Philipp Ent 33 (2): 171-184 ISSN 0048-3753 July-December 2019

DETECTION OF THE FALL ARMYWORM, Spodoptera frugiperda

(J.E. Smith) (LEPIDOPTERA: NOCTUIDAE) USING LARVAL
MORPHOLOGICAL CHARACTERS, AND OBSERVATIONS
ON ITS CURRENT LOCAL DISTRIBUTION
IN THE PHILIPPINES

Mario V. Navasero1*, Marcela M. Navasero1, Gideon Aries S. Burgonio1,

Karen P. Ardez1, Melvin D. Ebuenga1, Marie Joy B. Beltran1, Maeden B.
Bato1, Pablito G. Gonzales1, Gil L. Magsino1, Barbara L. Caoili2,
Aimee Lynn A. Barrion-Dupo3, and Minda Flor G.M. Aquino4*

1National Crop Protection Center, College of Agriculture and Food Science (CAFS), University
of the Philippines Los Baños (UPLB), College, Laguna 4031 Philippines
2Institute of Weed Science, Entomology and Plant Pathology, CAFS, UPLB, College, Laguna

4031 Philippines
3Institute of Biological Sciences, College of Arts and Sciences, UPLB, College, Laguna 4031

Philippines
4Department of Agriculture-Regional Crop Protection Center II, Ilagan, Isabela, Philippines

*Corresponding authors: marnavasero@[Link]; da_cvialrcpc02@[Link]

ABSTRACT

Detection surveys and identification using larval morphological

characters of the fall armyworm, Spodoptera frugiperda (J.E. Smith), were
conducted in several provinces in the Philippines from June to October
2019. The smooth body texture and presence of conspicuous black
pinaculae on the dorsal body segments of larvae distinguished S.
frigiperda from the other species of the genus and three other common
noctuid pests of corn in the Philippines, namely; S. exempta (Walker), S.
exigua (Hübner), S litura (Fabricius), S. mauritia Boisduval, Helicoverpa
armigera (Hübner), Mythimna separata (Walker), and Chrysodeixis
eriosoma (Doubleday). The markings along the inverted Y-shaped fronto-
clypeal suture and ecdysial line, except for C. eriosoma, is common to all
species but differ in breadth and color. Incidence of fall armyworm was
recorded in 17 municipalities in 10 provinces. Preliminary observations
indicated that the pest is damaging only to non-Bt corn. It infests all
growth stages of corn but showed apparent preference for earlier growth
stages of the host.

Key words: detection survey, fall armyworm, insect migration, invasive species,
Spodoptera frugiperda

