Tropical Fruit - Chap 11 7/5/03 1:05 PM Page 253

11 Diseases of Fig

Themis J. Michailides
University of California, Kearney Agricultural Center, Parlier, California, USA

Introduction which serves as a pollenizer, and the pistil-

late edible fig. Pollination is achieved by the
The common edible fig, Ficus carica, probably female fig wasp, Blastophaga psenes (Fig. 11.2),
originated in southern Arabia since the which develops in the syconium of the capri-
caprifig, ancestor of the edible fig, is still fig. The symbiotic relationship between fig
found there growing wild (Solms-Laubach, and fig wasps is an example of co-evolution
1885). The species name refers to Caria, an between plant and insect (Galil and Neeman,
ancient region of Asia Minor noted for its 1977; Beck and Lord, 1988a,b; Brostein, 1988).
figs. Fig spread through the Old World in Other related forms that are described as dis-
ancient times, and was first introduced into tinct species include F. geraniifolia, F. palmata,
the New World by Spanish and Portuguese F. serrata (the Indian form of F. carica), F.
missionaries (Cuba and Peru). pseudo-carica and several others that are used
in breeding programmes.

Origin and Taxonomy of Important

Species History

F. carica is a member of the mulberry The literature on figs is voluminous and

(Moraceae) family. It contains 60 genera and dates from ancient times (Condit and
possibly >2000 species of trees, shrubs, vines Enderud, 1956). Figs have been prized for
and herbs. Common edible figs and their both medicinal and dietary value. The early
pollinating counterpart, caprifigs, are the Greeks considered it an honour to bestow
only members of the subgenus Eusyce that either the foliage or the fruit, and winning
are cultivated for their fruit. athletes in the original Olympic games were
The edible ‘fruit’ of F. carica is unusual. It crowned with fig wreaths and given figs to
is a morphologically unique syconium that is eat. Figs were so highly valued that their
comprised almost entirely of vegetative export from ancient Greece was forbidden.
peduncular tissue. The true fruits are tiny, Athenians who informed authorities about
pedicellate drupelets, which are found those who illegally exported figs from
within the syconium (Fig. 11.1). F. carica is a Attica were named ‘sycophants’, a practice
gynodioecious species with two distinct from which the word took its modern
forms: the monoecious, non-edible caprifig, meaning (sykon in Greek means fig fruit).

© CAB International 2003. Diseases of Tropical Fruit Crops (ed. R.C. Ploetz) 253
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254 T.J. Michailides

Fig. 11.1. A fig syconium in cross-section depicting the location of female (f) and male (m) inflorescences
(diagram: T.J. Michailides).

Fig. 11.2. A female of the pollinator wasp of fig, Blastophaga psenes (diagram: T.J. Michailides).

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Diseases of Fig 255

Attributes and Present Status Caprifig trees produce three to four crops
of syconia annually (Fig. 11.3) (Brostein and
Fig fruit are enjoyed both fresh in areas Patel, 1992). The winter crop (mamme) is ini-
where they are grown, and dried throughout tiated in the autumn and matures in late
the world. They recently have been described winter, and the spring crop (profichi), initi-
as the fruit with the highest content of fibre ated in late winter, matures in mid-spring.
(Duxbury, 1986; Bamford, 1990) and minerals The profichi syconia contain the most pollen
and polyphenols (Vinson, 1999). and are used for the pollination of the edible
In 1999, world production was almost Smyrna-type figs.
1.2 Mt (FAO, 2000). With few exceptions,
the leading producers either were from the
Mediterranean region or were areas that Edible figs
had Mediterranean climates (Table 11.1).
Edible fig cultivars are divided into three
Currently, Turkey is the leading country in
horticultural categories, Smyrna, San Pedro
world fig production.
and Common, which are distinguished by
their cultural and pollination requirements.
There are >600 named cultivars, many of
Caprifigs
which are synonymous (Condit, 1955). They
are produced clonally to maintain their
Although the various crops of the fig and
desirable characters.
caprifig trees are given different names in
Smyrna-type figs are grown mostly in
different countries, the convention in this
Algeria, California, Greece, Portugal and
chapter is to refer to the edible, pistillate, cul-
Turkey, and usually require pollination in
tivated members of F. carica as ‘figs’ and the order to set fruit. Thus, they usually do not
inedible, monoecious forms that serve as produce a first, breba crop in summer when
pollinators as ‘caprifigs’. Caprifigs are essen- pollen is not available. However, the follow-
tial for pollination of the edible fruit of the ing cultivars can set a significant amount of
Smyrna-type cultivars. Their syconia contain breba crop: ‘Bardajik’ (Turkey), ‘Calimyrna’
both female and male flowers (hermaphro- (‘Lob Injir’) (California), ‘Cheker Injir’
dites). The fig wasp transfers pollen from the (Turkey), ‘Kassaba’ (Turkey), ‘Tameriout’
caprifig to pistillate flowers of the edible fig, (Algeria) and ‘Taranimt’ (Kabylia). In
a process known to fig growers as caprifica- California, the main cultivar, ‘Sari Lop’
tion. (renamed ‘Calimyrna’ = California plus
Smyrna), produces fair breba seedless crops,
but requires pollination to set a main crop.
Table 11.1. Leading producers of fig, Ficus carica,
The San Pedro-type figs initiate moderate
in 1999.a to large breba crops and an adequate main
crop (which requires pollination) on the
Country Production (t) same branch in the same season. Cultivars of
Turkey 260,000
it include ‘Banqhette’ (Morocco), ‘Dauphine’
Egypt 216,500 (France), ‘Gentile’ (England), ‘King’ and
Greece 80,000 ‘Lampeira’ (Italy, Portugal) and ‘San Pedro’
Iran 78,555 (Spain) (Condit, 1955). ‘King’, which is of
Morocco 70,000 minor importance, is the only San Pedro-
India 60,000 type fig used in the California fig industry
Spain 55,000 (Ferguson et al., 1990).
USA 46,000 Most common fig cultivars produce a
Algeria 45,000 moderate to large main crop without caprifi-
Syria 45,000
cation, and are thus parthenocarpic. However,
aSource: FAO (2000). Production figures are in climatic differences among locations can
metric tonnes. markedly alter the expression of partheno-

