Population Dynamics – Practical 2 Report

Divya Krishnamohan
Student ID: 200292988

K- Factor Analysis:
The holly-leaf miner
(Phytomyza ilicis)

Practical 2 Report

POPULATION DYNAMICS
BLGY 5101

(MSc. Biodiverity & Conservation)

Divya Krishnamohan
Student ID: 200292988
 Population Dynamics – Practical 2 Report
Divya Krishnamohan
Student ID: 200292988
K- Factor Analysis: The holly-leaf miner (Phytomyza ilicis)

Introduction:

The aim of this study is to determine the major causes of mortality in the holly-leaf miner
(Phytomyza ilicis) in the area of Leeds, by performing a k-factor analysis on data obtained by
dissecting local holly leaves.

The k-factor analysis is used to identify which of several mortality factors contribute most
significantly towards determining the population size of P. ilicis, as well as whether the mortality
factors involved are density dependent.

Life Cycle of P. ilicis

The holly-leaf miner is an agromyzid fly. During June, the female fly lays a single egg on the
base of the midrib on the underside of the leaves of the holly tree (Ilex aquifolium). When the
larva hatches, it slowly eats its way through the central tissue and eventually into the outer
parenchyma, where it remains beneath the epidermis, feeding on the parenchymatous tissue. The
‘mine’ so formed appears irregularly shaped and pale in colour, attaining its maximum size in the
month of March. Before pupating, the larva prepares a thin triangular area on the leaf cuticle
against which it fits a hinged emergence plate. The larva then forms a puparium which lies
pressed against the epidermis, with its anterior spiracles projecting through the attenuated area of
the cuticle. The adult form emerges from the puparium by breaking through the hinged plate. The
emergence of the adult is therefore identifiable on the leaf surface by a characteristic raised
triangular flap.

During its larval phase, P. ilicis is subject to various threats to its survival. Several
hymenopterous wasps parasitise either the larva or the pupa of P. ilicis and can be identified by
the presence of a pupa characteristic of the parasitic species, or the presence of an emergence
hole (usually a neat, round hole) that differs from that made by adult P. ilicis.

Another significant cause of mortality (especially during the last instar) is bird predation. The
blue tit (Parus caeruleus), in particular, is adept at ripping open mines and feeding on the larvae.
A mine surface that is torn is indicative of bird predation.

Further, an unknown number of diseases are seen to affect the holly-leaf miner larvae and pupae.
An intact mine or puparium containing no intact insect usually indicates that the individual has
succumbed to some disease.

In the following study, leaf mines present on leaves from 9 different holly trees were dissected
and the fate of the insect determined. The number of survivors (successful emergents), and
number of deaths attributed to parasitic wasps Chrysocharis gemma, Chrysocharis syma and
Sphegigaster flavicornis; bird predation; larval disease; and pupal disease were recorded. The
data were then analysed to identify the key factors of mortality influencing the population of P.
ilicis under study.
 Population Dynamics – Practical 2 Report
Divya Krishnamohan
Student ID: 200292988

Method:
The method followed was as specified in the practical handout, Practical 2. K-factor Analysis:
The holly-leaf miner (Phytomyza ilicis) by Dr. K. C. Hamer.

The methodology outlined was followed with the exception of one deviation – data collected by
Group- Tree 1 was transformed for the purpose of performing the necessary calculations of the
analysis, i.e., as no survivors were found, calculating the mortality factor k5 was not possible; to
resolve this, a value of +1 was added to the cumulative data collected by Group- Tree 1, achieved
by assuming that 1 survivor was found.
 Population Dynamics – Practical 2 Report
Divya Krishnamohan
Student ID: 200292988

Results:
Table 1.

Raw data, cumulative numbers and k values for data collected by Group- Tree1.

Life stage Numbers Cumulative Log numbers k value

(cause of (raw data) numbers (a0-6) (of a0-6)
mortality)
C. gemma 3 21 1.322 0.067
Larval disease 11 18 1.255 0.41
S. flavicornis 0 7 0.845 0.00
C. syma 0 7 0.845 0.00
Pupal disease 1 7 0.845 0.00
Bird predation 5 6 0.778 0.067
Survivors 1 1 0 0.778
Total 21 K=1.322

Table 2.

Cumulative data and k values of trees 1-9.

C. gemma Larval S. flavicornis C. syma Pupal Bird

disease disease predation
Tree a0 k0 a1 k1 a2 k2 a3 k3 a4 k4 a5 k5 K
1 21 0.06 18 0.41 7 0.00 7 0.00 7 0.06 6 0.77 1.322
2 26 7 22 0 9 0 9 0 3 7 2 8 1.415
3 23 0.07 21 0.38 9 0.00 8 0.47 8 0.17 8 0.30 0.760
4 27 3 27 8 9 0 9 7 9 6 5 1 0.829
5 18 0.04 18 0.36 15 0.05 13 0.00 13 0.00 9 0.30 0.477
6 15 0 15 8 8 1 7 0 7 0 4 1 1.176
7 16 0.00 14 0.47 8 0.00 7 0.00 7 0.25 4 0.09 0.903
8 25 0 24 7 11 0 11 0 11 5 10 7 1.398
9 22 0.00 22 0.07 8 0.06 8 0.00 8 0.16 5 0.17 1.342
0 9 2 0 0 6
0.00 0.27 0.05 0.00 0.24 0.60
 Population Dynamics – Practical 2 Report
Divya Krishnamohan
Student ID: 200292988
0 3 8 0 3 2
0.05 0.24 0.05 0.00 0.24 0.30
8 3 8 0 3 1
0.01 0.33 0.00 0.00 0.04 1.00
8 9 0 0 1 0
0.00 0.43 0.00 0.00 0.20 0.69
0 9 0 0 4 9

