Agricultural Meteorology, 20(1979) 411--425 411

© Elsevier Scientific Publishing Company, Amsterdam -- Printed in The Netherlands

THE THERMAL ENVIRONMENT OF THE CITRUS RUST MITE*

JON C. ALLEN and CLAYTON W. MCCOY

University of Florida, Institute of Food and Agricultural Sciences, Agricultural Research
and Education Center, Lake Alfred, FL 33850 (U.S.A.)
(Received March 13, 1978; accepted September 19, 1978)

ABSTRACT

Allen, J. C. and McCoy, C. W., 1979. The thermal environment of the citrus rust mite.
Agric. Meteorol., 20 : 411--425.

The distribution of citrus rust mite on individual fruit and in the whole citrus tree
suggests an avoidance of solar exposure. A model of fruit temperatures as affected by
solar radiation in different parts of the tree was constructed, and its predictions are
compared with field temperatures and rust mite distribution patterns. The fruit tempera-
ture model is a function of latitude, time of year, time of day, fruit diameter and
reflectance, position in the tree, atmospheric transmissivity and wind velocity. Where
model predictions indicate unfavorable conditions, rust mite population levels were low.
When the tree was divided into north--south, top-bottom quadrants, the north bottom
quadrant had the most favorable temperatures and usually the most rust mites; the south
bottom was also favorable and had high mite densities. The south top quadrant was least
favorable often having temperatures in the lethal range, and it also had the lowest rust
mite populations.

INTRODUCTION

P o p u l a t i o n m o d e l s o f e c t o t h e r m i c organisms always d e p e n d heavily o n

t e m p e r a t u r e to p r e d i c t d e v e l o p m e n t rates. M u c h has been said a b o u t the w a y
in w h i c h t h e d e v e l o p m e n t a l r a t e ' d e p e n d s u p o n t e m p e r a t u r e (Stinner et al.,
1 9 7 4 ; Allen, 1 9 7 6 ; Sharpe a n d D e m i c h e l e , 1 9 7 7 ) . In the case o f a t i n y
o r g a n i s m such as the citrus rust mite, P h y l l o c o p t r u t a oleivora (Ash.), living in
citrus trees, the p r o b l e m o f m i c r o c l i m a t e variation is as serious as the develop-
m e n t rate p r o b l e m . While d e v e l o p m e n t rates m a y be p r e d i c t e d a c c u r a t e l y f r o m
t e m p e r a t u r e , t h e t e m p e r a t u r e e x p e r i e n c e d b y t h e mites can be v e r y d i f f e r e n t
f r o m t h a t m e a s u r e d in a w e a t h e r shelter.
Several a u t h o r s have n o t e d t h a t rust mites on citrus fruit t e n d t o avoid the
bright sunlit side in f a v o r o f semishade ( H u b b a r d , 1 8 8 5 ; Y o t h e r s and Mason,
1 9 3 0 ; Albrigo a n d M c C o y , 1 9 7 4 ; Van Brussel, 1 9 7 5 ) . O n t h e o t h e r h a n d ,
s h a d e d groves and t h e s h a d e d side o f f r u i t do n o t e x h i b i t mite densities as
* Florida Agricultural Experiment Stations Journal Series, No. 1050.
412

high as semishade areas (Yothers and Mason, 1930; Muma, 1970; McCoy and
Albrigo, 1975). Several hypotheses are suggested by the aggregation of mites
around sunlit areas of which the following seem most plausible: (1) the
aggregation is due to a temperature gradient; (2) the aggregation is due to a
moisture gradient of either dew or water vapor; (3) some combination of
these two hypotheses. The picture is further complicated by the fact that the
rust mite is attacked by a fungal pathogen, Hirsutella thompsonii (Fisher et al.,
1949; McCoy and Knavel, 1969) whose infectivity is dependent on the
presence of free water and high h u m i d i t y (McCoy, 1978).
It is our purpose here to examine the effects of temperature on the
distribution pattern of rust mites in the tree. This is done by developing a
model of fruit temperatures as a function of latitude, time of year and
position in the tree. The model's predictions are then compared with
temperature measurements in the field and with data on rust mite population
dynamics from different parts of the citrus tree.

