Journal of Threatened
Taxa | www.threatenedtaxa.org | 26 April 2018 | 10(5): 11672–11678
The
invasive aphid Pterochloroides persicae (Cholodkovsky, 1899) (Hemiptera: Aphidoidea: Lachninae) recorded
on important fruit trees in Kashmir Valley, India
Govindasamy Mahendiran 1, Shahid Ali Akbar 2 & Mudasir Ahmad Dar 3
1 ICAR- National Bureau of Agricultural Insect Resources
P.B. No. 2491, H.A. Farm Post, Bellari Road, Hebbal,
Bengaluru, Karnataka 560024, India
2,3 Division of Plant Protection, Central Institute of
Temperate Horticulture, Srinagar, Jammu & Kashmir 191132, India
1 mahi.iari@gmail.com, 2 kingakbarali@gmail.com
(corresponding author), 3 mudasir753@rediffmail.com
Abstract: Pterochloroides persicae (Cholodkovsky,
1899) is reported here for the first time from the Kashmir Valley. The aphid is
seen to infest almond, peach, plum orchards in the region. Monitoring of the pest was carried out
in the peach and almond fields of the Central Institute of Temperate
Horticulture (CITH) during the years 2014–2016. Seasonality and
bio-rational management practices of the pest are discussed.
Keywords: Aphid, bio-rational, natural-enemies, pest, Pterochloroides persicae,
seasonality.
doi: http://doi.org/10.11609/jott.2968.10.5.11672-11678
Editor:
B.A. Daniel, Zoo Outreach
Organization, Coimbatore, India. Date of publication: 26 April 2018
(online & print)
Manuscript details: Ms # 2968 | Received 08
August 2016 | Final received 07 March 2018 | Finally accepted 25 March 2018
Citation: Mahendiran, G., S.A. Akbar & M.A. Dar (2018).
The invasive aphid Pterochloroides
persicae (Cholodkovsky,
1899) (Hemiptera, Aphidoidea:
Lachninae) recorded on important fruit trees in
Kashmir Valley, India. Journal of Threatened Taxa
10(5): 11672–11678;
http://doi.org/10.11609/jott.2968.10.5.11672-11678
Copyright:
© Mahendiran et al. 2018. Creative Commons
Attribution 4.0 International License. JoTT
allows unrestricted use of this article in any medium, reproduction and
distribution by providing adequate credit to the authors and the source of
publication.
Funding: Second author thanks the Department of Science and Technology (DST), Govt. of India, New
Delhi [N-PDF Fellowship program: File No: PDF/2015/000866].
Competing interests: The authors declare no competing interests.
Acknowledgements:
Sincere thanks are due to director CITH and ICAR for
facilitates made available at CITH. Sense of deep gratitude
towards Dr. Mariusz Kanturski
(Faculty for biology & Environmental Protection University of Silesia) for
suggestions and much acknowledged help. The authors would also like to
thank anonymous reviewers for helpful comments and suggestions about the
manuscript.
Aphids are among
the most destructive insect pests on cultivated plants in temperate regions (McGavin 1993).
They passively feed on the sap of phloem vessels in plants, creating
lack of vigor along with frequent transmission of
various viruses to their hosts (more than 50 aphid species are cited to be
vectors; (Robert & Bourdin 2001). Though enemies of farmers and gardeners
these represent a highly successful group of organisms on planet earth
(Blackman & Eastrop 1994; Piper 2007). About 4,702 species are known globally (Remaudiere & Remaudiere
1997), among which 1,015 species occur in the oriental region and 653 species
belonging to 208 genera have been reported from India, with a major portion
(310 species) being reported from the northeastern
region of India (Agarwala & Ghosh
1984, Verma & Das 1992; Ghosh
& Singh 2000; Ghosh & Ghosh
2006). Among globally known aphids,
around 250 species are dependent on agriculture and forestry species (Blackman
& Eastrop 1994; Aphid Species File 2016), 51
species have been found to affect more than 31 species of agricultural crops
from the Kashmir Division (Jammu & Kashmir, India) (Bhagat
1986a,b). Aphids represent a
dominant and economically important group of insects from the region.
