias pers. com.). To date no effort has been made to

lmprove leve1s af bl0109ica1 control, even though the phytoselld

fauna in cassava all agroeco10g1ca1 zones of northeast BraZl1 ls

les s dlverse than that in homo10gous zones in northwest S

20

(

Amer1ea (CIAT 1991, Moraes et al submitted) Introductl.on of

exot1c species or strains may provide an ecoloqically sound pest

management opt10n in northeast Brazil 1f specl.es or stra1ns whieh

are well adapted to sem1arid condltions can be found The

largest diversity of phytoselid species in a sem1arl.d area occurs

1n eoastal Ecuador (Braun 1993). Introduction of exotie natural

eneml.es for control of CGM 1n Brazil should be 1mplemented in

eonJunetion w1th other plant protection mea sures sueh as

augmentation and conservation of natural enemies, habitat

manipulation and the deployment of host plant resistance

The pOSSl.b11ity that the troph1c relationship between

Mononychellus and HanlhQt evolved in northwest Sk Ameriea

suggests that this are a should reeeive high priority as a souree

of phytoseil.d predators and fungal pathogens (H~ozyq1tes spp.) of

natural enem1es of CGM for introduction to Afr1ca and Braz11.

A hlqh degree of host specificity of Neozyq1tes spp. can be

dedueed from the difficulty of eulturing this fungus on

art1f1c1al media (Alvarez 1990, Evans 1991). Since our data

p01nt to differenees in morphotype composition and ecolog1cal

adaptation between CGM from northeast Brazil and Africa, we

recommend broadeninq the effort to introduce fungal pathogens

from northeast Brazil to lnelude Neozyqltes speciesjstral.ns

obtained elsewhere 1n the Americas

The use of ecologieal homologue mapping fer priorit1zing the

seareh fer natural enemies and decidinq whieh natural enemies to

lntroduce to particular regions of the Afriean cassava belt has

been proposed by Yaninek & Bellotti (1987), and 1S compatlbl.le

21

(

w1th a strategy based on searchlng 1n the area of or1g1n of the

trophlc relationship between CGM and cassava.

ACltNOWLEDGKENTS

We thank Ruben Escobar for his hard work on manag1ng the

foreign exploration and setal length databases, and the CIAT Land

Use Program for assistance in map preparation. We are indebted

to three anonymous internal reviewers from CIAT and to steve

Yan1nek and Lucy Rogo for their comments and suggestions for

improvement of the manuscript

i

22

(

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Yan~nek, J S & Bellott~, A C 1987 Exploration for natural enem~es of cassava green m~tes based on agrometeorolog~cal cr~ter~a pp. 59-67 ~ R1)ks, O & Mathys, G , (Eda ) Proc Seminar on Agrometeorology and Crop Protect~on ~n the Lowland Hum~d and Subhum~d Tropics, Cotonou, Ben~n 7-11 July, 1986 World Meteorolog~cal organ~zat~on, Geneva.

Yan~nek, J S & Herren, H R 1988 Introduction and spread of the cassava green mlte, MQoonychellus taoal0a (Bondar) (Acari Tetranychldae), ~n exot~c pest ln Afrlca and the seaerch for appropriate control methods a reVlew Bull Ent Res. 78 1-13.

Yaseen, M. & Benoett F D 1977. Distribution, blology aod population dynam~cs of the green cassava m~te ~n the Neotropics. pp 197-202 In Proc. 4th symp. lnt. Soc Trop Root crops Cali, Colombla. (Cock, J., Maclntyre, R & Graham, M. Eds.) IORC, Ottawa. IORC-080e

Yaseen, M., Bennett, F O, OeVoogd, W & Girling O J. 1979. Investigatlons on the cassava mita Mooooychellus taoalQa (Bondar) and lts natural enemles An tha Neotroplcs and East Afrlca Flnal Report. October 1976-March 1979 ClaC 20pp

Yaseen, M 1988. Invest~gatlons of cassava mites of tanaJoa complex and the~r natural enemles ~n the Neotroplcs Vllth. Symposlum of the Int SOCo for Trop. Root Crops, Gosier, Guadalupe, 1-6 July 1985

Zuluaga, l. 1971 Llsta prellmlnar de acaros de lmportancla economica en Colombia Acta Agron. 21 119-131