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256 T.J. Michailides

Fig. 11.3. A diagram of Blastophaga psenes flights and caprifig crops during a calendar year under
California conditions. The fig wasp has three major flights in late March to early April, early June, and from
August to October. Caprifig trees produce at least three major crops per year in winter (mamme), spring
(profichi) and summer (mammoni). The spring crop caprifigs are collected by growers, and hung on trees in
Calimyrna orchards (diagram: T.J. Michailides).

carpy in common figs. Common cultivars the producing country. Grade defects include
include: ‘Adriatic’ (Italy), ‘Beall’ (California), insect infestation; smut, mould and endosep-
‘Black Spanish’ (Spain), ‘Blanche’ (France), sis; sour rot; and filthy or worthless fruit.
‘Bourjassotte Grise’ (UK), ‘Brown Turkey’
(Turkey), ‘Celeste’ (Malta), ‘Dottato’ (Italy),
‘Mission’ (Spain) and ‘Troiano’ (Italy) (Condit Fig Diseases
and Swingle, 1947). Condit (1955) described
470 major cultivars in this class. Fig spoilage was mentioned in the Old
Fig fruit are marketed either dry or fresh. Testament of the Bible (de Lagarde, 1881), and
Figs for drying are allowed to ripen fully and continues to have a major impact on the pro-
partially dry on the tree. Partially dried figs duction of this fruit. Many diseases, some of
then fall to the ground where they continue to which can be quite destructive, affect the fig
dry and are later collected by hand and dried plant and fruit. Detailed information on 18 of
under the sun. Since commercial figs are har- the most significant diseases and disorders
vested every 2 weeks, disease and insect are found below. Fungi cause most of these
infestation is common due to their extended problems, but two are caused by bacteria, one
contact with the soil. Grade standards vary by by viruses, and another is physiological. Table

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Diseases of Fig 257

11.2 lists relatively minor problems of this Acervuli consist of a few layers of pseudo-
crop that could not be covered in detail. parenchymatous tissue, upon which are
borne small, parallel conidiophores. Conidia
are borne singly on the tips of the conidio-
Anthracnose phores. The pathogen is described in detail in
Chapter 1.
This disease was described in Germany and Mummified fruit are a source of conidia
Louisiana (Stevens and Hall, 1909; Edgerton, for new infections; they are dispersed in rain
1911). Both foliage and fruit are affected. and irrigation water. Removal of sources of
Leaf lesions have a dark brown margin, and inoculum, such as infected fruit fallen on the
affected leaves often turn brown, dry at the ground or hanging on the tree (Matz, 1918)
edges, and may abscise. Occasionally, spots and pruning all the dead limbs and twigs, may
and lesions can be found on leaf petioles help to reduce the disease (Edgerton, 1911).
and blades. Symptoms on fruit range from ‘De Constantine’ and ‘Celeste’ are resistant.
localized, sunken lesions to a general rot of
the entire fruit. Pink, slimy masses of coni-
dia are produced in acervuli on lesions, and Armillaria root rot
affected fruit may remain on the trees as
dried ‘mummies’. Armillaria root rot is the most important of
Anthracnose is caused by Colletotrichum several different root diseases in California
caricae (teleomorph: Glomerella cingulata). that are caused by basidiomycetes
Conidia are hyaline and generally cylindrical (Adaskaveg and Ogawa, 1990). Leaves on
or slightly elliptical. Setae, long, black, hair- affected trees are discoloured, severely wilted
like structures, are present in acervuli. and abscise prematurely (Thomas et al., 1948).

Table 11.2. Miscellaneous diseases and disorders of fig, Ficus carica.

Disease Cause

Alternaria internal rot Alternaria alternata, Curvularia spp.,

Dendryphiella vinosa, Epicoccum purpurescans,
Stemphylium botryosum and Ulocladium atrum
Alternaria leaf spot Alternaria alternata and Alternaria spp.
Branch wilt Hendersonula toruloidea
Brown leaf spot Phyllosticta sycophilla
Canker Tubercularia fici
Cephalosporium leaf spot Cephalosporium fici
Dagger nematode Xiphinema index
Frost dieback Abiotic disorder
Immature fruit drop Abiotic disorder
Leaf blight Rhizoctonia solani AG 1-IB
Leaf spot Cercospora fici
Lesion nematode Pratylenchus vulnus
Limb blight Erythricium salmonicolor (synonym: Corticium salmonicolor )
Macrophoma canker and fruit rot Macrophoma fici
Ormathodium spot Ormathodium fici
Reniform nematode Rotylenchulus macrodoratus
Root rots and wood decay Inonotus cuticularis, I. rickii, Pleurotus ostreatus and
Schizophyllum commune
Sclerotinia shoot blight Sclerotinia sclerotiorum
Sclerotium blight Sclerotium rolfsii
Stem gall and canker Nectriella pironii
Sunburn Abiotic disorder
Thread blight Corticium (Ceratobasidium?) stevensii