A MODEL OF TEMPERATURES ON CITRUS FRUIT

We wish to compare surface temperatures at different positions on individual

fruit, at different positions in the tree and at different times of year. This
means that the model must include the effects of solar elevation and azimuthal
angles as well as the heat balance on individual fruit. Our mathematical
notation will be as shown in the Notation.

Energy balance on individual fruit

Equations for the dynamic energy exchange for irradiated citrus fruit
have been obtained by Poppendiek (1953), but for simplicity we have
adopted the equilibrium state equations of Thorpe (1974) and particularly
those of Smart and Sinclair (1976). At equilibrium the energy balance can be
written as:
Energy absorbed = Energy lost by + Energy lost by + Energy lost by
from radiation convection transpirational long wave (1)
cooling radiation

The last two terms, transpirational cooling and long wave radiation, wilt be
ignored as they were by Smart and Sinclair {1976}. Their arguments show that
these quantities are normally quite small compared to convection losses for
grapes under field conditions. Citrus fruit (Valencia orange) has a water vapor
resistance comparable to (Albrigo, 1977) or perhaps even greater than (Nobel,
1975} the grapes used by Smart and Sinclair. Thus, the transpirational
cooling should be no greater on citrus than on grapes. On the input side, only
direct beam solar radiation will be considered.
For the temperature distribution at equilibrium within the spherical fruit,
we can use the fact that the rate of change in the temperature gradient in any
NOTATION

a a l b e d o ( r e f l e c t a n c e ) of spherical f r u i t surface ~ 0.6 for

Valencia oranges (Gaffney, 1973)
2
A area o n t h e surface o f spherical tree m

AA small finite e l e m e n t of tree surface area m2

day J u l i a n day n u m b e r (1 = Jan. 1 ) day
D d i a m e t e r of spherical f r u i t m
h convective heat transfer coefficient Wm-1 °C-I
H solar h o u r angle m e a s u r e d f r o m solar n o o n radians
Io solar b e a m i n t e n s i t y n o r m a l to f r u i t surface Wm-2
IoA average solar b e a m e n e r g y f l o w r a t e t h r o u g h a n area, A, o f W
t h e tree

I 0 QUAD average solar b e a m f l u x t h r o u g h a p a r t i c u l a r tree q u a d r a n t Win-2

u n i t v e c t o r s in t h e n o r t h , east a n d vertical d i r e c t i o n s
ks t h e r m a l c o n d u c t i v i t y of t h e f r u i t (~--0.46 W m -1 ° C - l ) Wm-10C-1
( B e n n e t t e t al., 1 9 7 0 )
ha t h e r m a l c o n d u c t i v i t y of air ( 0 . 0 2 5 5 W m -1 ° C - 1 ) Wm-1 °C-1

n o r m a l v e c t o r to t h e tree surface
distance from center of fruit m
R radius o f f r u i t m
solar b e a m v e c t o r Win-2
So solar c o n s t a n t ( 1 3 6 0 Wrn-2) Wm-2
t hour of the day hr
T e l e v a t i o n o f fruit t e m p e r a t u r e a b o v e a m b i e n t °C
Tcen t e m p e r a t u r e e l e v a t i o n a b o v e a m b i e n t at f r u i t c e n t e r °e
Tmax temperature elevation above ambient at ~ = 0 and r = R °C
("hotspot")
Train t e m p e r a t u r e e l e v a t i o n a b o v e a m b i e n t at/~ = 1 8 0 ° a n d r = R °C
-1
U wind velocity m sec