Table 1 Inventory of Mononychellus spec~es on cassava ~n

the Amer~cas.

f~elds resent

Brazll 52 M. tanal0a 29 M DlankJ. 1 M maqreqorlo 1

CUba 43 M canbbeanae 23

Ecuador 132 M !;; ,11:: ;¡'Qbe,m,u: 65 M. mcgreqorl 13

Colombia 869 M. tanalOª 420 M. !;; a UQQ!il2mUI 71 M. m!::g¡;:~gQ¡;:. 59 M. plankJ. 5

Mexico 27 M canbb!ilanae 16 •

N~caraqua 2 11 canbbe2D2e 1

Honduras 3 M canbbe2nae 3

Panama 17 M !::2UQQ!ilana!il 13 11 tanaJoa 3

Venezuela 84 M taDalQa 76 11. can.bb!ilanae 36

Peru 14 11 mcqregor. 5

Paraguay 7 M· tanalQa 5

Trinldad , 12 11 !;;an!:l!:l!il5lD5l1i: 11 Tobago

11 tanaJOa 8

Guyana 2 1 1

Table 2 Mononychellus specles reported ln the Neotrop1cS

I Specles

t:I bispidosetus t:I hyptis2

1:1 estrada¡)

t:I ervthnnae2

1:1 flabellosetus 1

ti chapalensis2

1:1 wl11ardlae2

t:I walnste¡n 2

t:I eysenbardtiae2 t:I tephrosiae2

ti psldium4

ti yilancens.l.s 1:1 bonelari 5

t:I mcgreqQu 6

(contlnuedl

j Country

MeX1Coª Hexlcoª Hexlcoª Nicarªguaª Mex1COª Mexl.coa

Mexlcoª Hexl.coª Hexl.coª Mexlcoª Mexlcoa

Mexlcoa

Brazlla

Brazll

ColombIa BraZ1l Coloml:na

Trlnldad Argentlnaª Panama Peru

Reference

Beer & Lang 1958, Tuttle ~ Al 1976 Tuttle ~ Al 1974, 1976 Tuttle ~ Al 1976 Baker & Pritchard 1962 Tuttle ~ al 1976 Beer & Lang 1958 Tuttle ~ Al 1976 Tuttle ~ al 1976 Tuttle ~ al 1976 Tuttle ~ al 1976, 1977 Tuttle ~ al 1976 Tuttle ~ Al 1974 1976 Estebanes & Baker 1968, Tuttle ~ al 1976 Paschoal 1971c Paschoal 1970a, Flechtmann & Baker 1970, Yaseen & Beonett 1978b

Urueta 1975 Yaseen 1978b , Samways & Cl0clo1a 1980, urueta 1975 Yaseen & Bennett 1977b , 1975b

Guerrero & Bellottl 1980, CIA~ Yasseen & Bennett 1978b

Prltchard & Baker 1955 CIATc CIATc

Table 2 (cont) MODonychel1us specles reported ln the Neotroplcs

I Specles

~ mamhotl

H progreslyus8d

ti tapaJQa9

I Country

Venezuela Trimdad BollVla, ColombIa Venezuela Paraguay, Trimdad Branl

Paraguay

Costa RIca ColombIa

Venezuela

Hanl Guyana, TrInIdad Bahamas, SurInam French Guyana Panama

a Reports on host plants other than Man¡hot spp b Unpubhshed

Doreste 1980, 1981 CIBC 1982b

Yaseen 1978b

Yasean 1978b

Yaseen 1988 Yasean 1978b

RefereDce

Bondar 1938, Paschoal 1971a, Flechtmann & Abreu 1973, Flechtmann & Bastos 1972, Costa 1975, TUttle ~ al 1977, Yaseen & Bennett 1977b , 1978b , Flechtmann 1987, Yaseen 1978b , Farias ~ al 1978, 1979, 1981, Samways & ClocIo1a 1980, ClATc Aranda & Flechtmann 1971, ClA~, Yaseen & Bennett 1977b

Salas 1978 Urueta 1970, 1975, Yaseen & Bennett 1977b , 1978, Guerrero & Bellott! 1980, ClA~ QUIroz 1977, Yaseen & Bennett 1978b , Doreste 1981, CIA~ Lenolr ~ al 1981 Yaseen & Bennett 1977b , 1978b , CIA~ Yaseen 1977b , Yasean & Bennett 1978b