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INTRODUCTION

Currently, the fall armyworm (FAW), Spodoptera frugiperda (J.E. Smith), is

plaguing corn across Africa and Asia. FAW is native to the Americas (Greathead
& Greathead, 1992), distributed from Uruguay and Northern Argentina to
Florida and Texas in southern United States, including Central America and the
Caribbean Islands (Sparks, 1979; Ashley et al., 1989; Nagoshi & Meagher, 2014).
It migrates from the American tropical to semi-tropical regions to the temperate
states in North and South America during summer season (Jeger et al., 2017).
The maximum migration flight recorded for FAW was 1,600 km (Rose et al., 1975
as cited by Huesing et al., 2018). In spite of its migratory behavior, it was kept
confined in its native range by the vast Atlantic Ocean on the east and the Pacific
Ocean on the west, the expanses of which are beyond the FAW dispersal flight
range. However, FAW was introduced to Africa in 2016 (Goergen et al., 2016;
Nagoshi et al., 2018) presumably by trade as it has been intercepted at
quarantine points in Africa (Huesing et al., 2018), indicating that it was able to
hurdle one of those natural barriers, thus enabling it to further expand its
geographic range. In Europe, there were also reported interceptions of larvae
carried by air-freight on vegetables, fruits or herbaceous ornamentals from the
New World (Seymour et al., 1985 as cited in CABI, 2019).
FAW was detected in Yemen (FAO, 2018a; IPPC, 2019a) and India (Ganiger
et al., 2018; IITA, 2018; Sharanabassapa et al., 2018) in mid-2018 and quickly
spread in the latter by the end of the year (EPPO, 2019). The population of FAW
introduced to India in 2018 was presumably from Africa, or from the same
source as in Africa, evidence of which was provided by Nagoshi et al. (2019)
showing genetic homogeneity between the South African and Indian FAW
populations. Since its incursion to India, it rapidly spread eastward by the end of
2018 to Bangladesh (Palma, 2018; FAO 2019a), Myanmar (FAO, 2019a; IPPC,
2019b), Thailand (FAO, 2018b), and Laos (FAO, 2019a & b). The most recent
record of countries invaded by FAW in 2019 includes Malaysia (Anonymous,
2019a) and China (IPPC, 2019c; Shrikanth, 2019) in January; Indonesia (FAO
Indonesia, 2019; IIPC, 2019d) and Vietnam (FAO 2019d; Vietnam News, 2019) in
March; Egypt (Anonymous, 2019b), Cambodia (Kunthear, 2019), Taiwan
(Charlier, 2019), and South Korea (FAO, 2019e; IIPC 2019e) in June; and Nepal
(Guragain, 2019) and Japan (Anonymous, 2019c; IIPC, 2019f) in July. Using the
Species Distribution Models to predict the geographic expansion of the
distribution of FAW, Early et al. (2018) identified the Philippines as one of the
countries facing high risk of possible invasion. S. frugiperda is not included in
the earlier listings of pests of corn in the Philippines (Gabriel, 2000; Caasi-Lit,
2015).
The Philippines had experienced several outbreaks of native and invasive
species in recent years. Two of these involved armyworm species under the
genus Spodoptera. First was the black armyworm, S. exempta (Walker), in 2010
that brought considerable damage to corn, rice, and sugarcane in Southern
Tagalog provinces and in the province of Bataan (Navasero et al., 2011; Aguilon
et al., 2015). Second was the onion or beet armyworm, S. exigua (Hübner) in
2016 (Navasero, 2017; Navasero et al., 2019), which up to the present, remains

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the main pest problem in onion production in mainland Luzon. The latter also
attacks corn in the early growth stages of the host. Since, three of the primary
host plants of S. frugiperda, namely; corn, rice, and sugarcane, are major
economically important crops in the country, it poses a big threat to the
Philippine economy.
This paper reports the early detection of FAW in 10 provinces and
identification of other incidence areas in the Philippines including preliminary
observations on its field biology, nature of damage and natural enemies.

MATERIALS AND METHODS

Collection
Detection surveys were conducted in 14 Philippine provinces from June to
October 2019. Live larvae were collected from various localities in Luzon and
were taken to the laboratory located at the National Crop Protection Center
(NCPC), University of the Philippines Los Baños (UPLB), in Laguna Province. Few
of them were preserved in 85% ethyl alcohol as reference samples and others
were reared on fresh leaves of open pollinated corn up to late instars and adults.
Larvae collected from the Visayas and Mindanao were directly preserved in 85%
ethyl alcohol.

Identification of FAW.
Based on pertinent literature (Levy & Habeck, 1976; King & Saunders,
1984; Huesing et al., 2018), larvae found in corn in the areas visited, that
possess morphological characteristics distinctive of FAW were collected. Larvae
of FAW were also compared to larvae of other species of Spodoptera and other
noctuids associated with corn.
Representative adult moths ensuing from field collected larvae of FAW were
preserved as voucher specimens kept at the NCPC, UPLB. Some were deposited
at the Entomology Section of the UPLB Museum of Natural History, College,
Laguna (Accession numbers UPLBMNH LEP-04643 to 04676).

Nature of damage, type, and growth stages of corn attacked

Characteristic signs of damage by FAW observed in the field were noted.
Similarly, type of corn (traditional or genetically modified) and growth stages of
corn present in incidence sites were also recorded.

RESULTS AND DISCUSSION

Identification of FAW
It has been established that FAW can be identified even in its larval stage
(Levy & Habeck, 1976; King & Saunders, 1984; Huesing et al., 2018). Easily
recognizable characters are the presence on the dorsum within the pale-colored
area, of four, conspicuous, raised, black pinaculae or “spots” bearing the long