During the present
study, the alien invasive aphid Pterochloroides
persicae (Cholodkovsky,
1899) of temperate crops is reported here for the first time from the Kashmir
Valley. From India, the species was
previously reported sporadically only from the states of Himachal Pradesh and
Punjab, on peach (Bindra & Bakhetia
1970; Mann et al. 1979). Review of the pest status, biology and control measures are
provided here. The sporadic
appearances of the species in the valley suggest that it has not flourished in
the region yet but considering the economic importance, its presence in the
region should be taken seriously and immediate measures for its control and
regional quarantine should be enforced.
Materials and
Methods
Specimens were
collected by hand picking method.
Weekly observations on aphid population were recorded using standard
University of California, integrated pest management (UC IPM) sampling
protocols for aphids. The weather
parameters used are relative to the place and not exact and were procured from
the regional metrological department.
Taxonomic analyses were conducted using Olympus SZX16 stereo zoom
microscope. For
digital images, ProgRes0 CapturePro v.2.8.0. evolution digital camera was used on the same microscope
with Combine ZP-Montage software. Later, images were cleaned with Adobe
Photoshop CS6. The collected
material has been deposited in the Biosystematics Laboratory of the Central
Institute of Temperate Horticulture (CITH–Srinagar) for future reference.
Results and
Discussion
Distribution: Pterochloroides
persicae also called brown peach aphid, woody
aphid, clouded peach bark aphid, cloudy-winged peach aphid depends on its host
plants Prunus spp. (almond, apricot, peach)
and Citrus species (Ciampolini et al. 1997). The species is well documented with
distributional records available for many regions (Blackman & Eastrop 1994; Ateyyat & Abu-Darwish 2009; Knowledge Bank-Plantwise
2015). The species is persistent (Liotta & Maniglia 1993),
serious (Kairo & Poswal
1995), invasive (Ciampolini
et al. 1997; Bounfour et al. 2005) with
distributional records suggesting gradual extension towards the eastern
Mediterranean and central Asia (Kairo & Poswal 1995).
Damage: Pterochloroides
persicae feeds on the brunch cortex (phloem),
which results in general weakening of the young fruit trees, withered branches
and reduced yield (Stoetzel 1994). The species is not listed as
transmitting a virus (Chan et al. 1991) but healthy densities result in
premature fruit drop, leaf curling, an irregular curvature of twigs, stunted
growth and sooty mold fungal growth development, due
to excessive honey dew production (Mann et al. 1979; Hondru
et al. 1986). The species does not
depend on attendance by ants for its survival as it can efficiently eject its
honeydew but ants are seen collecting existing honeydew deposits (Talhouk 1977).
Although polyphagous, peach is the most
preferred host plant of the aphid, with regards to its enhanced fecundity and
developmental rates, peach is also considered as the best
suited host plant for its mass rearing. The species is most prolific in its
rates of reproduction at 200C temperature (Khan et al. 1998). The colonies constantly move away from
exposed areas of sun and actively move on the trees but spread to neighbouring
trees is seen to be very poor (Mann et al. 1979).
Not much is known
about the natural enemies of the aphid with Chrysoperla
carnea, Coccinella
undecimpunctata, Pauesia
antennata and Coccinella
algerica as other alien invasives
are considered as most promising bio-control agents of the pest (Mirabzadeh et al. 1998; Abd-Rabou
2008; Rakhshani et al. 2005; Mdellel
& Kamel 2012). Entomopathogenic
fungi against the species include Beauveria
bassiana (Hypocreales, Cordycipitacae); Metacordyceps liangshanensis
and Metarhizium anisopliae
(Hypocreales, Clavicipitaceae)
(Tsinovskii & Egina
1972). Yeast, vegetable oil,
mineral oil, potassium sulphate + detergent have been trialed
successfully against the control of the species (El-Salam 2001; Braham et al.
2007). Several chemicals at different
concentrations and varying compositions have also been successfully used
against the aphid: Dimecron (phosphamidon)
or diazinon, Etiol, Anthio (formothion) and Lebaycid (fenthion) at 0.2%
concentration (Velimirovic 1997); 0.05% of malathion and phosalone (Mann et
al. 1979); Confidate¨ (Imidaclorpid)
and Chlorcyrin¨ (Chlorpyrifos
and Cypermethrin) (Ateyyat
2008); Patron¨ (diflubenzuron) and Trivap¨ (Cyromazine) as biorational insecticides (El-Salam 2001; Ateyyat & Abu-Darwish 2009);
ULV sprays of dimethoate and monocrotophos
applied at rates of 135 and 100 g a.i./ha,
respectively, and 0.025% high-volume sprays of chlorpyrifos,
dimethoate, leptophos, monocrotophos, oxydemeton-methyl,
permethrin, phosalone and quinalphos (Sandhu & Sohi 1978); non-specific insecticides parathion, dimethoate, mevinphos and acephate along with the addition of some wetting agent (Ciampolini & Martelli
1980). Use of Phosalone
has also been advocated (Sandhu & Sohi 1978) for its effectiveness and for being harmless to non target insects.