Total 19 0.02 18 0.33 84 0.02 79 0.03 73 0.14 53 0.42 0.985

3 8 1 3 7 4 3

1.50

1.25
Total life-time mortality (K)

1.00

0.75

0.50

0.00 0.02 0.04 0.06 0.08

Mortality caused by C. gemma (k0)

1.50

(K)
1.25
ality
Mort
time
1.00
Life-
Total
0.75

0.50

0.10 0.20 0.30 0.40 0.50

Mortality caused by Larval Disease (k1)
 Population Dynamics – Practical 2 Report
Divya Krishnamohan
Student ID: 200292988

1.50

1.25
Total life-time mortality (K)
1.00

0.75

0.50

0.00 0.01 0.02 0.03 0.04 0.05 0.06 0.07

Mortality caused by S. flavicornis (k2)

1.50

1.25
Total life-time mortality (K)

1.00

0.75

0.50

0.00 0.20 0.40

Mortality caused by C. syma (k3)
 Population Dynamics – Practical 2 Report
Divya Krishnamohan
Student ID: 200292988

1.50

1.25
Total life-time mortality (K)
1.00

0.75

0.50

0.00 0.05 0.10 0.15 0.20 0.25 0.30

Mortality caused by Pupal Disease (k4)

1.50

1.25
Total life-time mortality (K)

1.00

0.75

0.50

0.00 0.20 0.40 0.60 0.80 1.00

Mortality caused by Bird Predation (k5)

Fig.1. Relationship between the mortality factors (k0-5) and total life-time mortality (K) to
determine which mortality factors are key factors.
 Population Dynamics – Practical 2 Report
Divya Krishnamohan
Student ID: 200292988

0.08

0.06

Mortality caused by C. gemma (k0)

0.04

0.02

0.00

16.00 18.00 20.00 22.00 24.00 26.00 28.00

Cumulative population (a0)

0.50

Disease 0.40
(k1)by Larval
Mortality caused

0.30

0.20

0.10

15.00 17.50 20.00 22.50 25.00 27.50

Cumulative population (a1)
 Population Dynamics – Practical 2 Report
Divya Krishnamohan
Student ID: 200292988

0.07

0.06

0.05
(k2)by S. flavicornis
Mortality caused
0.04

0.03

0.02

0.01

0.00

7.50 10.00 12.50 15.00

Cumulative population (a2)

0.40

Mortality caused by C. syma (k3)

0.20

0.00

7.00 8.00 9.00 10.00 11.00 12.00 13.00

Cumulative population (a3)
 Population Dynamics – Practical 2 Report
Divya Krishnamohan
Student ID: 200292988

0.30

0.25

0.20 (k4)
Mortality caused by Pupal Disease
0.15

0.10

0.05

0.00

2.500 5.000 7.500 10.000 12.500

Cumulative population (a4)

1.00

0.80
Mortality caused by Bird
Predation (k5)
0.60

0.40

0.20

0.00

2.000 4.000 6.000 8.000 10.000

Cumulative population (a5)

Fig. 2. Relationship between cumulative populations (a0-5) and mortality factors (k0-5) to
determine density dependence of mortality factors.
Table 3.

Kendall’s tau_b correlations between total life-time mortality (K) and mortality factors
(k0-5) to determine which mortality factors are key factors.

K correlated with Kendall's tau_b Sig.

k0 0.365 0.189
k1 0.278 0.297
k2 -0.567 0.049
k3 0.471 0.121
k4 -0.141 0.600
k5 0.551 0.043

Table 4.
 Population Dynamics – Practical 2 Report
Divya Krishnamohan
Student ID: 200292988
Kendall’s tau_b correlations between mortality factors (k0-5) and their initial densities or
cumulative populations (a0-5) to determine which mortality factors are density dependent.

ax correlated with kx Kendall's tau_b Sig.

a0, k0 0.122 0.661
a1, k1 0.514 0.058
a2, k2 0.110 0.718
a3, k3 0.191 0.551
a4, k4 -0.120 0.667
a5, k5 0.209 0.452

key factor not a key factor

density
dependent -- Larval disease

density Bird predation C. syma

independent S. flavicornis C. gemma
Pupal disease

Fig. 3. Classification of mortality factors affecting the population of P. ilicis in Leeds.

 Population Dynamics – Practical 2 Report
Divya Krishnamohan
Student ID: 200292988

Discussion and Conclusion:

Of a total of 193 holly-leaf mines examined, 10.36% of the cases were identified as having adult
P. ilicis emerge successfully. It is evident that the population of holly-leaf miners under study are
subject to a very high mortality rate.

