a new genus of mite (acari acaridae) phoretic on bees (ctenocolletes)
a new genus of mite (acari acaridae) phoretic on bees (ctenocolletes)
a new genus of mite (acari acaridae) phoretic on bees (ctenocolletes)
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Rec, West, A"st, ;'VI"", 1984.11 (2): 77·86<br />
A New Genus <str<strong>on</strong>g>of</str<strong>on</strong>g> Mite (Acari: Acaridae) Phoretic<br />
<strong>on</strong> Bees (Ctellocolletes) in Australia<br />
A.Fain*<br />
Abstract<br />
A <str<strong>on</strong>g>new</str<strong>on</strong>g> <str<strong>on</strong>g>genus</str<strong>on</strong>g>, Ctenocolletacarus, and three <str<strong>on</strong>g>new</str<strong>on</strong>g> species, C. [<strong>on</strong>g£rostr£s, C. grand£or<br />
and C. brev£rostr£s, are described from hypopial nymphs from three species <str<strong>on</strong>g>of</str<strong>on</strong>g> <strong>bees</strong><br />
(Ctenoco[[etes, Stenotritidae) from Western Australia.<br />
Introducti<strong>on</strong><br />
The <str<strong>on</strong>g>new</str<strong>on</strong>g> <str<strong>on</strong>g>mite</str<strong>on</strong>g>s described herein are known <strong>on</strong>ly from hypopial nymphs obtaincd<br />
by T.F. Houst<strong>on</strong>, Westcrn Australian Museum, Perth (WAM), from adult <strong>bees</strong><br />
(Ctenocolletes, Stenotritidac). Most were secreted in the abdominal tergal<br />
pouchcs <str<strong>on</strong>g>of</str<strong>on</strong>g> females, but some occurred under the cdges <str<strong>on</strong>g>of</str<strong>on</strong>g> the abdominal terga<br />
and <strong>on</strong> other parts <str<strong>on</strong>g>of</str<strong>on</strong>g> thc bodies <str<strong>on</strong>g>of</str<strong>on</strong>g> male and female <strong>bees</strong> (Houst<strong>on</strong> in press).<br />
The holotypes <str<strong>on</strong>g>of</str<strong>on</strong>g> the <str<strong>on</strong>g>new</str<strong>on</strong>g> species are deposited in the WAM. Paratypes are also<br />
deposited there, in thc Institut royal des Sciences naturelles de Belgique, Bruxelles<br />
(IRSNB) and in the British Museum (Natural History), L<strong>on</strong>d<strong>on</strong> (Bl\INH).<br />
The host <strong>bees</strong> are in the WAM and the Western Australian Department <str<strong>on</strong>g>of</str<strong>on</strong>g><br />
Agriculture, Perth (WADA).<br />
Type Species<br />
Ctenoco[[etacarus [<strong>on</strong>g£rostr£s sp. novo<br />
Systematics<br />
Acari: Acaridae<br />
Genus Ctenocolletacarus gen. novo<br />
Diagnosis<br />
With characters <str<strong>on</strong>g>of</str<strong>on</strong>g> hypopial nymphs <str<strong>on</strong>g>of</str<strong>on</strong>g> the Acaridae, but differing from all<br />
other genera <str<strong>on</strong>g>of</str<strong>on</strong>g> this family by prol<strong>on</strong>gati<strong>on</strong> <str<strong>on</strong>g>of</str<strong>on</strong>g> antero-dorsal extremity <str<strong>on</strong>g>of</str<strong>on</strong>g><br />
idiosoma into a well-developed rostrum. Resembling Caloglyphus Berlese, 1923<br />
(= ?Sancassania Oudemans, 1916) and Lackerbaueria Zachvatkin, 1941 but differing<br />
by presence <str<strong>on</strong>g>of</str<strong>on</strong>g> a rostrum, its l<strong>on</strong>ger sternum (together with epimeres It)<br />
* Institute <str<strong>on</strong>g>of</str<strong>on</strong>g> Tropical Medicine, 155, Nati<strong>on</strong>alestraat, Antwerp, Belgium.<br />
77
A New Genus <str<strong>on</strong>g>of</str<strong>on</strong>g> Mite Phoretic <strong>on</strong> Bees<br />
almost reaching coxae III and its transversely striated dorsal shields. Differing<br />
from Laekerbauen'a in coxal fields III being separated medially (fused in Laekerbaueria).<br />
Resembling Paraeeroglyphus Fain and Beaucournu, 1973 in epimeres<br />
Ill-IV forming closed coxal fields (narrowly separated medially) but differing<br />