solar a z i m u t h angle m e a s u r e d clockwise f r o m N o r t h radians

angle b e t w e e n a p o i n t in or o n t h e f r u i t a n d t h e solar b e a m radians
7 angle b e t w e e n t h e solar b e a m a n d a n o r m a l v e c t o r t o t h e radians
tree surface
5 solar d e c l i n a t i o n angle: (--) w i n t e r , (+) s u m m e r radians
A b o u n d a r y layer t h i c k n e s s m
c solar e l e v a t i o n angle radians
l a t i t u d e : (+) N o r t h , (--) S o u t h (-~ + 0 . 4 9 r a d ( 2 8 ° N ) at radians
our latitude)
e l e v a t i o n angle in t h e tree c o o r d i n a t e s y s t e m radians
0 a z i m u t h angle in t h e tree c o o r d i n a t e s y s t e m radians
P r a d i u s o f spherical tree m
T zenith atmospheric transmissivity
4l 4

direction is zero, i.e. :

V2T = 0 (2)
where 2 is the Laplacian operator. The solution to eq. 1 can be written as:
--/°(-1--a)fe0s~--" R'I V (2n + 1)Pn(O)Pn(cOsfl) I' r ~"
Ar(r, f l ) - 2(ks+hR) -o , , . - - a ) 7-0 2(n+2i(n--1}(nks+hR)~R-!
(3)
after Thorpe (1974) and Smart and Sinclair (1976). Pn(O)and Pn(cos fl) are
Legendre polynomials of degree n, and n takes only even values in the sum-
mation, r is the distance from the center of the fruit, and fl is the polar angle
between a point and the solar beam.
By assuming a linear temperature gradient along the solar beam, Smart and
Sinclair (1976) were able to use the energy balance equation (1) to derive
simplified expressions for the increase above air temperature at the center,
hotspot and backside of a spherical fruit. These expressions are:
Tcen = I 0 (1 -- a)/4h (4)
Tmax=Io(1 a)(ks + 4hR)/{(Rh + ks)4h} (5)
Tmin = I0(1 a)ks/{(Rh + ks)4h} (6)
The heat transfer coefficient, h, is calculated from:
h = ka/A + 2ka/D (7)
(Nobel, 1975; Smart and Sinclair, 1976) and the boundary layer thickness,
A, (in meters) adapted from Nobel's eq. 4 is obtained from:
D
A : 2-8~.87~vD}~:~ (8)
where ka is the thermal conductivity of air (0.0255 Wm -1 °C-1), D is the
diameter of the sphere (m) and v is the wind velocity in m sec -~ .
We have found t h a t computer calculations of Tmax using eq. 5 and the
exact solution (eq. 3) for 6 cm diameter oranges involved disagreements of
less than 1%. Smart and Sinclair (1976) also f o u n d that eqs. 5 and 6 gave
values very close t o eq. 3. For this reason, we have used eqs. 5 and 6 to
describe the range of temperatures on the surface o f citrus fruit as a function
of I0, the solar beam intensity.

Average energy flux in a tree quadrant

The intensity o f the solar beam, I0, in eqs. 4, 5 and 6 is dependent on

several variables. Some of these, such as latitude, time of year, time of day
and position in the tree can be easily predicted or measured with considerable
accuracy. Others such as cloudiness and wind velocity may have predictable
 415

S. N

S?U~H/AAf
~ P /~/ NORTH

SOUT~ EAST
BOTTOM
Fig.1. Spherical s h a p e used t o r e p r e s e n t a citrus tree. P o r t i o n m o r e t h a n 35 ° b e l o w c e n t e r
is n o t used. H e a v y lines i n d i c a t e q u a d r a n t d e s i g n a t i o n s , ~ a n d ~ are a z i m u t h a n d e l e v a t i o n
angles at the c e n t e r of t h e area e l e m e n t AA.

effects but be entirely whimsical in time and space.

