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For pub11cat1on 1n B1olog1cal Control
March 15, 1993
Untll 30 June Address correspondence to
Telephone Telefax Electronlc Mal1 (Bltnet)
1 July - 31 August
As of 1 September
Ann R Braun Cassava Program Centro Internatlonal de Agrlcultura Trop1cal Apartado Aereo 6713 Call, Colombla
57-23-675050 ext 373 57-23-647243 ABRAUN@CIATCOL
9 Wlleatlllll Clase Leamlngton Spa Warw1ck~111re CV3~ 6PL u/u ted l<i/lgdom
Internatlonal Po tato Center ESEAP Reglan c/o Centfal Research Inst1tute
'r----~~-~--~-~"-~"~.=;~~-' ~~~a~o~~r~~~~Sl!;RIFC)
l Telephone Telefax
v
~raPllY w1th
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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
'-,I:¡.,k, '11\
CCM~U LIB~ARY
,
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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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
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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)
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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
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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
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