Yasean U78b

CIA~

c Speclmens deposlted In a reference collectlon at Centro ~nternaclonal de Agrlcultura Tro¡ncal (CIAT)

(Contmued)

Table J

Cluster

1

2

3

4

5

Mean dorsocentral setal lengths of female Mononychellus tanaloa from 266 s~tes ~n the Amer~cas

Mean length Range

Morphotype n Seta (¡.1m) (¡.1m)

Very short 29 DC1 20 37 17 85-22.24

DC2 21.45 17 85-25.50

DC3 24 87 20 40-28 22

Short 73 DC1 24 05 20 40-26.78

DC2 25 32 22 95-28 90

DC3 30 68 26 07-37 05

Intermed~ate 32 DC1 25 64 22 10-28 05

DC2 28 26 25 50-30 60

DC3 38 83 33 15-45 05

Long 94 DCl 28.85 23.80-34.00

DC2 34.99 28.90-45 90

DC3 46 35 35 21-61 20

Very long 38 DC1 36 17 30 60-54.40

DC2 44 82 34 00-56 10

DC3 54 19 44 20-64 60

..

I

· ,

Table 4 Distribution of dorsocentral setal length morphotypes ~n Mononychellus

tana10a spec~mens from the Amer~cas.

, spec~mens/morphotypel

No

spec~mens

Country measured 1 2 3 4 5

Braz~l 611 28 6 55 8 12.1 1 O 2 5

Colomb~a 716 2.9 15.5 24.3 16.8 40.5

Venezuela 440 1.6 25.9 48 9 18 6 5 O

Tr~nidad 64 O O 7.8 84 4 6 25 1 6

Panama 18 O O 94.4 5.6 0.0 O O

Paraguay 11 36 4 9 1 0.0 45.5 9 1

Guyana 2 0.0 O O O O 100 O O O

1 l=very short, 2=short¡ 3=~ntermed~atei 4=long¡ 5=very long. ,

• •

1111111111111111111111111 1: I

Table 5 Geoqraph~c d~str~but~on of Mononyche11us tanaJoa dorsocentra1 seta1

length morphotypes ~n Bra~~l.

% s~tes/morphotypel

No

s~tes

Regl.on2 sampled 1 2 3 4 5

North 5 100 O O O O O O O O O

Northeast 54 27 a 70 4 1 a 0.0 o o Central West 16 12 5 12 5 la a 31 3 25 O

Southeast 8 12 5 62.5 12 5 12 5 0.0

South 1 100 O O o O O 0.0 O O

1 l=very short, 2=short¡ 3=~ntermed~ate, 4=10ng, 5=very long

2 Sta tes sampled l.n reg~ons are North-Amazonas, Northeast-Ceara, P~au~1 Bah~a,

Alagoas, Sergl.pe, Paralba, Pernambuco, Maranhao, Central West-Matto Grosso do , Sul, Bras~lla O F , Southeast-Sao Paulo, South-Santa Catar~na.

Table 6. Variab~l~ty w~th~n dorsocentral setal morphotypes of Mononychellus tanaloa from the Amer~cas.

% spec~mens/morphotype1 No

females Horphotype measured VS S 1 L VL

vs 208 60.4 39 6 0.0 0.0 0.0

s 589 5.5 88 6 5.9 o O 0.0

1 518 o 4 42.0 29.0 27.8 o 8

L 218 0.0 10.3 12.1 63.5 14 2

VL 329 o O 1 4 2.1 26.0 70 6

1 VS=very short, S=short, I=~ntermed1ate, L=long, VL---very long.

Table 7. Geographic d1str1but1on of MonQnychellus tana10a dorsocentral setal length morphotypes in BraZ1l

% sltes/morphotype1

No.