As a starting point for an energy flux model, some sort of citrus tree
geometry must be assumed. We have used two different shapes to represent
citrus trees: (1) spherical and (2) icosahedron (virus particle-like polygon
having t w e n t y equilateral triangular faces). Both of these shapes gave virtually
the same results with the sphere having some computational advantages. For
this reason we utilize the spherical model t h r o u g h o u t as represented in Fig. 1.
The complete sphere is n o t used, only that portion down to a plane 35 ° below
the center since this seemed to approximate the observed profile of most
citrus trees. The " s o u t h t o p " quadrant is bounded on the north by the east--
west plane and on the b o t t o m by a plane 10 ° above center; the " s o u t h
b o t t o m " quadrant is bounded on the north by the east--west plane and on
the top and b o t t o m by the +10 ° and --35 ° planes, respectively. The north top
and b o t t o m quadrants are analogously defined.
The total direct beam solar energy flow rate through any area, A, of the
spherical tree is given by the integral of the dot product between the solar and
normal vectors taken over the area, i.e.:

I0 A = f f S " N dA (9)
A

This integral is difficult to evaluate analytically because of complicated

limits involving both the time varying solar terminator and the fixed quadrant
416

boundaries. We have, therefore, chosen to use the approximation:

IOA -~ ~ S " Ni~Ai
A
or:
Io A ~ ISI IN I cos TiAAi (10)
A
where ~ A i is a (small) finite area element and 7i is the angle between the
solar and normal vectors at the " c e n t e r " of this element. Computer evaluation
of eq. 10 is relatively easy since cos 7 becomes negative for 7 > 90°, and: these
negative parts can be excluded from the summation thus avoiding the shaded
part of the tree.
In order to evaluate eq. 10 we must first know 7, the angle between the
solar vector, S, and the normal vector, N, at any point on the spherical tree
surface (Fig.l). If the finite area element A A i is b o u n d e d by the spherical
coordinate angles Oi, 0_i+1 and ¢i, ¢i+1 and has average coordinates_ _
-Oi = (0i + 0i+1)/2 and ¢i = (¢i + ~i+1)/2, then the normal vector at 0i, ¢i can
be written as:
Ni ~ -"
~ l = i cos ~i sin 0i +J cos ¢i cos 0i + k sin ¢i (11)

where, for the moment, J/VI will be taken as 1. In the same coordinate system
(Fig.l), the solar vector can be written as:

S icosesina+ cosecosa+ sine (12)

Isl
where, 181 will also be taken as 1 and a and e are the solar azimuth (measured
clockwise from north) and elevation angles. If we momentarily shorten the
notation of 11 and 12 to:
N=A1i+A +A a (13)
and:

then since INI = IS] = 1, cos 71can be written as:

cos T = AIB1 + A2B2 + A3B3 (15)
The solar elevation angle, e, can be calculated from the well-known
formula:
sin e = sin k sin 5 + cos k cos 5 cos H (16)
(Gates, 1962, pp. 58--59; Stapleton et al., 1973; Schultze, 1976). The solar
hour angle, H, measured as time from solar noon in radians is given by:
H = (12 -- t)Tr/12 (17)
 417

where t is hour on a 24-h cycle. Corrections for non-central time zone location
and elliptical earth orbit are available in Stapleton et al. (1973). The declina-
tion angle, 5, may be found in Solar Ephemeris Tables or, more simply,
calculated from:
5 = --0.40773 cos [(11.36 + day) 2n/365] (18)
where day is Julian day number (A. Green, personal communication, 1978).
The solar azimuth angle, ~, can be calculated from:
sin ~ = cos 5 sin H/cos e (19)
(Gates, 1962; Stapleton et al., 1973). In this expression, we have removed a
negative sign from in front of Stapleton's formula (A33, p. 40) to put the
azimuth angle in our coordinate system (measured clockwise from north,
e.g., 0 in Fig.l). In addition, Gates' formula (eq. 2.13, p. 59) in terms of
zenith angles should be divided by sine rather than cosine of zenith angle.
Eqs. 11--19 allow us to calculate cos 3'i in eq. 10. The spherical area
element, AAi, in eq. 10 may be calculated from:
-Oi+ 1 cPi+l

AAi=P2 jl re) cos~)d@dO

i i

or

AAi = p2 (Oi+' _ Oi) (sin @i+1--sin ~i) (20)