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258 T.J. Michailides

These symptoms may be confused with those from California where the disease is consid-
caused by asphyxiation, drought, insects and ered the second most common fruit decayer
Rosellinia root rot. Death of trees usually fol- (Doster et al., 1996). In its early stages, small,
lows periods of excessive wind, drought or brown, soft spots with white mycelia
heavy fruit loads (La Massèse et al., 1984). develop on the interior flesh of green fruit,
The causal agent, Armillaria mellea, is usually at the eye end of the figs. In contrast
described in Chapter 1. Its wide host range to smut, water-soaked areas do not appear at
and ability to survive as a saprophyte make it the lower part of the fig around the eye.
a difficult pathogen to control. If roots of pre- When the fruit is ripe, the pathogens pro-
viously grown plants exhibit signs of A. mel- duce abundant condiophores and conidia in
lea during the preparation of new plantations, the fruit cavity (Plate 72A and B).
two fumigants, carbon bisulphide and Twenty-three species of Aspergillus have
methyl bromide (most effective), can be used. been reported on fig in California, only a few
of which produce an Eurotium teleomorph
(Doster et al., 1996). Among these, E. amstelo-
Aspergillus mould dami (anamorph: A. amstelodami) may be the
most common. Other species include: A.
Although Aspergillus mould occurs in other flavus (Fig. 11.4), A. parasiticus (Fig. 11.5) and
countries, the only published work comes A. tamarii in Section Flavi, and A. alliaceus, A.

Fig. 11.4. (A) Culture of Aspergillus flavus on Czapek agar after 1 week at 25°C, (B) conidial heads (note
splitting), (C) conidial head, (D) SEM of a conidium and (E) SEM of a conidial head of the fungus (from CMI
descriptions nos 91 and 1251).

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Diseases of Fig 259

Fig. 11.5. SEM of (A) a conidial head and (B) conidium of Aspergillus parasiticus (from CMI description
no. 994).

melleus, A. ochraceus and A. sclerotiorum in 1.25–2.5 cm in length. These small surface

Section Circumdati. Although only a small cracks may cover much larger cankers (to
percentage of figs are affected by Aspergillus 15 cm) on the cambium (Hansen, 1948).
mould, it should be considered a serious Cankers can girdle and kill limbs.
problem since some of the agents produce Bacterial canker is caused by Pseudomonas
carcinogenic aflatoxins (A. flavus and A. para- fici. The bacterium is Gram negative, 0.5–0.6
siticus) or ochratoxins (A. alliaceus, A. melleus, ⫻ 1.5–2.6 ␮m, motile with four to five fla-
A. ochraceus and A. sclerotiorum). The conidia gella, and has optimum and maximum tem-
of A. flavus are pale green and finely rough- peratures for growth of 15 and 35–37°C
ened, those of A. parasiticus are dark green respectively. Colonies on nutrient agar are
and very rough, and those of A. ochraceus are white to creamy, and on King’s B medium
2.5–3.0 ␮m, finely rough-walled and ochre– produce a fluorescent pigment.
yellow to olivaceous. Infected shoots should be pruned (Pilgrim,
Figs become more susceptible as they 1950a). A 2% Bordeaux mixture should be
mature (Doster and Michailides, 1997). applied when buds are opening, followed by a
Wounding does not increase infection or 0.75% mixture after they are open, a 1% mix-
aflatoxin contamination in very immature or ture at the end of April, and a 1.5% solution in
very mature figs, and insect damage does August. In California, the Adriatics and capri-
not predispose fruit to infection. However, figs are more susceptible than Kadotas,
insects could play an important role in trans- Calimyrnas and Missions (Pilgrim, 1951).
mitting spores of these fungi. Reducing dust
in orchards may reduce the colonization of
figs by these pathogens. Botrytis limb blight (dieback) and fruit rot

First reported by Prunet (1903), the disease is

Bacterial canker now widely recognized (Massee, 1911;
Condit and Stevens, 1919; Manziano et al.,
Bacterial canker is the only bacterial disease 1990). It can blight a significant number of
that has been reported on figs. In California, shoots in trees during wet and cool springs.
it has only been found on limbs that are >4
years old, but in Italy it causes dark lesions
Symptoms
on leaves and elongated lesions and wilting
on new shoots. Symptoms on limbs start as Initial infections start at the base of young
wet spots, which dry later, causing cracks shoots that wilt, bend and develop light

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260 T.J. Michailides

green to brown leaves. Blighted shoots and Isolates from fig orchards produce
fruits of both fig and caprifig can be covered sparse to abundant conidia and sclerotia.
with abundant buff sporulation of the Sclerotia are small, hard, black survival
pathogen (Figs 11.6A and B). Under extreme structures and often are observed firmly
wet conditions, infected midribs and the sur- attached to the outside of decayed fig fruits
rounding areas of leaves turn black (Fig. 11.6B). Carpogenic germination to
(Manziano et al., 1990). produce apothecia can occur after a dor-
mant period of 2–6 months, but has not
Causal agent been reported in fig orchards.

The disease is caused by Botrytis cinerea

(teleomorph: Botryotinia fuckeliana), a cos- Epidemiology
mopolitan ascomycete on numerous crops. It B. cinerea can infect through injuries on fruit,
is described in Chapter 1.
and leaf and fruit scars to cause cankers
below and above the entry point (Fig. 11.6B).
Infected shoots blight, resulting in fruit
blight. Conidia from blighted shoots and
fruits are disseminated by air and initiate
new infections during cool, rainy weather on
susceptible tissues (leaf or bud scars) or
weakened shoots and fig fruits. English
(1962) showed that winter injury to fruits or
branches followed by infection by B. cinerea
caused shoot dieback.
This disease is more common on caprifigs
than on ‘Calimyrna’ figs. Since the pathogen
sporulates on caprifigs as late as July, these
shoots probably constitute an important
inoculum source (Ricci, 1972). Conidia conta-
minate the mamme crop of caprifigs and,
unless these are treated before storage, a sig-
nificant percentage decay. Although control
of this disease in figs has not been studied,
dipping the mamme crop in iprodione
(Rovral 50W) before cold storage can signifi-
cantly reduce the disease.