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primary dorsal setae as follows: arranged in a line on the 2nd and 3rd thoracic
segments, arranged in trapezoid on the first seven abdominal segments, and
arranged in a square on the second from last abdominal segment (Figure 1). The
arrangement of primary dorsal setae is the same among other local species of
noctuids associated with corn, namely: Spodoptera exempta (Walker), S. exigua
(Hübner), S litura (Fabricius), S. mauritia Boisduval, Helicoverpa armigera
(Hübner), Mythimna separata (Walker), and Chrysodeixis eriosoma (Doubleday),
however, conspicuous pinaculae are absent (Figure 2). Only some larvae of H.
armigera, particularly the dark-colored form (Figure 2f), may possess similar
spots but can be distinguished from FAW by the rough and bumpy texture of the
body in contrast to the smooth texture in the latter. The inverted Y-shaped fronto
-clypeal suture (branches) and ecdysial line (base) is also present in all the
species mentioned but differ in the extent and breadth of color bands to totally
absent in C. eriosoma (Figure 3)

Figure 1. Distinguishing morphological features of the fall armyworm, Spodoptera

frugiperda. a. spots on dorsum. b. inverted Y-shaped yellowish band
along the fronto-clypeal suture and the ecdysial line. c. close-up of the
spots arranged in a square on the second to the last abdominal
segment.

Figure 2. Larvae of noctuids attacking corn in the Philippines. a. Spodoptera

exempta. b. S. exigua. c. S. litura. d. S. mauritia. e. Mythimna separata.
f. Helicoverpa armigera. g. Chrysodeixis eriosoma.
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Figure 3. Heads of larvae of noctuids attacking corn in the Philippines. a.

Spodoptera exempta. b. S. exigua. c. S. litura. d. S. mauritia. e.
Mythimna separata. f. Helicoverpa armigera. g. Chrysodeixis
eriosoma.
Detection and identified incidence areas of FAW
FAW was collected in 10 of the 14 provinces visited. Identified incidence
areas are listed in Table 1 and indicated on its local distribution map (Figure 4).
The presence of FAW was first recorded in Piat, Cagayan. Three larvae of FAW
were collected on June 7, 2019 from Bt corn in mid-whorl stage. This was also
the first photo-documentation of a larva that was then suspected as S.
frugiperda. One of those specimens was among the samples of noctuid larvae
sent to the NCPC and was the very first to have been morphologically examined.
Considering there has been no previous record yet of S. frugiperda in the
Philippines (Gabriel, 2000; Caasi-Lit, 2015), this became the basis of the initial
and unpublished report on the occurrence of the FAW in the country (Navasero
& Magsino, 2019).
It was in Gonzaga, Cagayan where more than 100 FAW larvae were
collected on two occasions, i.e., July 20 and 25, 2019. Different growth stages
of open-pollinated and Bt corn were present in the area but FAW larvae were
only collected from open-pollinated corn. Most of the larvae were collected in
early to mid-whorl growth stages and were estimated to be in their 3rd to 4th
instars. Only a few late instar larvae were collected from corn in the late-whorl
stage and none was collected from those in the reproductive stages. Infestation
level was estimated at 30% in the affected fields. In Solana, also in Cagayan
province, the presence of FAW larvae in one field of open-pollinated corn at
tasseling stage was detected on July 25, 2016.
The few individuals of FAW found in Piat, Cagayan in Bt corn field were
probably from the refuge plants and should not be interpreted as an indication of
resistance to Bt corn. Follow-up visit in Piat three weeks from the date of first
detection did not result in further collection of additional specimens. So far, the
genetic marker linked with resistance to the Cry1Fa toxin expressed in Bt corn
was not found in Togo, one of the first territories in Africa where FAW has been
detected (Huesing et al., 2018). In addition, field performance of single-gene Bt
corn further suggests that Bt resistance alleles may not be present in the FAW
populations currently in Africa (Huesing et al., 2018). One of the authors of this
paper (G.L. Magsino), during his visit to Vietnam in July 2019 had provided
firsthand evidence of how open-pollinated corn varieties were devastated by FAW
but not the Bt corn products with two modes of action.
In Gonzaga and Solana, Cagayan, detection of FAW were all from open
pollinated corn. Predominance of FAW in mid-whorl to late whorl among the
different growth stages of corn present in the collection site in Gonzaga could be
an indication that the pest prefers the early growth stages. Similarly, in Ilocos