Biology: The life-cycle
of the species is complex, having alteration of parthenogenetic
and sexual generations, apterous and alate forms exhibited and persistently overlapping. Anholocyclic
and holocyclic means of life cycle of the species
from Lebanon was detailed by (Talhouk 1977). Holocyclic
populations with sexual phase in autumn have only been observed in cooler
regions with oviparous females very similar to apterous
viviparous females, except for their larger size (Archangelsky
1917; Wieczorek et al. 2013). Environmental conditions predominately
influence population dynamics of the species with 18 overlapping generations
reported in a year (Darwish et al. 1989); however
fewer offspringÕs are produced per female in the winter than in summer (Velimirovic 1977).
It takes 66–82 hr, 64–80 hr and 144–152 hr for the
first, second and third instar to complete development; 16–48 hr, 122–280 hr and
8–24 hr for the pre-oviposition,
oviposition and post-oviposition
periods. 17–55 nymphs/female are raised in the entire reproductive life
with an estimated reproductive rate of 4.51 nymphs/female per day (Velimirovic 1977; Darwish et al.
1989). Khan et al. (1998) reported
4-instars with eight days nymphal period and affirmed
that average life span of the aphid is 41 days.
Pterochloroides persicae (Cholodkovsky)
Lachnus persicae Cholodkovsky,
1899: 472
(Images 1–9)
Material examined: CHT-IC/NPDF/B1-3, India: Kashmir,
Srinagar, CITH, 33.590N & 74.500E, 1,500m, 122 Apterous viviparous and 32 Alate
viviparous females, 10.v.2014, 12.x.2014, 21.iii.2015, 17.v.2015, 11.viii.2015,
11.iv.2016, 16.v.2016, 28.v.2016, 09.vi.2016, 21.xi.2016, coll. G. Mahendiran and S.A. Akbar.
Description: Apterous
viviparous and alate viviparous females are the most
conspicuous form in the colony, when fully grown these are large in size,
having morphometric ranges of body length and width as 3.75–4.00
mm and 2.61–2.70 mm respectively; color
dark brown to black with some white patches, dorsum of abdomen with a double
row of large tubercles (Images 1–3); antennae six segmented; forewing
with large distinctive dark patches and long rostrum easily demarcates the
species from other con-generic members (Images 4–9). Sexual forms (oviparous females and
males) although previously reported, have only recently been detailed and
described by Wieczorek et al. (2013).
Seasonality (Images
7–22): Periodic
monitoring of the aphid was carried in non-native peach and almond fields of
the Central Institute of Temperate Horticulture (CITH) during 2014–2016
growing seasons. CITH is situated
at 33.590N and 74.500E at an altitude of 1,640m. Eggs were initially sighted from the
second week of November with few nymphs also present but infrequent in occurrence
(Image 10). These eggs are mostly
laid on the stems and are light brownish in colour but change to dark brownish
to blackish-brown in colour with time.
These eggs cover the entire surface of branches and twigs and are easily
sighted (approximately 600–750 eggs/10cm2) (Image
11). These eggs are comparatively
bigger and thicker shelled than other aphid eggs. The aphid overwinters in the egg stage
and these eggs begin to hatch from the first week of March. By the middle of March almost 70–80
% of the overwintering eggs are hatched.