Endosepsis

Endosepsis occurs in California, Greece,

Turkey and possibly other areas.
Endosepsis is also called ‘brown rot’, ‘eye-
end rot’, ‘pink rot’ and ‘soft rot’, terms that
describe various phases and symptoms of
the disease (Caldis, 1927). In California,
endosepsis, smut and souring diseases can
Fig. 11.6. (A) Sporulation of Botrytis cinerea on a
cause combined losses of up to 50%
fig shoot affected by Botrytis blight (dieback) and (B) (Hansen, 1928). Handpicking and sorting of
a canker initiated from a fruit affected by Botrytis rot fruit for later treatment also raise the costs
(photos: T.J. Michailides). of production in some districts.

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Diseases of Fig 261

Symptoms less common are F. solani and F. dimerum

(Michailides et al., 1987; Wilson and
Any cultivar that requires pollination by the
Ogawa, 1979).
fig wasp can be affected. Green fruit exhibit
F. lactis produces microconidia in chains
brown or rust coloured areas on flower stig-
on polyphialides and few, small falcate
mas, flower bases or the entire flower. Upon
macroconidia (Gerlach and Nirenberg, 1982).
ripening, the brown streaks become
Some of the microconidia are pyriform.
yellow–brown spots that involve a number of
Colonies are white or light pink, with aerial
flowers and can develop on any part of the
hyphae. Unless indicated otherwise, the dis-
pulp, but usually are found near the eye. No
cussions below pertain to this fungus.
external symptoms are evident at this stage.
When fruit soften, the skin appears water
soaked, usually around the eye in a circular Epidemiology
spot and extending down the sides to the
The pathogen overwinters in mummified
neck. The water-soaked areas gradually turn
mammoni (summer crop) caprifigs and in
pink or purple. In other cases, only a small
mamme (winter crop) caprifigs until April of
water-soaked ring appears around the eye,
the following year (Plate 73). Propagules of
and a drop of dense, clear or amber syrup is
the pathogen can also be found on the sur-
exuded. This is common in ‘Calimyrna’, but
face of caprifig and ‘Calimyrna’ trees
not caprifig, fruits. The pulp becomes amber
(Michailides et al., 1987, 1996). Conidia of
and is considerably darker than the beige
the fungus are introduced from these loca-
colour of healthy fruit (Fig. 11.7). Affected
tions into caprifig and ‘Calimyrna’ fruit by
‘Calimyrna’ fruit will dry and appear healthy
female fig wasps as they enter to lay eggs.
if the pulp is not greatly affected. However,
Since the pathogen sporulates at approxi-
even slightly affected pulp has off flavours. A
mately the same time that adult female
‘slip-skin’ condition can also develop whereby,
wasps emerge, the wasps become contami-
the fruit epidermis is removed easily.
nated with conidia and transfer them to
healthy, developing fruit of the following
Causal agents caprifig crop, or from profichi caprifigs to
‘Calimyrna’ fruit during the caprification
The most common endosepsis agent is
process (Caldis, 1927). Parthenocarpic culti-
Fusarium lactis (O’Donnell et al., 1998), and
vars are not infected by F. lactis unless the
wasps have entered them accidentally. The
pathogen first invades the dead stigmas, but
eventually fills the interior of the fruit as it
begins to ripen.
Endosepsis decreases with distance from
an inoculum source, decreasing faster to the
south than in other directions (Michailides
and Morgan, 1998). Although endosepsis can
complete as many cycles per year as its vector
in caprifigs (polycyclic disease), there is no
secondary spread in ‘Calimyrna’ and
‘Smyrna’ figs (Michailides and Morgan, 1998).

Management
In California, control efforts are directed
towards reducing the proximity of
‘Calimyrna’ to caprifig trees and producing
Fig. 11.7. Endosepsis on a ‘Calimyrna’ fig (photo: clean female fig wasps. Caprifigs are now
T.J. Michailides). planted in separate orchards, usually far

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262 T.J. Michailides

from ‘Calimyrna’ crops. Excess caprifig trees endosepsis is reduced in the following pro-
and those in abandoned orchards are fichi fruit. Profichi figs that show external
destroyed (Smith and Hansen, 1935). symptoms of discoloration should be dis-
Dipping mammes in a solution of benomyl, carded before hanging them in ‘Smyrna’ fig
chlorothalonil, DCNA and potassium sorbate trees for pollination (Fig. 11.8). In
lowers the incidence of F. lactis, as well as California, caprifig growers test profichi for
Rhizopus and Alternaria (Obenauf et al., 1982). infection by Fusarium spp. (Michailides et
Good sanitation practices are even more al., 1994). If the caprifig orchard is close to
beneficial than chemical treatments the edible fig orchard, surplus profichi
(Michailides et al., 1996). Discoloured should be destroyed before they are mature
mammes should be discarded, since wasps (Smith and Hansen 1935). Avoiding over-
from these fruits usually are contaminated caprification is another way to reduce
with the pathogen (Michailides and Ogawa, endosepsis in ‘Smyrna’ type figs
1989). By treating disease-free mamme fruit, (Michailides and Morgan, 1994).

Fig. 11.8. Diagram depicting pollination in non-edible caprifigs and edible ‘Calimyrna’ figs, the life cycle of
fig wasp, Blastophaga psenes, and the disease cycle of fig endosepsis caused by Fusarium lactis and other
Fusarium spp. Depending on weather conditions, one to two additional wasp and disease cycles can be
completed during the summer and early autumn in California. The circular arrows inside the caprifigs
indicate successful development and emergence of adult female wasps through the fig ostiole.