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Table 1. Sites where fall armyworm, Spodoptera frugiperda, was collected/

documented.
Province/Town/ Date
Coordinates Collector(s)
Barangay (2019)
Cagayan: Piat: 17°50'8.62"N, 07-Jun M.F.G.M. Aquino & E.S. Ilorea
Minanga 121°29’28.21"E
Cagayan: Gonzaga: 18°17'30.10"N, 20-Jul M.F.G.M. Aquino & E.S. Ilorea
Patel 122°3’33.25"E
Cagayan: Gonzaga: 18°17'30.10"N, 24-Jul R.P. Arapan, M.V. Atienza, G.B.
Patel 122°3’33.25"E Katimbang, M.M. Navasero &
M.V. Navasero
Cagayan: Gonzaga: 18°17'32.03"N, 24-Jul R.P. Arapan, M.V. Atienza, G.B.
Migrafil 122°3’32.15"E Katimbang, M.M. Navasero &
M.V. Navasero
Cagayan: Solana: 17°44’52.53"N, 25-Jul M.F.G.M. Aquino
Dassun 121°42'33.81"E
Ilocos Norte: Laoag: 44 18°11’58.10"N, 25-Jul Ardez, M.J.B. Beltran, G.A.S.
-Zamboanga 120°33’6.93"E Burgonio, K.P. & R.P. Urriza
Ilocos Norte: Laoag: La 18°11’18.82"N, 25-Jul Ardez, M.J.B. Beltran, G.A.S.
Consolacion 120°36’14.34"E Burgonio, K.P. & R.P. Urriza
Nueva Ecija: San Jose 15°47’4.39"N, 30-Jul M.T. Lontoc & N.M. Navasero
City: Palestina 121°0’22.99"E
Nueva Ecija: San Jose 15°47’2.75"N, 30-Jul M.T. Lontoc & N.M. Navasero
City: Palestina 121°0’24.10"E
Batangas: San Juan: 13°51’25.17"N, 28-Aug R.P. Arapan, K.P. Ardez, M.V.
Janao-Janao 121°21’53.42"E Atienza & G.A.S. Burgonio
Quezon: Candelaria: 13°54’39.54"N, 28-Aug R.P. Arapan, K.P. Ardez, M.V.
Malabanan 121°25’56.5"E Atienza & G.A.S. Burgonio
Quezon: Tiaong: Kipot 13°51’14.13"N, 02-Sep M.V. Atienza, G.B. Katimbang,
121°19’5.24"E M.M. Navasero & M.V.
Navasero
Quezon: Tiaong: 13°56’38.39"N, 02-Sep M.V. Atienza, G.B. Katimbang,
Ayusan 2 121°17’41.76"E M.M. Navasero & M.V.
Navasero
Laguna: Bay: Tranca 14°8’40.27"N, 10-Sep G.B. Katimbang
121°15’32.94"E
Laguna: Los Baños: 14°0’6.03"N, 121° 20-Sep M.V. Atienza & G.B. Katimbang
Batong Malake 0’9.14"E
Laguna: Nagcarlan: 14°0’6.03"N, 121° 27-Sep M.T. Lontoc & N.M. Navasero
Wakat 0’9.14"E
Zamboanga del Norte: 8°30’29.28"N, 17-Sep G.A.S. Burgonio, M.D. Ebuenga
Katipunan: Daang 123°18’20.48"E & M.V. Navasero
Lungsod
Zamboanga del Sur: 7°4’58.68"N, 122° 17-Sep G.A.S. Burgonio, M.D. Ebuenga
Zamboanga City: 13’15.2"E & M.V. Navasero
Sangali
South Cotabato: 6°11’55.3"N, 125° 19-Sep G.A.S. Burgonio, M.D. Ebuenga
Polomolok: Magsaysay 3’15.79"E, & M.V. Navasero
South Cotabato: 06°09’33.02"N, 19-Sep M.D. Ebuenga & M.V. Navasero
General Santos City: 125°12’18.57"E
Katanggawan
Leyte: Baybay: 10°40’58.04"N, 11-Oct R.P. Arapan, M.V. Atienza, G.P.
Banawan 124°48’48.58"E Cueto, T. Reyes & R.P. Urriza
Leyte: Sta. Fe: Gapas 11°12’56.90"N, 11-Oct R.P. Arapan, M.V. Atienza, G.P.
124°55’27.60"E Cueto, T. Reyes & R.P. Urriza

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Figure 4. Incidence sites of Spodoptera frugiperda in the Philippines. (Generated

using QGIS Development Team. 2016. QGIS Geographic Information
System. Open Source Geospatial Foundation. URL [Link]