Using eqs. 11, 12 and 20, eq. 10 becomes:
IOA ~-- ISI [NIp2 ~ {(cos ~i sin0i cos e sin a + c o s T / c o s 0i cos e cos a +
A

sin ~i sin e) (Oi+, -- Oi) (sin ~i+, -- sin ¢i)} (21)

where the summation is to be taken over a "quadrant" of the tree and
negative values (shaded portions) are excluded.
Since assumingthat p2 = 1 is equivalent to assuming a spherical tree of 1-m
radius, IOA must be divided by the area of the quadrant to obtain average
incident energy flux per quadrant. The two top quadrants have an area of:

ATO P = p2 ( ~ - - 2 ) {sin(90°)-- sin(10°)}

p2 2.596 m2 (22)
and the b o t t o m quadrants have an area of:

ABOT = p2 ( ~ _ _ ~ ) {sin(10o)_ sin(_35o)}

= p2 2.347 m2 (23)
from eq. 20. The sum in eq. 21 must be divided by eq. 22 or 23 for top and
418

b o t t o m quadrants respectively. In addition the magnitude of the solar vector

is given approximately by:
I SI = So rcsce (24)

(Allen, 1974; Schulze, 1976). So is the solar constant, 1360 W m -2, (Monteith,
1973), v is the zenith path atmospheric transmissivity (about 0.8 in central
Florida) and e is the solar elevation angle given by eq. 16.
From eqs, 22, 23 and 24, eq. 21 can be written as:
IOQuAD ~ p2 (SoTCSCe/AQUAD) ~ {(cos ~i sin 0i cos e sin a +
QUAD

COS ~i COS 0i COS e COS O~+ sin ~i sin e) (0i+1- - Oi) (sin ¢i+, -- sin ¢i)} (25)
w h e r e IOQuAD is the average solar energy flux in W m -2 incident upon a
particular tree quadrant, AOUAD is the area of the quadrant from eq. 22 or
23 and the summation is to be taken over the sunlit part of the quadrant in
question. In all our computer calculations, (Oi+l -- Oi) and (¢i+1 -- ¢i) = 5° ~-
0.087266 rad.
To simulate average fruit surface temperatures in a tree quadrant,/0QUA D
is substituted for I0, the solar beam intensity, in eqs. 4, 5 or 6.

COMPARISON OF THE MODEL WITH FIELD DATA

Three exposed Valencia oranges in each tree quadrant were fitted with
thermistor temperature probes wired to a twelve-point recorder. The probes
were held tightly against the outside surface with rubber bands and a drop of
heat sink c o m p o u n d to insure good thermal contact. The three fruit in each
quadrant were positioned " a r o u n d " the quadrant dividing it into three equal
parts. This was done to obtain a more representative average " h o t s p o t "
temperature for the quadrant than would have resulted from random spatial
selection. Air temperatures were measured with a hygrothermograph in a
standard weather shelter in the same grove.
Two days of particular interest are shown graphically in Fig.2. The average
fruit temperatures by quadrant for September 28, 1977 are shown in the four
right hand panels of Fig.2. The main purpose here is to demonstrate the effect
of clouds on fruit temperatures. This particular day had clear sky followed by
cloud cover followed by clear sky again, and the depressing effect of clouds
on fruit temperatures is quite obvious. Cloud cover will reduce direct solar
intensity by about 80%, (Brooks, 1964; Schulze, 1976); i.e., eq. 22 would be
multiplied by 0.20. This fact and the data in Fig.2 suggest that cloud cover is
a major factor in the thermal environment of the rust mite on exposed citrus
fruit.
The effects of solar radiation on fruit temperatures on a clear day, September
30, 1977, are shown in the four left-hand panels of Fig.2. The south top
quadrant has average hotspot temperatures 10°C above air temperature, while
 419
CLEAR (9-30-77) CLOUDY (9-28-77)
--Model • Fruit * Air
[ z.lll& SOUTH SOUTH
TOP
,,%
30