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Diseases of Fig 263

Fig foot rot Fig mosaic

Fig foot rot was first reported in Brazil, and Fig mosaic has been reported in the British
in 1982 in Japan (Kato et al., 1982). Initial Isles, California, Greece, Italy, Libya, New
symptoms are wilting twig tips followed by South Wales and Yugoslavia. The first com-
leaf fall. Branches dry and entire trees may plete description of the disease was given by
die. Longitudinal sections of the trunk show Condit and Horne (1933). In California, it is
necrosis in cortical tissues and the cambium. thought to have originated on ‘Samson’
The disease is caused by Ceratocystis fim- caprifigs that probably were introduced from
briata (anamorph: Chalara sp.). Its perithecia Asia Minor in 1882. The disease then spread
are superficial or immersed in the substrate, through cuttings from this cultivar (Condit
globose, 90–210 ␮m in diameter, with a neck and Horne, 1933, 1941). It is also common on
29–715 ␮m in length, 25–37 ␮m at the base edible figs.
and 15–25 ␮m wide at the tip. The number
of ostiolar hyphae ranges from eight to 14,
Symptoms
with a mean width of 2–3 ␮m. The
ascospores are unicellular, hyaline, hat On leaves, mosaic spots are distinctive, con-
shaped and 2.7 ⫻ 6.4 ␮m (Valarini and trasting with the normal green colour of the
Tokeshi, 1980). The fungus produces macro- foliage (Fig. 11.9). The margins of the yellow
conidia and endoconidia. spots blend gradually through a light yellow
Valarini and Tokeshi (1980) assumed that colour into the dark green of healthy tissue.
the beetle Xyleborus ferrugineus was a vector Mosaic spots or lesions may be scattered uni-
of the pathogen based on its frequent associ- formly over the surface of the leaves or may
ation with diseased plants. In the field, appear as irregular patches of light green dif-
‘Branco’ and ‘Portugues’ were resistant fused widely throughout the leaf blade. Later
when they were used as rootstocks for ‘Roxo in the season, a rust-coloured band develops
de Valinhos’ (Valarini and Tokeshi, 1980). along the border of the mosaic spots, appar-

Fig. 11.9. Foliar symptoms caused by fig mosaic virus (photo: T.J. Michailides).

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264 T.J. Michailides

ently caused by the death of epidermal or Fruit splitting

subepidermal cells. Malformed leaves, the size
and shape of which vary greatly, also develop. This is a severe problem in some years, espe-
Mosaic spots on fruits are similar to those cially in ‘Calimyrna’ and ‘Adriatic’ figs.
on leaves. Premature fruit drop is sometimes Splitting is the result of sudden changes in the
associated with the presence of fruit or leaf internal fruit pressure brought on by either
symptoms on certain cultivars (Condit and cool temperatures and high humidity as the
Horne, 1943). fruit matures or showers during the fruit-
ripening period (Obenauf et al., 1978). Splitting
of ‘Calimyrna’ also can result from over-capri-
Causal agent
fication (too many seeds are developed).
Fig mosaic virus causes fig mosaic. To date, it Splitting begins in the eye of fruit, and
has not been characterized or isolated. ruins the fruit, since the pulp is exposed to
Although Quacquarelli (1971) isolated insect and fungus attack. Over-caprification
Sowbane mosaic virus from figs with mosaic can be prevented by reducing the number of
symptoms, it does not cause fig mosaic caprifigs or by isolating caprifig plantings.
(Plavsic and Milicic, 1980). Insect control may prevent further spoilage of
Symptoms of fig mosaic are not transmissi- split fruits.
ble mechanically. They are associated with
polymorphic particles that are present only in
symptomatic plants, and are either spherical Phomopsis canker
and 120–160 nm in diameter or elongated and
200 nm in length (Plavsic and Milicic, 1980). Phomopsis canker, which is also known as
fig canker, occurs on all commercial culti-
vars of fig in California. ‘Kadota’ is seriously
Epidemiology
affected, ‘Calimyrna’ suffers minor damage,
Fig mosaic is not seedborne, but can be and ‘Mission’ and ‘Adriatic’ are rarely
transmitted by grafting and females (but not attacked (English, 1951). The disease has
males) of the eriophyid mite, Aceria ficus also been reported in Italy (Manziano et al.,
(Flock and Wallace, 1955; Smith, 1972). A sin- 1990).
gle female is capable of transmission, and
minimal acquisition and infection feeding
Symptoms
periods are <15 min. For optimal transmis-
sion, a feeding period of 24 h on the source Dead bark and wood develop on pruning
and test plants is necessary (Smith, 1972). wounds or on injuries caused by frost or
Symptoms are produced <10 days after feed- sunburn. The bark in the older portions of
ing (Jeppson et al., 1975). cankers becomes bleached, cracked and
sunken (Wilson and Ogawa, 1979). Pycnidia
of the pathogen develop on the outer bark
Management
layers. Cirrhi of conidia exude from pycni-
‘Black Mission’ is the most seriously dam- dia under humid conditions. Elongation of
aged cultivar, and ‘Kadota’ and ‘Calimyrna’ elliptical cankers results in yearly zonations,
are the least affected. ‘Black Bursa’ is affected with the fungus sporulating more profusely
in Turkey. F. palmata, or progeny with F. in the outermost zone. Enlarged cankers gir-
palmata as the male parent, appear to be dle and kill branches on which the withered
immune (Condit and Horne, 1933). foliage remains.
Care must be taken to ensure that stocks
are not propagated from mosaic-affected
Causal agent
trees (Khalil, 1982). Apical meristem culture
and thermotherapy of alternating light (16 h The disease is caused by Phomopsis cineras-
5000 lux at 37°C) and dark (8 h at 34°C) are cens (teleomorph: Diaporthe cinerascens). Its
effective (Gella et al., 1997). pycnidia are black, 250–500 ␮m, globose and