Newly emerged nymphs are most frequently sighted from the second week of
March; moving upwards towards the small twigs for
feeding (Image 12). These grow and
attain full maturity and start to produce young ones from the second week of
April (approximately one month from the date of hatching) (Image 13). First noticeable population build up of
the species is observed during the second week of May with several batches of
45–60 individuals easily seen along the underside of tree
trunks. Alate
forms are mostly seen from the beginning of the second week of May (Image
14). Population density peaks in
the month of May and decreases towards the latter half of the month and more in
the early weeks of June. The
species gradually becomes inconspicuous during the early weeks of July. The second population build up starts
towards the end of July. The population of the species gradually increases in
the month of August along with the appearance of a few alate
forms. These alates,
however, are most conspicuous during the latter half of August, concurring with
dispersal and an increased degree of infestation of other trees and evident
presence of several coccinellids availing
opportunities for egg laying.
Second noticeable population build up of the species becomes most
palpable during the latter half of October, with the underside of tree trunks
occupied by the species along with large areas of blackened soil surface near
the tree trunks (Images 15–17). The population of the species continues
to increase in September and in the early weeks of October; however a drastic
decline in the population occurs in the month of December. Longevity of females ranges from 41–60
days during which she lays several nymph batches (35–43 nymphs per
female).
Probably six to
eight generations of the aphid species are prevalent in a season from the
region; however, ambiguous numbers observed could not be concluded and
confirmed. The species was most
prolific during the months of April, May, September and October with the
population density of late season generation much higher compared with the
early season. Mean average
temperature recorded during the early season months (April, May) was 120C
with maximum and minimum temperatures been 18.60C max and 8.60C
min; dew point of 120C, precipitation of
31.0mm and wind speed of 9km/hr.
Mean average temperature of 160C with maximum and minimum
temperatures of 80Cmax and 230Cmin; dew point of 170C,
precipitation of 10.0mm and wind speed of 89km/hr for
the late season (September, October, November). The aphid was seen most prolific at mean
temperature of 20–22 0C. The species was seen to be active at
temperatures as low as 30C.
There exists a definite relationship between the temperature and the
population dynamics of the species, however in an open system with multiple
environmental factors no definite correlation was confirmed. The species can be used as an ideal
model for the concept of adaptive phenotypic plasticity (Halkett
et al. 2004) which states the investment in sexual
reproduction to be proportional to winter severity and forecasting climate
predictability; however any definite inference on the subject is beyond the
scope of this paper and would require much more information. The aim of this
review is to encourage understanding of this fascinating aspect of aphid
biology and to promote more studies in this direction.
Several safe
chemical combinations were used (Neem seed kernel
extract-NSKE, Neem plant extract; Soyabean,
Oliver, Rosemary Lavender oils; MetOH extract of Crocus
sativus, Spilanthes
ciliate, Datura stramonium,
Juglans regia
and Vim gel soap solution) at various concentrations against Pterochloroides persicae. Days after spray (DAS) data showed that
all the combinations were prolific against the invasive species. Data recorded seven days after spray
showed that in the beginning all the treatments provided cent percent control of the pest; after 15 days pest population
remained minimal in rosemary oil + soap solution (@ 0.1ml+10ml /litre of water)
whereas negligible resurgence was observed in other combinations which also
subsided to zero in following weeks.
These chemicals were responsible for causing membrane disintegration on
contact and the repellant nature of the spray also
dispersed the surviving aphids (Image 18).
The data generated is under evaluation for further use and
recommendation. Some of the
prominent coccinellids found associated with the
aphid during the study were Hippodamia
variegate (Goeze, 1777), Oenopia
conglobata (Linnaeus, 1758), Priscibrumus
uropygialis (Mulsant,
1853) and Adalia tetraspilota
(Hope, 1831). Several un-identified
parasitoids, syrphids, Vespa
sp. were also seen frequently associated with the nymphs and adults of the
alien species (Image 19–21).
The aphid was also seen predominately attended by Formica sanguinea Latreille, 1798 for
its honey dew.
References
Abd-Rabou, S. (2008). Mass production,
releasing and evaluation of the Lady Beetle, Coccinella
undecimpunctata (Coleoptera:
Coccinellidae), for control of aphids in Egypt.
Archives of Phytopathology and Plant Protection 41:
187–197.
Agarwala, B.K. & A.K. Ghosh (1984). A checklist of Aphididae
of India. Records of Zoological Survey of India,
Occasional Paper 50: 1–71.
Aphid Species File (2016). http://aphid.speciesfile.org/Common/basic/Taxa.aspx?
Taxon Name ID = 1159459 accessed on 7 May 2016.