Norte, Zamboanga del Norte and Sur, and South Cotabato (Polomolok), FAW
larvae were collected from open-pollinated corn in mid- to late-whorl stages. In
South Cotabato (General Santos City), 2nd to 3rd instar larvae were found on 12-
day old seedlings. At about 6-7 days from egg-laying to 2nd or 3rd instar,
oviposition had occurred at only 2-3 days after seedling emergence.
In the African continent, aside from South Africa, corn plants being
cultivated are open-pollinated varieties, the annual loss on production was
estimated to reach 21-53% (Abrahams et al., 2017). However, area-wide loss in
percent production projected in Africa may not apply in the Philippines
considering that Bt corn accounts for about 50% of the total cultivated area for
the crop. To date, about three months after it was first detected in Cagayan
Valley, incidence of FAW remained sporadic and far from reaching outbreak
level. It is projected that FAW may not become a major problem in Cagayan
Valley where transgenic corn is 90% of the total corn plantings. However, among

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the incidence areas, infestations of more than 70% were observed in

municipalities of Candelaria and Tiaong in Quezon, and the municipality of San
Juan in Batangas, where Bt corn planting is not yet extensive. Sample of
damages of FAW on different growth stages of corn documented from the
different sites are shown in Figure 5.

Figure 5. Damage of Spodoptera frugiperda on different growth stages of

corn. a. high infestation in a field 12 days after sowing. b. early
whorl stage. c. mid-whorl stage. d. on ear. e. on tassel.

New associations may arise between the newly arrived alien fall armyworm
and the natural enemies of other species of Spodoptera in the Philippines (i.e., S.
litura, S. exigua, S. exempta, and S. mauritia). Mermithid nematodes were
recovered from FAW larvae in both Cagayan and Ilocos Norte, and parasitoids (a
braconid and an ichneumonid) had been reared out from samples also from
Cagayan. A fungus-infected larva was recovered from experimental fields in the
University of the Philippines Los Baños, Laguna. Such new associations involving
more species had been documented in Africa (Cruz et al., 2018; Sisay et al.,
2018; Kenis et al., 2019) and in India (Shylesa et al., 2018; Chormule et al.,
2019) which have their own native species of Spodoptera. Continuous inventory
of natural enemies of FAW must be conducted and their impact on the FAW
population must be determined.
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The FAW spreads rapidly once introduced in an area. In China, it was first
reported in January 2019 (FAO, 2019b) and by June, it had spread to half of
China’s provinces (Shrikanth, 2019). In the case of Taiwan, it was first detected
on June 10, 2019 in Miaoli Country but a month later it was already present in
17 out of 22 administrative areas (Wang & Wu, 2019). Attempts on pre-emptive
elimination or limiting its spread from incidence areas to other places in the
Philippines through quarantine measures may not be feasible at all, and as
mentioned, its distribution based on the results of surveys (Figure 4) shows its
presence even in the Visayas and Mindanao. At the rate that it is being detected,
and considering its migratory behavior, FAW may have already spread
throughout the country. The prevailing tropical conditions and availability of
host plants in the Philippines may be favorable for FAW. Hence, it is expected
that the pest will continue to persist. Efforts should focus on managing the
populations of FAW to prevent them from attaining outbreak levels.
FAW is a polyphagous and voracious pest in its native range and in
countries it has invaded. It is expected that its introduction into the country may
also be a problem in crops other than corn, like rice, sugarcane and other
vegetables like onion, which are reported hosts of FAW (Pashley, 1986; Lu &
Adang, 1996 as cited by Levy et al., 2002; Montezano et al., 2018). Its possible
attack on onion should be another big concern. Aside from onion, FAW may also
become a problem on sugarcane but to a lesser extent. In India, Chormule et al.
(2019) found an infestation of less than 5% on sugarcane.
This is the first report of the fall armyworm in the Philippines. The pest can
be managed with available tools such as insecticides, Bt corn with functionally
effective modes of action, identification of and augmentation with natural
enemies, and clean cultivation practices.