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Diseases of Fig 265

immersed in the bark from which their osti- Phytophthora fruit rot
oles emerge. They produce two types of pyc-
nidiospores on host bark: ␣-conidia, which Phytophthora fruit rot is caused by
are elliptic–fusoid and 6–9 ⫻ 2–2.5 ␮m; and Phytophthora palmivora, which is described in
β-conidia, which are filiform, 20–25 ⫻ 1 ␮m, Chapter 1. It was first reported in Japan, and
mostly hooked and on short pedicels. is also known in Florida, India, New South
Wales and Taiwan (Nisikado et al., 1941; El-
Gholl and Alfieri, 1984).
Epidemiology
The fungus infects both wounded and
P. cinerascens is a wound parasite. The fre- non-wounded green fruits, creating water-
quent association of Phomopsis canker with soaked lesions that are covered with white
pruning wounds explains the disease’s fluffy mycelium of the pathogen. The disease
importance on ‘Kadota’, a cultivar that is is more common in the lower part of the tree,
pruned heavily (Hansen, 1949a, b), and but fruits and limbs in higher parts of the
infection through leaf scars probably occurs. trees are also affected. In Japan, the pathogen
The same fungus causes a canker on weep- caused a white powdery rot on fig fruits,
ing fig, F. benjamina, and a twig blight on the leaves, young buds and young stems (Katsura
same species indoors (Hampson, 1981; and Yamamoto, 1969). Isolates of P. palmivora
Anderson and Hartman, 1983). from ‘White Genoa’ figs caused rot on apple,
The fungus survives in cankers and on Japanese pear, persimmon, aubergine, tomato
pruned branches that are left on the and potato (Nisikado et al., 1941).
ground. During rainy weather, the spores
are exuded from pycnidia and washed
Root-knot nematodes
down along the branches and trunk of the
tree. They germinate and start new cankers
Root-knot nematodes are probably the most
wherever they come in contact with an
important and widespread parasites of figs
unhealed wound. Pruning tools contami-
(McBeth, 1949). Although the exact species of
nated with pycnidiospores can disseminate
Meloidogyne that are involved have not been
the fungus.
reported, M. arenaria has been isolated from
Canker extension on ‘Kadota’ trees is
roots of ornamental figs (Davide, 1979). Trees
restricted between April and October, appar-
in sandy soil are very susceptible (Matz, 1918).
ently due to active growth of the host. Root-knot nematodes cause characteristic
However, from November to February, they knots or galls on fig roots. The adult female
are highly susceptible (English, 1952). is pearl white, pyriform and ~2 mm long.
Egg masses as large as her body are often
Management found near or attached to her body, but they
may also be found outside the gall. In the
Although removing and burning infected juvenile stages, this nematode is long and
branches from young trees is beneficial, slender. The adult males retain their vermi-
older trees with a high incidence of cankers form shape while the adult females become
may need to be cut back to the trunk and pyriform as they reach maturity. The
treated with a protective fungicide (English, females usually develop just beneath the
1953). Young, non-bearing trees should be surface of the root and, as a result of their
pruned late in the dormant season, and growth and secretions, induce the formation
pruning tools that are used on affected trees of galls. Heavily infected roots may actually
should be disinfested with dilute sodium die. As a result, affected plants have reduced
hypochlorite (2500 ␮m ml⫺1) (Pilgrim, absorption and transport of water and nutri-
1950b; English, 1953). Because vigorous ents, and exhibit stunting, chlorosis and
trees appear to be least susceptible, cultural reduced fruit production (McBeth, 1949).
practices that are designed to keep the trees To date, no research has been done to
in good health should be followed. explore resistance to nematodes in Ficus spp.

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266 T.J. Michailides

Fumigation before the establishment of a fig Severe cases on fig result in premature defo-

nursery can limit the dissemination of liation, leaving only immature fruit that may
nematodes, but in orchards is very expen- drop (McKenzie, 1986).
sive and may need to be limited only to the
planting rows (McKenry and Thomason, Causal agent
1975; Carles, 1985).
Rust is caused by Cerotelium fici (Laundon
and Rainbow, 1971). Pycnia and aecia of this
Rosellinia root rot fungus are unknown. Uredinia are hypophyl-
lous and scattered on the fruit or sometimes
Rosellinia root rot was recorded in figs grown grouped around the edge of leaves or in dew
drop runs on lamina (McKenzie, 1986).
the Aegean plains in Turkey (Igriboz, 1940;
Urediniospores are globose, obovoid, ellip-
Karaca, 1979). It made the Turkish fig industry
soidal, or angular, 19–30 ⫻ 15–23 ␮m and
move gradually to the more hilly areas of the
sparsely echinulate. Telia have been reported
Aegean Region. Rosellinia root rot is a more
only twice (Laundon and Rainbow, 1971).
common and devastating disease of fig than
Teliospores are hyaline, smooth, barrel
Armillaria root rot (La Massèse et al., 1984).
shaped, oblong or ovoid, 14–22 ⫻ 10–13 ␮m,
Armillaria root rot and Rosellinia root rot form in loose chains and tend to separate.
can be distinguished reliably only on the
basis of underground signs. Rosellinia neca-
trix (anamorph: Dematophora necatrix) is Epidemiology
described in Chapter 1. It produces cottony No information is available on physiological
white mycelia on root surfaces that are sig- specialization in C. fici, and its life cycle is
nificantly different from the rhizomorphs incompletely known (Laundon and
and mycelial plaques formed by A. mellea. Rainbow, 1971). In the southern USA, it is
As an ascomycete, R. necatrix also does not believed to overwinter as urediniospores on
produce basidiomes. fallen leaves that infect young leaves in the
Controlling Rosellinia root rot is extremely spring (Krezdorn and Adriance, 1981).
difficult. Fumigants, such as allyl bromide,
ammonium hydrosulphide, carbon bisul- Management
phide, carbon tetrachloride, chloroform,
chloropicrin, ethylene dibromide and forma- Fallen leaves should be removed from the
lin pentachlorethane have all failed to control orchard and destroyed. Fig rust can be con-
the disease. Removal of all infected roots trolled with sulphur dusts, 0.05% Vitavax
(Sinhri and Mishra, 1984), or Bordeaux mix-
from planting sites is also futile since the fun-
ture when applied to young leaves (Laundon
gus can survive in small pieces of dead roots.
and Rainbow, 1971). Other fungicides, such
as zineb and maneb, are also effective if
applied before the appearance of disease
Rust
(Assawah et al., 1966).
Fig rust is widely distributed on F. carica,
other species of Ficus, Morus alba, M. nigra Smut
and species of Broussonetia and Maclura. The
disease has been reported in Bermuda, Smut is one of the most studied diseases of
Egypt, India, Italy, New Zealand, the USA figs. It is not a true smut, but refers to the
(Arkansas and Florida), Venezuela and black powdery conidia of the causal fungi
Yugoslavia (Matz, 1918; Dale, 1958). that form on affected figs.