Archangelsky, P.P. (1917). On the biology of Pterochloroides persicae Chol. Reports of Turkestan
Entomological Station, Tashkent, 70pp.
Ateyyat, M.A. (2008). Evaluation of some insecticides
against the Peach Trunk Aphid, Pterochloroides
persicae (Cholodkovsky)
(Homoptera: Lachnidae), on
peach in Jordan. IOBC/WPRS Bulletin 37: 19–27.
Ateyyat, M.A. & M.S. Abu-Darwish (2009). Short communication. Insecticidal activity of
different extracts of Rhamnus dispermus (Rhamnaceae)
against Peach Trunk Aphid, Pterochloroides persicae (Homoptera: Lachnidae). Spanish Journal of Agricultural
Research 7(1): 160–164.
Bhagat, R.C. (1986a). Aphid pests of fruit
trees and their natural enemies in Kashmir Valley, India. Indian Journal Agricultural Sciences
56(7) 532–534.
Bhagat, R.C. (1986b). On aphid pests of fruit
trees and their natural enemies in Kashmir Valley, India. Indian Agriculturist 30(3) 229–235.
Bindra, O.S. & D.R.C. Bakhetia (1970). The chemical control of the Peach
Stem Aphid, Pterochlorus persicae (Cholodkovsky).
Journal of Research, Punjab Agricultural University 7:
48–51.
Blackman, R.L. & V.F. Eastrop (1994). Aphids on the WorldÕs Trees.
An Identification and Information Guide. CABI,
Wallingford, UK, 476pp.
Bounfour, M., F. Jebbour & J. Wadjinny (2005). Biological traits of
invasive insect species harmful to Moroccan agriculture, pp. 95–100.
In: Plant protection and plant health in Europe: introduction and spread of
invasive species, held at Humboldt University, Berlin, Germany.
Braham, M., P. Edison & N. Ncira (2007). Efficacy of koalin, spinosad and malathion
against Ceratitis capitata
in Citrus orchards. Bulletin on Insectology
60(1): 39–47.
Chan, C.K., A.R. Forbes & A. Raworth
(1991). Aphid-transmitted
viruses and their vectors of the World. Agricultural
Canada Research Branch Technical Bulletin, 216pp.
Ciampolini, M. & M. Martelli (1980). Appearance in Italy of the Peach
Trunk Aphid Pterochloroides persicae (Cholodk.). Bollettino di Zoologia Agraria
e di Bachicoltura 14: 189–196.
Ciampolini, M., I. Farnesi & A. Capella (1997). The spread of the woody
aphid of Prunus (Pterochloroides
persicae). Informatore Agrario 53: 105–107.
Darwish, E.T.E., M.B. Attia
& M.O. Kolaib (1989). Biology and seasonal activity of
giant Brown Bark Aphid Pterochloroides persicae (Cholodk.) on peach
trees in Egypt. Journal of Applied Entomology
107: 530–533.
El-Salam, S.A.A. (2001). Toxicity and
biochemical effects of some safe alternative materials against Pterochloroides persicae
Chlod. Individuals (Order Homoptera: Fam. Aphididae) on
peach trees. Egyptian Journal of Agricultural
Research 79: 1387–1397.
Ghosh, A.K. & L.K. Ghosh (2006). The Fauna of India and Adjacent Countries: Homoptera, Aphidoidea. Zoological Survey of India, Calcutta, 244pp.
Ghosh, L.K. & R. Singh
(2000). Biodiversity of
Indian Insects with special reference to Aphids (Homoptera:
Aphididae). Journal of Aphidology 14: 113–123.
Halkett, F., R. Harrington, M. Hulle«, P. Kindlmann, F. Menu, C.
Rispe & M. Plantegenest
(2004). Dynamics of
production of sexual forms in Aphids: theoretical and experimental evidence for
adaptive Òcoin-flippingÓ plasticity. The American
Naturalist 163(6): 113–125.
Hondru, N., G. Mărgărit
& I. Popa (1986). A new aphid pest of fruit orchards
- Pterochloroides persicae
Cholodkovsky (Homoptera - Aphidina - Lachnidae). Analele
Institutului de Cercetări
pentru Protecţia Plantelor 19: 151–154.
Kairo, M.T.K. & M.A. Poswal (1995). The brown peach aphid, Pterochloroides
persicae (Lachninae: Aphididae): prospects for IPM with particular emphasis on
classical biological control. Biocontrol
News and Information 16: 41–47.