sltes Reglon2 sampled VS S 1 L VL

North 5 100 O O O 0.0 0.0 0.0

Northeast 54 27.8 70 4 1.8 0.0 0.0

Central West 16 12 5 12 5 18.8 31.3 25.0

Southeast 8 12 5 62 5 12.5 12.5 O O

South 1 100 O O O O O O O O O

1 VS=very short, s=short, I=lntermedlate; L=long, VL=very long

2 States sampled 1n reglons are. North-Amazonas, Northeast-Ceara, P1au1, Bah1a,

Alagoas, Serglpe, Paralba, Pernambuco, Maranhao¡ Central West-Matto Grosso do

Sul, Brasll1a D F , Southeast-Sao Paulo, South-Santa Catarlna ,

i

Table 8. Dlstribution of MODonychellus taoaJoa dorsocentral setal length morphotypes ln Brazil.

t speclmens/morphotype1

No speclmens

Rec:Jlon2 measured VS S 1 L VL

Nortb 15 100.0 O O 0.0 0.0 0.0

Northeast 459 29.4 64.5 6 1 0.0 O O

Central West 73 16.4 17.8 38.4 6 8 20.5

Southeast 58 13.8 55.2 31.0 0.0 0.0

South 6 100 O 0.0 O O O O O O

1 VS=very short, s=short, I=lntermedlate¡ L=lonq¡ VL=very long. 2 States sampled ln reglons are. North-Amazonas¡ Northeast-Ceara, Plaul, Bahla,

Alagoas, Serglpe, Paralba, Pernambuco, Maranbao¡ Central West-Matto Grosso do Sul, Brasllia D.F.¡ Southeast-Sao paulo; South-SanWa Catarlna.

Table 9 Comparative trequencies ot anomalies in the sensory and tactile setae of Mononychellus taDaloa morphotypes

/MorPhotype2 Anomaly type1

n 1 2 33 44 5

very short 23 14 1 6 7 O

short 20 10 O 9 10 O

intermedlate 25 14 O 6 10 O

long 29 20 O 7 9 O

verv long 25 21 O 3 3 1

1

2

3

4

1 = normal; 2 = mascullnlzed; 3 = unequal number of rlght and left sensory or tactlle setae; 4 = more or les s than 5 tactlle setae on tarsus l; 5 = more or less than 9 tactile setal on tibia l. The probability of possessing normal morpholoqy ls equal for al1 morphotypes (X2 = 5 951 df = 4, p = O 20, N5).

Morphotype was determined by cluster analysis of 1engths of dorsocentral setae 01, 02 and 02.

The probability of possessinq an unequal number of riqht and left sensory and tactile setae is equal for all morphotypes (X2 - 4 96, df a 4, P - 0.29, N5).

The probabillty of possesslng more or less than 5 tactile setae on tarsus l ls equal for all morphotypes (X2 = 5 73, df - 4, P - 0.22, N5).

(

Fl.gure Legends

1 The geographl.cal dl.stributl.on of H. tanal0a, H

carl.bbeanae, H plank. and H. mcgregorl. in the Americas

Fl.g 2 Varl.ation 1n the length of the dorsocentral setae 01, 02,

and 03 of H tanaJQa (Morphotypes: A = very short¡ B =

short, e = l.ntermediate¡ o = long).

F1g. 3 eontinuous gradient 1n variability in the mean length of

the dorsocentral setae D1, 02, and 03 of H. tanaloa from 266

sites 1n the Americas and their distribution 1n morphotypes.

Fig 4 Geographical distribution of H. tanaJQa morphotypes in

the Americas.

Fig. 5 Frequency of H. tanaJoa morphotypes in huml.d (HL),

seasonally dry (SOL), and semiarid (SAL) lowlands of the

Americas (Morphotypes: VS = very short, S = short, 1 =

lntermediate, L = long, VL = very long).

Flg 6 Aedeagi ol male H taDaloa from Colombia (A), Venezuela

(B) and Brazil (e)

(

F1g 7 Band1nq patterns for a- and B-esterases from H. tanaJoa

from the Caribbean coast (Malagana (lanes 1-31 and Arjona

Clanes 4-6], BOlivar, colombia, Luruaco (lanes 7-9],

Atlant1co, Colombia)

Fig 8 Banding patterns for a- and B-esterases from H tanaJoa

w1th short dorsocentral setae from an 1nterandean valley

(Palmira, Colombia; lanes 1-3) and Nortbeast Brazil (Cruz

das Almas, lane 8), and with long setae from an interandean

valley (palmira, colombia¡ lane 4-7).