ACKNOWLEDGEMENT

This initiative highlights public-private partnership in addressing the FAW

problem. We would like to thank the following individuals and their institutions
for the invaluable contribution primarily in the collection of FAW specimens.
First, the NCPC staff for assistance in collecting and rearing FAW larvae – G.B.
Katimbang, M.V. Atienza, M.N. De Panis-Javier, R.N. Candano, R.P. Arapan, R.P.
Urriza, and T.B. Reyes. Second, our partners from local government units – Z.V.
Amargo, F.P. Valdez W. Carandang of Tiaong, Quezon; Municipal Agriculturist
F.L. Baclig of Gonzaga, Cagayan and his staff; Ilocos Norte Provincial Corn
Coordinator, J.L. Macoco, and Laoag City Agriculture Office, Cherry P.B.
Joaquin. Third, the staff of regional field office of RCPC II and field monitor,
Edmar S. Ilorea; RCPC IX Center Chief P.C. Jover; Brgy. Chair V.F. Pasilan of
Magsaysay, Polomolok, South Cotabato. Fourth, our partners from the industry -
Jedeliza B. Ferrater of East-West Seeds; Mark T. Lontoc and Nelson M. Navasero
of Eurofins Agrosciences; Corteva Agrisciences, for sponsoring the very first
activity on FAW awareness campaign in March 2019 facilitated by its former
staff, Henry B. Dupo, Jr. Lastly, the Department of Agriculture-Bureau of
Agricultural Research for funding NCPC’s quick response initiatives on new and
emerging crop pests.
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Volume 33 July-December 2019 Number 2

TABLE OF CONTENTS

Marita D. Salinas-Labe, Augusto C. Sumalde,

Leonila A. Corpuz-Raros, and Virginia R. Ocampo
Biology of a Spider Mite, Tetranychus urticae Koch and a Phytoseiid Predator,
Proprioseiopsis lenis Corpuz & Rimando and its Biological Control Potential .............. 97-111

Jay-R M. Millanes and Pio A. Javier

Biology of the Predatory Water Bug, Diplonychus rusticus (Fabricius)
(Hemiptera: Belostomatidae), on Prey Aedes aegypti L. (Diptera: Culicidae)
Wrigglers ..................................................................................................................... 113-128

Merdelyn T. Caasi-Lit, Mark Bryan T. Lontoc, and Edwin A. Benigno

Efficacy of Corn Event TC1507 and the Hybrid Stacked Product
TC1507xMON810xNK603 Against Secondary Lepidopteran Pests
of Corn in the Philippines ............................................................................................. 129-141

Maria Dulce J. Mostoles, Aimee Lynn A. Barrion-Dupo,

Maria Amabel A. Capricho, and Alberto T. Barrion
Mark-Recapture Method: A New Dimension in the Management of the
Rice Black Bug, Scotinophara sorsogonensis Barrion et al. (Hemiptera:
Pentatomidae) in the Bicol Region, Philippines ........................................................... 143-158

Aaron V. Ogot, Romnick A. Latina, and Barbara L. Caoili

Molecular Identification and Preliminary Genetic Diversity Analysis of Tobacco
Stem Borer, Scrobipalpa aptatella (Walker) (Lepidoptera: Gelechiidae) from
Misamis Oriental and Bukidnon using Cytochrome c Oxidase Subunit I Gene ……… 159-170

Mario V. Navasero, Marcela M. Navasero, Gideon Aries S. Burgonio,

Karen P. Ardez, Melvin D. Ebuenga, Marie Joy B. Beltran, Maeden B. Bato,
Pablito G. Gonzales, Gil L. Magsino, Barbara L. Caoili,
Aimee Lynn A. Barrion-Dupo, and Minda Flor G.M. Aquino
Detection of the Fall Armyworm, Spodoptera frugiperda (J.E. Smith) (Lepidoptera:
Noctuidae) Using Larval Morphological Characters, and Observations on its
Current Local Distribution in the Philippines ………...............……………………....….. 171-183

Abstracts of Papers Presented at the Scientific Sessions of the Philippine

Association of Entomologists, Inc., held during the 51st Anniversary and
Annual Scientific Conference of the Pest Management Council of the
Philippines, Inc., 2-5 July, 2019, Coron Westown Resort, Coron, Palawan ………….. 185-214

Reviewers for Volume 33 …………………………………....…….….….….….….…..….. 215-216

Author Index .….….….…..……….….….….…..…….….….….…..…….….….….…..….. 217-219

Subject Index .….….….………….….….…….….….….…..…….….….….…..….…..…... 220-224