The k-analysis revealed that there were two key mortality factors – bird predation, with a
positive correlation value of 0.551 (P=0.043) and S. flavicornis, with a negative correlation
value of -0.567 (P=0.049). (Refer Fig. 1, and Table 3).

The incidence of bird predation was recorded in 17.1% of the cases; however the ‘killing power’
of this mortality factor was seen to be 62.26% (almost 10% more than the next most significant
mortality factor, larval disease). A positive, significant correlation indicates that an increased
incidence of bird predation would significantly influence the total life-time mortality, causing it,
in effect, to rise.

In contrast, S. flavicornis was recorded in 2.59% of the cases, having a ‘killing power’ of 5.95%.
A negative but significant correlation could indicate that a lower occurrence of S. flavicornis
could result in a higher total life-time mortality value. An interpretation of this result could be
that given that the causes of mortality occur in a time series, i.e., each successive mortality factor
acts on the proportion of survivors that have escaped the influence of the preceding mortality
factor, a higher percentage of cases with S. flavicornis infections means that there are fewer
individuals that are subject to mortality caused by a factor further down the time series (such as
bird predation) which has a killing power with an influence 10 fold that of S. flavicornis.

Correlations of the different mortality factors with their initial densities revealed that only one
factor was density dependent (marginally) – larval disease, 0.514 (P=0.58). Logically, one can
conclude that an increased density of P. ilicis results in a higher proportion of individuals that get
infected with larval disease as higher densities favour rapid spread of contagion.

Of the four possible types of mortality that can act on the holly-leaf miner (refer to Fig.3),
P. ilicis in Leeds is affected by three types – density independent key factors, and non key factors
that are both density independent and dependent. Larval disease is the one factor that is density
dependent and though it is a not a key factor it may be noted that it was recorded in 50.25% of
the cases.
 Population Dynamics – Practical 2 Report
Divya Krishnamohan
Student ID: 200292988
Recommendations:
In the event that Phtyomyza ilicis is considered a pest species:

Studies have shown that a good bio-control agent should impose spatially density-dependent
mortality on the victim population (Beddington et. al 1978). In this study however, larval disease
is seen to be the only density-dependent factor, albeit, a non-key factor. Larval disease therefore,
cannot be used as a bio-control agent as the analysis reveals that it does not significantly
influence life time mortality, added to which the specific disease is unknown (it could be one of
several that afflict P. ilicis).

Of the factors analysed, bird predation is seen to contribute most significantly towards the life
time mortality rate of P. ilicis. The blue tit, (Parus caeruleus) is a well known predator of the
holly-leaf miner. One may recommend encouraging bird predators such as the blue tit in areas
targeted for reducing P. ilicis by erecting nest boxes (rather than placing bird feeders). Blue tits
are aggregate feeders and providing nesting sites beside a potential food source such as an Ilex
stand (having leaf mines) is likely to have the desired effect of reducing P. ilicis numbers.

Elsewhere, the use of the parasitoid wasp Chrysocharis gemma has proven successful (Clausen
1978). Other studies have indicated that C. gemma is a density dependent key factor. A possible
explanation for the discrepancy in results is the scale of this study and perhaps a prevalence of
habitat variables that do not encourage C. gemma in the locality from which samples were
obtained.

In the event that Phytomyza ilicis is considered an important and threatened species:

In order to conserve the holly-leaf miner, it would be imperative to reduce the factors that
threaten its survival. From this study it is apparent that bird predation poses a significant risk,
and measures to reduce the incidence of birds feeding on P. ilicis, such as providing alternative
food sources, or discouraging aggregations, may reduce larval mortality caused by this source.

Further, narrow spectrum insecticides, or even the use of pheromone baiting, may be employed
to target the parasitoid wasp species (assuming they exist).

Research on the diseases that affect P. ilicis may provide valuable information on how to control
and treat larval and pupal diseases.

Increasing plantations of the food plant Ilex aquifolim may help in boosting P. ilicis populations,
though little is known about how host selection takes place or what specific characteristics of the
host plant are desirable.
 Population Dynamics – Practical 2 Report
Divya Krishnamohan
Student ID: 200292988

References:
Books and journals:

Heads, P. A., & Lawton, J. H. (1983). Studies on the natural enemy complex of the holly leaf-
miner: the effects of scale on the detection of aggregative responses and the implications
for biological control. Oikos, 40, 267-276.

Lewis, T., & Taylor, L.R. (1967). Introduction to Experimental Ecology. London: Academic
Press.

Valladares, G., & Lawton, J. H. (1991). Host-Plant Selection in the Holly Leaf-Miner: Does
Mother Know Best? The Journal of Animal Ecology, 60, 227-240.

Websites:

Flint, M. H., & Doane, C.C. (1996). Understanding semiochemicals with emphasis on insect
sex pheromones in integrated pest management programs. Retrieved November 21, 2006
from the University of Minnesota, Radcliffe’s IPM World Textbook website:
http://ipmworld.umn.edu/chapters/flint.htm