Symptoms Symptoms
Angular leaf spots are pale and reddish In its early stages, small, brown, soft spots
brown due to the formation of uredinia. develop on the interior flesh of green fruit

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Diseases of Fig 267

while they are still on the tree (Phillips et al., Michailides et al. (1991) showed that beetles
1925). Decay usually begins at the eye end infested with A. niger and caged with
and progresses as water-soaked areas healthy ‘Calimyrna’ figs did not increase
around the eye. When fruit are ripe, the fun- levels of smut relative to fruit that were
gus produces abundant black masses of coni- inoculated with A. niger alone, but that
dia in the fruit cavity (Fig. 11.10). dusting figs with soil powder and A. niger
did.
Thrips were also implicated in carrying
Causal agents
fig spoilage fungi including A. niger
The disease is caused by species of Aspergillus (Hansen, 1929; Smith and Hansen, 1931;
in Section Nigri. Initially, it was thought that Davey and Smith, 1933; Baker, 1939). The fig
only one species, A. niger, was involved; it is mite, although present in essentially all
described in Chapter 1. However, Doster et al. ripening figs, has not been implicated as a
(1996) showed that five distinct, black-spored carrier of any fruit pathogens (Hansen and
taxa are involved in California: A. niger var. Davey, 1932). In the Smyrna fig district of
niger, A. niger var. awamori, A. japonicus var. Turkey, Hagan (1929) proposed that an ant,
aculeatus, A. japonicus var. japonicus and A. Formicomus ionicus, which entered ripening
carbonarius. A. niger vars niger and awamori fruit was a prolific disseminator of smut
were responsible for 93% of the affected spores and a greater threat in Turkey than
‘Calimyrna’ figs in 1993 and >99% of the Carpophilus spp.
‘Conadria’ figs in 1994 (Doster et al., 1996).
Management
Epidemiology
Fruit of the ‘Black Mission’ and ‘Kadota’ cul-
Hodgson (1918) first proposed that insects tivars are affected less than those of
were responsible for transmitting smut. ‘Adriatic’ and ‘Calimyrna’ (Phillips et al.,
Transmission tests showed that the dried 1925). Removing old fruit culls and refuse on
fruit beetle, Carpophilus spp., and the vine- which dried fruit beetles and vinegar flies
gar fly, Drosophila melanogaster, were able to might persist and preventing dust in the
transmit spores of the fungus to edible figs orchard may help to reduce decays caused
(Phillips et al., 1925). More recently, by these fungi.

Fig. 11.10. Smyrna type fig with smut caused by Aspergillus Section Nigri (photo: T.J. Michailides).

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268 T.J. Michailides

Soft rot eyes have sterile pulp (Phillips et al., 1925).

Affected figs develop a pink colour and
The disease was first described in Louisiana later become water soaked. A pink syrupy
(Edgerton, 1911). It also occurs in California, liquid exudes through the eye, dropping on
Italy, Texas and Turkey (Taubenhaus, 1936; to the leaves or congealing at the eye. The
Manziano et al., 1990). Mature fruit become gas bubbles and characteristic alcohol
mushy, are eventually covered by mycelium odour that develop indicate fermentation
and sporangia of the pathogen, and ulti- of the pulp. In later stages, the pulp disinte-
mately fall from the tree. grates and usually is covered by a white
Soft rot is caused by Rhizopus stolonifer scum. Affected fruits lose firmness, sag,
(Fig. 5.3) and Mucor spp. It is found regularly turn black, shrivel and either drop or hang
in the cavity of ‘Calimyrna’ and ‘Conadria’ on the twig. A dead spot or ‘eye canker’
figs (T.J. Michailides and M.A. Doster, often develops where the peduncle is
unpublished data). A similar soft rot of fig attached to the tree.
fruit is caused by Choanophora cucurbitarum
(Taubenhaus, 1936).
Causal agents
Soft rot is very common when fruits are
ripening during rainy weather in the sum- Souring is caused by yeasts and bacteria
mer. Rains during ripening result in cracks (Mrak et al., 1942). From affected fruit, Miller
near the eye. Airborne sporangiospores of and Phaff (1962) recovered species of yeast in
the causal fungi are abundant and can infect Saccharomyces and Pichia, and of the apiculate
fruit through these cracks. Spores are also yeasts Hanseniaspora kloeckera, Candida and
disseminated by the dripping juice of Torulopsis. Approximately the same yeast
affected fruit, rainwater, wind and insects flora are found on the exterior and in the
(Edgerton, 1911). intestinal tract of a beetle, Carpophilus
Nearly all large, soft-skinned cultivars are hemipterus, which feeds on soured figs.
very susceptible to this decay, while smaller Because Koch’s postulates were not per-
figs, such as ‘Celeste’ and ‘Reine Blanche’, formed with each of these yeasts, it is not
are less so. During conducive conditions, lit- known whether they all cause souring.
tle can be done to control this disease.
Edgerton (1911) suggested that figs should
Epidemiology
be harvested regularly and that ants be kept
out of trees. Insects transmit the organisms that cause
souring (Howard, 1933). Beetles and fruit fly
species that have been implicated as vectors
Souring apparently cannot enter the fig until its osti-
ole opens upon ripening, and yeasts that
Fermentative spoilage or ‘souring’ of figs is a cause fermentation were not present in
common problem. It occurs while fruit are experimental figs until after the ostioles had
still on the tree. opened. Periodic swarms of the cotton leaf
worm moth, Alabama argillacea, also transmit
the disease during feeding (Hull, 1929).
Symptoms
Symptoms are most distinct on fruit of
Management
parthenocarpic cultivars, such as ‘Adriatic’,
that have not been caprified (Caldis, 1930). In Exclusion of insects from trees almost com-
caprified figs, the symptoms of souring may pletely eliminated souring (Phillips et al.,
be confused with those of endosepsis. The 1925), although it did not prevent smut and
symptoms are noticeable only when the fruit mould (Davey and Smith, 1933). Souring
ripens and the eye is wide open. was also reduced when figs were protected
Deterioration of any kind does not begin either by bagging or by closing the ostiole
before the eye opens, and figs with closed with tanglefoot (Howard, 1933).