" ,

Seal. 1 32000000 approx

Specles

• M tanaJoa o M canbbeanae

q M mcgregon

O M plankl

o

• • e l' .' ,

~ :~ .{ . ~. .:I' .OJ-

:~ J.y

.' ~~~1iI ¿U 1 v Q:" . ' ., .

• • • • • •

•

•

Seale 1 35000 000 approx

•

• ••• ---1------------/. ,,------------------

",/ _, ._ 6 A

... / ,,¡. •• • • • ,/ • ",,6> •

, A A

'E ro /' / --~-"-At3-:~~: .. --------------------~ //// '~_r_¡,--__;r------.J

,/ t,/ .:a"'~ " --------- /'. ~ / " ---/~-----~ // /' O---(tOO-,L-------/' o &.1 ---

_________ .L/ ~ __ ~¡------------ • .f 'Oh. / - ..... --.p.

20S----------4i~'/~--~~--3&~"~'--~~·-~-~-·-/---58 ' , '. ., 44 ' • 30 ...

D2l~m) 1617

Setal

Cluster Leng\h

-=----1 • Very Shoet

2 o Shoet

31>. \n\ermedl¡ile

4 A Long

5-Vely Long

.. San Andrés

~-~ . . ,- ':r~ I~)C Panama

• .Iil

1'1 -• •

A Seal. 1 14 500 000 approx

A • " • l+l ... • ",}., " ... tI • . , ..

VJ I'J

o

Colombia

Seal. 1 22 600 000 approx

T",,,,hd& ToIJflt}o

··'---.~LJ=-"' ,_ l' .~--=-X.! ~

(.1

o

Venezuela

•

X Selal LOIl!Jlh (¡mI -----~--

D1 D2 03 Morpholype -----~

" 204 215 249 Very Short

'" 24 O 253 307 Short o 256 203 33 3 Me¡Jlum

• 289 35 O 464 Long o 362 448 542 Very Long

50 ~VS ~S [B 1

40 0TIIIIIl L §VL

>- JO U Z LrJ :::> O LrJ o:: 20 .....

10

o -1----

HL SDL SAL

AGROECOLOGICAL ZONE

(

) .

J

• if~ " , ;;'-

:r t ,:..

, ~ ... ~

~ '<~ , , ... .- • " ,~

" " - '\. .. . , ~. \~- ,

" " , -::; '" -.- .~ " ',: . ..t ~ .-

, - -" .

~"""1-,. .. ' ~ - ... , • ,

~

-~ ... ~ -,

1 2 3 4 5 6 7 8 9

r

, , ' ..... -1 .... iii, \ 1 1"'" • -.'.;

, >

I

1 2 3

..... _ ... -­,

4 5

t t ¡ 1

6 7 8

Append~x

The estLffiate of the quadrat~c d~scr~m~nat~on score ~s

dO(X)= lnlsl+(X-X 1S-'(X-X )-21n(P) .j J l Ji J; 1

x = vector of values, DI, D2, D3, to be ass~gned to a

morphotype

S~ = sample covar~ance matr~x ~n morphotype ~.

x = 1

sample mean vector of morphotype ~.

p~ = pr~or probab~l~ty of membersh~p ~n morphotype ~.

Allocate X to morphotype nk ~f the quadrat~c d~scr~inat~on

A o A o acore d

k (X) ~s the largest of the d. I ~ = 1,2,. .9. The Si

are as followsl

[' ". 2.176 2.37'] S = 2 176 3 900 2.926 ,

2 373 2 926 4 620

U 969 0.146 -O ''']

S = 146 1 321 1 097 2

704 1 097 7 OBO

U 01B O 544

-O "'] S = 544 1 972 -O 026 1

B37 -O 026 9 352

[' 761 2 420 1 '3S] S = 2 420 12 181 8 730 o •

1. 435 8 730 25 062

(17 182 11 539 7 8"] S = 1~ 539 23 528 9 180 5

887 9.180 20 101

..

The X are as followa. ,

[" 371] ~ x = 21 454

1

24 868

[24047] X = 25 319

2

30 681

[25 640] X '" 28 261

)

38 833

[" '52] X '" 34.989 • 46 355 •

["016'] X ::: 44 817 s

54 187