20
SOUTH SOUTH
BOTTOM

A
30
o

20 i , , I I I I I |

^I NORTH
It TOP
NORTH
uF TOP

L~J
I-- 20 j I I i I I

nL NORTH I NORTH
vt BOTTOM •~ BOTTOM

20 ~
6 12 18 24 6 6 12 18 24 6
SOLAR TIME

Fig.2. Temperature cycles on fruit (u) compared with air temperature (+) for cloudy and
clear days. The solid line represents simulated temperature using eqs. 5 and 25.

the n o r t h b o t t o m s h o w s o n l y a 1 - - 2 ° C increase. S i m u l a t i o n of t h e s e
t e m p e r a t u r e s requil'es w i n d v e l o c i t y to calculate the h e a t t r a n s f e r c o e f f i c i e n t ,
h, f r o m eqs. 7 a n d 8. Wind v e l o c i t y o n t h a t d a y was a p p r o x i m a t e l y 3 m sec -1
at 18.3 m as m e a s u r e d b y t h e N O A A N a t i o n a l W e a t h e r Service at L a k e l a n d ,
Florida. A v e l o c i t y o f 3 m sec -1 at 18.3 m will be r e d u c e d b y a p p r o x i m a t e l y
2 / 3 at t h e c a n o p y t o p (Albrigo, 1976}. In a d d i t i o n , a n o t h e r 50% r e d u c t i o n
w o u l d s e e m t o be in o r d e r in t h e b o t t o m h a l f o f t h e tree ( V a n E i m e r n et al.,
1 9 6 4 ) . T h u s w i n d v e l o c i t y was a s s u m e d to be 1 m sec -1 in t h e t o p half o f the
t r e e a n d 0.5 m sec -1 in t h e b o t t o m half.
Eq. 25 was u s e d t o c a l c u l a t e IOQUAD, t h e average solar e n e r g y f l u x f o r e a c h
t r e e q u a d r a n t , at 15 m i n intervals. This e n e r g y was t h e n u s e d as I 0 in eq. 5 to
o b t a i n t h e e l e v a t i o n o f average h o t s p o t t e m p e r a t u r e o v e r air t e m p e r a t u r e . T h e
420

temperature increase from eq. 5 was then added to observed air temperature
to produce the solid lines in Fig.2. In spite of a rough wind velocity estimate
and a sample size of only three fruit per quadrant, the model agrees remarkably
well with the observed data,

RUST MITE OUTBREAKS IN TREE QUADRANTS

Two studies were done in which rust mite densities on exposed fruit
(Valencia oranges) were followed during unsprayed outbreaks in different
tree quadrants. In this section we will compare the behavior of these popula-
tions with simulated fruit temperatures. The actual fruit temperature during
these outbreaks cannot be reproduced because of the absence of solar radiation
data, but fruit temperatures can be simulated for different times of year and
different wind velocities.

Simulation of fruit temperature

Typical temperature regimes on fruit for our latitude (28°N) are shown for
the summer and winter solstices in Fig.3 and for the vernal and autumnal
equinoxes in Fig.4. Air temperature was assumed to be sinusoidal (5.5 °
amplitude) around a 35-yr average for the day with a peak at 15h00. Tmax
or Train of eq. 5 or 6 was added t o this ambient temperature cycle to produce
the upper and lower limits of temperature on the fruit as indicated by the
pairs of lines in Figs.3 and 4. All fruit temperatures must lie between these
lines for the given set of conditions. Wind velocities of I and 4 m sec -1 (top
quadrants) and 0.5 and 2 m sec -1 ( b o t t o m quadrants) are illustrated.