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Diseases of Fig 269

Surface mould or contact spot greenish to yellow-dark greenish conidio-

phores of A. flavus and A. parasiticus.
Surface moulds or contact spots are caused
by Alternaria alternata, Aspergillus niger and
Causal agents
Cladosporium herbarum. Other fungi that are
involved include A. flavus, A. parasiticus and A. alternata produces conidiophores on the sur-
occasionally other species of Aspergillus and face of infected areas, bearing long, sometimes
Penicillium (T.J. Michailides, unpublished branched, chains. A. niger produces conidio-
data). Some of these diseases were first phores bearing heads of conidia in chains on
reported in California (Smith and Hansen, the surface of infected areas. Both are
1931). All fig cultivars are susceptible, but described in Chapter 1. A. flavus, A. ochraceus
Alternaria surface mould is most severe on and A. parasiticus are described under
‘Kadota’ figs. Alternaria surface rot is a limit- Aspergillus mould in this chapter.
ing factor in the production of fresh fruit or Conidiophores of C. herbarum are erect, arising
fruit for canning (Bewaji et al., 1977). in tufts from the epidermal cells. It is described
in Chapter 6 under Cladosporium spot.
Symptoms
Epidemiology
Surface moulds occur on both green and ripe
fruit, developing mainly on the surface of C. herbarum, A. alternata and Aspergillus spp.
fruit that are in contact with other fruit (Fig. are common in the air, on plant surfaces and
11.11). Symptoms differ according to the in soil (Ingol, 1971), usually developing on
causal agent. For C. herbarum, initial dark dying or dead tissues of plants. On figs, C.
olive green specks develop and enlarge to herbarum and A. alternata can be found on tis-
become depressed and yellowish olive in sues of green fruit damaged by frost or sun-
colour. The first symptoms of Alternaria fruit burn and on wounded green or mature fruit.
rot are water-soaked areas, which soon are While C. herbarum predominates on green
covered with olivaceous conidia of the fun- fruit, A. alternata is a serious problem on ripe
gus. The first symptoms of Aspergillus infec- fruit (English, 1954). If rains prevail during
tions are water-soaked areas, which soon are harvest, Alternaria rot becomes a problem in
covered with white, then yellow, and finally ‘Calimyrna’ fruit, usually developing on the
black conidiophores of A. niger or yellow- surface where figs touch.

Fig. 11.11. Surface or contact spot on ‘Calimyrna’ figs (photo: T.J. Michailides).

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270 T.J. Michailides

An important source of inoculum is the market fruit, preharvest and postharvest

picking box. Wooden boxes become contami- sprays with chlorothalonil reduced
nated with fungi during the season, and 14 Alternaria surface rot more effectively than
different species were isolated from the inner benomyl or maneb (Bewaji and English,
surfaces of picking boxes, with C. herbarum 1976; Bewaji et al., 1977).
and A. alternata being most prevalent Picking the fruit before it becomes over-
(Harvey, 1956). Wooden boxes have been ripe reduced Alternaria rot, and a modified
replaced with plastic ones that are easier to atmosphere of ≥23% CO2 during storage
disinfest and keep clean. and transit was as effective as immediate
storage at 0°C (Brooks and McColloch,
1938). To prevent contamination of har-
Management
vested fruits, picking boxes should be kept
Sprays with protectants, such as zinc clean and washed regularly.
coposil and Dithane Z78 (zinc ethylene bis-
dithio-carbamate), in the first week of
September resulted in reduced disease Acknowledgements
caused by C. herbarum and A. alternata
(English, 1954). A combination of benomyl, I thank M.A. Doster and D.P. Morgan for
potassium sorbate, DCNA (Botran 75W) their technical assistance in the studies on
and chlorothalonil appeared to be very endosepsis and smut, J. Doyle and L.
promising in reducing Alternaria rot inci- Ferguson for useful discussions on fig, and
dence in profichi caprifigs (Obenauf et al., the California Fig Instutute (Fresno,
1982). Maneb is registered for spraying California) for financial support of research
‘Kadota’ figs to control Alternaria rot in on diseases of figs.
California (Dibble et al., 1972). On fresh-

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