Khan, A.N., I.A. Khan, M.A. Poswal & S. Ahmad (1998). Mass rearing/culturing of brown peach
aphid, Pterochloroides persicae
(Kholodkvoskii) (Lachninae:Aphididae) under laboratory condition. Sarhad
Journal of Agriculture 14:355–368.
Knowledge Bank-Plantwise
(2016). http://www.Plantwise.org/KnowledgeBank/Map/GLOBAL/Pterochloroides_persicae/
accessed on 10 May 2016.
Liotta, G. & G. Maniglia (1993). Variations in infestations of the
almond tree in Sicily in the last fifty years. Acta Horticulturae
373: 277–285.
Mann, G.S., D. Singh & A.S. Dhatt
(1979). Chemical
control of Pterochloroides persicae (Cholod.), a
sporadic pest of peach and almond in Punjab. Indian
Journal of Agricultural Sciences 49: 895–897.
McGavin, G.C. (1993). Bugs of the World. Infobase
Publishing, 550pp.
Mdellel, L. & B.M. Kamel (2012). Prey
consumption efficiency and fecundity of the Ladybird Beetle Coccinella
algerica Kovˆr (Coleoptera: Coccinellidae)
feeding on the Giant Brown-bark Aphid, Pterochloroides
persicae (Cholodkovsky)
(Hemiptera: Lachninae). African Entomology 20: 292–299.
Mirabzadeh, A., A. Sahragard
& M. Azma (1998). Functional response of Chrysoperla carnea Steph. larvae to Pterochloroides persicae,
Uroleucon (Urol.) cichorii
and Aphis craccivora. Seed and Plant
14: 29–34.
Piper, R. (2007). Extraordinary Animals: An Encyclopedia of Curious and Unusual Animals. Greenwood
Press. 6–9pp.
Rakhshani, E., A.A. Talebi,
P. Stary, S. Manzari &
A. Rezwani (2005). Re-description and biocontrol
information of Pauesia antennata
(Mukerji) (Hym., Braconidae, Aphidiinae),
parasitoid of Pterochloroides
persicae (Chol.) (Hom., Aphidoidea,
Lachnidae). Journal of the
Entomological Research Society 7(3): 59–69.
Remaudiere, G. & M. Remaudiere (1997). Catalogue of the WorldÕs Aphididae. Paris, 426pp.
Robert, Y. & D. Bourdin
(2001). Aphid
transmission of potato viruses, pp. 195–225. In: Loebenstein, G., P.H. Berger, A.A. Brunt & R.H. Lawson
(eds.). Virus and Virus-Like Diseases of Potatoes and
Production of Seed- Potatoes. Kluwer, Dordrecht, Netherlands.
Sandhu, G.S. & A.S. Sohi (1978). High-volume and ultra-low-volume
spraying for the control of peach stem-aphid in Punjab. Pesticides 12: 15–17.
Stoetzel, M.B. (1994). Aphids (Homoptera: Aphididae) of potential importance on Citrus in the United
States with illustrated keys to species. Proceedings of
Entomological Society Washington 96: 74–90.
Talhouk, A.S. (1977). Contribution to the knowledge of
almond pests in East Mediterranean countries. VI. The sap-sucking pests.
Zeitschrift fur Angewandte
Entomologie 83: 248–257.
Tsinovskii, Y.P. & K.Y. Egina (1972). The use of Entomophthora
fungi in the control of aphids, pp. 73–94.
In. The pathology of insects and mites: Patalogiya nasekomykh i kleshchei.
UC IPM Home page (2016). http://www.ipm.ucdavis.edu/ accessed on
10 May 2017.
Velimirovic, V.A. (1977). Contribution to the study of peach
aphid Pterochloroides persicae.
Zastita Bilja 28(1): 3–7.
Verma, K.D. & S.M. Das
(1992). The Aphididae of north-west India
(with special reference to Aphids of Jammu and Kashmir State). Ashish
Publication House, New Delhi, i–viii+171pp.
Wieczorek, K., M. Kanturski & L. Junkiert
(2013). The
sexuales of Giant Black-bark Aphid, Pterochloroides persicae
(Hemiptera, Aphidoidea: Lachninae). Zootaxa 3626(1): 094–098.