Rust mite population dynamics

Rust mite population densities by tree quadrant in two different groves are
shown in Fig.5. Waverly grove has a wide tree spacing (6.1 x 9.1 m) with small
trees having considerable open space around t h e m and little shading of one
tree by another. Bay Lake grove is a hedgerow spacing (4.6 x 7.6 m) of large
overlapping trees with rows running east--west which would be expected to
accentuate any north--south differences. Both groves were 4.05 ha blocks, and
eight fruit were sampled f r o m each quadrant of four randomly selected trees
on each sample date.
Both groves had a summer mite o u t b r e a k with the b o t t o m of the tree
showing the highest mite densities (Fig.5). For unknown reasons, Bay Lake
grove had a m u c h higher popuiation on the north b o t t o m at one c o u n t than
occurred in any other sample in either ~ o v e . This may simply b e d u e to our
random sample including a few unusually high density trees. It cannot be
easily explained on the basis of temperature differences. North and south
b o t t o m show nearly identical temperature regimes on June 21 (Fig.3), and it
seems unlikely than an east--west hedgerow would drastically alter this
 421

JUNE2[ DECEMBER21
SOUTH TOP f SOUTH TOP
--I msec -L 35 ~1 msec -I
35
.... -- I ...... 4 m s ec - I

2G

t5
i L h i L L L i i

SOUTH BOTTOM I" SOUTH BOTTOM

35 --05 m sec t 35I --0.5 m see"~
..... C - I ...... 2 m $ec -t

25
~ e 25

15
i A i i i i "~- i i i

NORTHTOP NORTH TOP

--I mse¢ "l ~ 1 msec -I
~ 35 ..... -I
35
..... 4 m sec -I

25

co
15 15
i i ~ i h h

NORTHBOTTOM NORTH BOTTOM

--0.5 m sec "~ 35 ~ 0 . 5 lit SeC"1
35 ..... -i ...... 2 m sec-I

2G

15 15
i i i

4 B ,2 ,~ 2'0 2'4 ,~ ,~ ,'~ 2'0 2'4

SOLAR TIME

Fig.3. S i m u l a t e d t e m p e r a t u r e s o n fruit f o r t h e s u m m e r a n d w i n t e r solstices in c e n t r a l

F l o r i d a . The t w o solid lines i n d i c a t e t h e m a x i m u m a n d m i n i m u m t e m p e r a t u r e s for
1 m sec- 1 w i n d ( t o p q u a d r a n t s ) a n d 0.5 m sec- 1 w i n d ( b o t t o m q u a d r a n t s ) . T h e t w o d o t t e d
lines i n d i c a t e t h e r a n g e f o r 4 m sec -1 a n d 2 m sec -1 w i n d t o p a n d b o t t o m , respectively.

situation. (In fact, the temperatures reported in Fig.2 are from the Bay Lake
grove and fit the model quite well.)
Hobza and Jeppson (1974) reported an o p t i m u m temperature for citrus
rust mite at 24.5°C. Reed et al. (1964) f o u n d that temperatures of 32.2 ° _
37.8°C were lethal to citrus rust mite in 2--3 h. Therefore it is not surprising
t h a t the n o r t h and south top quadrants with an average m a x i m u m temperature
range between 33 ° and 38°C (1 m sec -1 wind) show very low rust mite
population densities. Even 4 m sec-1 winds do not reduce average tem-
peratures out of the lethal range in the top of the tree on June 21 (Fig.3).
422

MARCH21 SEPTEMBER 2J
SOUTHTOP SOUTH TO,
35 - - I m SeC-I 3~i / \ --,m,ec"

25

IS
i i i i

F SOUTHBOTTOM [ SOUTH BOTTOM

35} ~ O . 5 m sec -~ 35~ ~ --0.5 mSiC "t
°~*'}
~ 2 , 5 2m
25I/
sec -I
t ~ 2" .'c''

15

'~ i i L J i i

f NORTH TOP NORTH TOP

~_~ 35 --I m sec "i 35 ~ 1 m SgC"1
4 m SeC "1 .... ¢-I

25

IS
"~ i i A i i A

NORTH BOTTOM I- NORTH BOTTOM

35 - - 0 5 m sec -I 35 L --O.Sm sec -j
.... 2 m see-' | ~ 2 m.c-t

25

t5
"r
,'2 ,'~ 2'0 2', ~ "4 8. . 12. . 16. . 20. 24 4
SOLAR TIME

Fig.4. Same as Fig.3 for the vernal and autumnal equinoxes.

Waverly grove exhibited a winter outbreak during the warm winter of

1 9 7 4 / 1 9 7 5 which is of some interest (Fig.5). The quadrant differences in
winter tend to diminish somewhat; however, the north bottom is highest,
and the south top is lowest in rust mites as one might e x p e c t if overheating is
involved. The average curve in Fig,3 f o r December 21 is not into the lethal
zone for the south top. The average curve can be a bit misleading, however,
since only an occasional day o f lethal temperatures would be required to
reduce population levels. Even though the temperatures for November 1974
to January 19,75 are only slightlyabove average, there were 41 days in these
months with the maximum air temperature above 26°C and 6 days in
November having a maximum of at least 30°C. This puts the fruit temperature
 423

WAVERLY GROVE

50

==40 : ; NORTH TOP

*-----,k NORTH BOTTOM S"
g ----,L SOUTH TOP .~.
o.---.o SOUTH BOTTOM
~3o

~20 It -
// tit "

I0
• ~ .' ' ~ : ~
~ H A R V E S T . ~ ~
", .',.'a
I t " I i ; , t -I
NOV DEC JANIFEBIMARIAPRIMAYIJUNEd, ILYiAUG SEPT OCT NOV DEC JAN
1973 ! 1974 11975

BAY LAKE

• • NORTH TOP
160 •m.--- D NORTH BOTTOM
140 ' ~ ' - - " SOUTH TOP !i
° " - - " ° SOUTH BOTTOM !i
120

== ioo li
LL. l i
=
o
BO
i I
"~ so i L
o~ 40 ! !
I ,e- ,-R
• 20 _ ~ " ] k ~ . i "'~,
- . H,LRVEST " * g ~ V ' - " " ~ ' . ~ - " - ' ~ , , , ~
OCT' NOV'DEC' JAN'FEB'MAR'APR'MAY'JUN'JUL'AUG'SEP'OCT' NOV'DEC'JAN'
1974 1975

Fig.5. Rust mite population dynamics in different quadrants in two different Valencia
orange groves in central Florida.

range for Fig.3 well into t h e lethal z o n e several times and this w o u l d be
e n o u g h t o r e d u c e rust mites in the s o u t h t o p o f the tree.

SUMMARY AND CONCLUSIONS

A m o d e l o f citrus f r u i t t e m p e r a t u r e s was c o n s t r u c t e d based o n average solar

e n e r g y flux in tree q u a d r a n t s . This m o d e l is a f u n c t i o n o f air t e m p e r a t u r e ,
424
latitude, time of year, time of day, fruit diameter and reflectance, position in
the tree, atmospheric transmissivity and wind velocity. Diffuse radiation,
radiation heat loss, latent heat loss and wind direction are ignored. The model
was f o u n d to agree remarkably well with t em perat ure data from the field and
is useful in simulating a variety of conditions affecting fruit temperatures.
When the model's predicted temperatures were com pared with rust mite
distribution patterns in the tree~ the rust mite populations were low where
excessively high t em pe r at ur e s were predicted. It seems that the general popu-
lation distribution in the tree can be predicted on the basis of t em perat ure
alone. It is true however, t h a t high h u m i d i t y favors the devel opm ent of the
rust mite's immature stages (Pratt, 1957; H o bza and Jeppson, 1974). F u r t h e r
study is needed to explain the aggregation of rust mites around sunlit areas
on individual fruit, perhaps by using the model developed here to calculate
water loss rates over the fruit surface. These water flux rates will be strongly
affected by temperature, and t h e y will, in turn, affect the b o u n d a r y layer
vapor pressure in which the rust mites live.

ACKNOWLEDGEMENTS

The mathematical advice of Dr. James H. St am per was greatly appreciated.

This study was s u p p o r t e d by CSRS Grants PL 89--106 and 701--15--58 and
also by NSF Grant G B - - 34718 to the University of California and the
University of Florida.

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