in presence <str<strong>on</strong>g>of</str<strong>on</strong>g> rostrum, l<strong>on</strong>ger sternum, presence <str<strong>on</strong>g>of</str<strong>on</strong>g> 5 foliate setae <strong>on</strong> tarsi IV and<br />
<strong>on</strong>e saucer-like seta <strong>on</strong> tarsi I-Il.<br />
Descripti<strong>on</strong><br />
Body oval. Dorsal surface slightly c<strong>on</strong>vex. Sejugal furrow str<strong>on</strong>g, well forward.<br />
Dorsum <str<strong>on</strong>g>of</str<strong>on</strong>g> body almost completely punctate. Dorsal setae short, either thin or<br />
spinous, except l<strong>on</strong>g piliform Q 5. Oil glands opening between setae hand Q 2.<br />
Venter: Palposoma completely covered by rostrum, l<strong>on</strong>ger than wide, with<br />
anterior margin slightly incised, bearing two solenidia a and two pairs <str<strong>on</strong>g>of</str<strong>on</strong>g> short<br />
lateral setae. Sternum loosely c<strong>on</strong>nected posteriorly with epimeres Il. Two narrow<br />
paramedian poorly sclerotized pregenital sclerites present. Suctorial plate large,<br />
wider than l<strong>on</strong>g. Anterior suckers distinctly smaller than posterior suckers;<br />
c<strong>on</strong>oids approximately <strong>on</strong> same line as posterior suckers (Fain 1973). Legs rather<br />
short and thick, ending in a well-developed curved claw.<br />
Chaetotaxy <str<strong>on</strong>g>of</str<strong>on</strong>g> body: Setae v i spinose; v e short, thinner than v i; s ex very<br />
thin. Other setae present: d 1 to d 5, Q 1 to Q 5;h; sh;ga;gm;gp. Setae ex I,<br />
ex III and g p modified into c<strong>on</strong>oids (Fain 1973).<br />
Legs: tarsi I-Il with 1 l<strong>on</strong>g simple seta and 2 short setae (<strong>on</strong>e thin and <strong>on</strong>e spinelike).<br />
Tarsi III with 7 foliate and 1 piliform setae. Tarsi IV with 5 foliate and 3<br />
unequal piliform setae (1 very l<strong>on</strong>g, 1 l<strong>on</strong>g and 1 short). Tibiae I-Il with 2-2-1-1<br />
setae. Genua 2-2-1-0. Femora 1-1-0-1. Trochanters 1-1-1-0. Solenidia: Tarsus I<br />
with w 1 slightly dilated; w 3 slightly pore apical and tapering at apex: w 2<br />
cylindrical, more basal. Tibiae I-IV with a well-developed solenidi<strong>on</strong>.<br />
Key to Species <str<strong>on</strong>g>of</str<strong>on</strong>g> Ctenocolletacams<br />
1 Idiosoma prol<strong>on</strong>ged anteriorly by a short rostrum.<br />
Most <str<strong>on</strong>g>of</str<strong>on</strong>g> dorsal setae spinose. Dorsal striati<strong>on</strong>s forming<br />
a network<br />
C. brevirostn's sp. novo<br />
Idiosoma prol<strong>on</strong>ged anteriorly by a l<strong>on</strong>g rostrum.<br />
Dorsal setae thin, piliform. Dorsal striati<strong>on</strong>s not<br />
forming a network . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 2<br />
2 Rostrum l<strong>on</strong>g and narrow. Posterior suckers slightly<br />
wider (18-19 I-lm) than anterior suckers (13-15 I-lm).<br />
Idiosoma 340-426 I-lm l<strong>on</strong>g<br />
C. l<strong>on</strong>girostris sp. novo<br />
Rostrum relatively shorter and thicker. Posterior<br />
suckers almost twice as wide (33-36 I-lm) as anterior<br />
suckers (18-20 I-lm). Idiosoma 548-585 I-lm l<strong>on</strong>g<br />
C. grandior sp. novo<br />
78
Holotype<br />
A. Fain<br />
Cte/locolletacanLS IO/lgirostris Sp. nov.<br />
Figures 1-6, I 0<br />
WAM 83/169, hypopus <strong>on</strong> slide, ex tergal pouches <str<strong>on</strong>g>of</str<strong>on</strong>g> Ctenocolletes nichols<strong>on</strong>i 9; 200 km<br />
N <str<strong>on</strong>g>of</str<strong>on</strong>g> Geraldt<strong>on</strong>, Western Australia, 20.viii.1971, T.F. Houst<strong>on</strong>.<br />
I'aratypes<br />
Western Australia<br />
Ctenocolletes centralis; 50 hypopi ex tergal pouches <str<strong>on</strong>g>of</str<strong>on</strong>g> 9 (WAM 82/1890),12.5 km ENE <str<strong>on</strong>g>of</str<strong>on</strong>g><br />
Anketell HS (28°02 's, 118°51 'E), 6-7 .ix.1981, '1'.1". Houst<strong>on</strong> (WAM 32 hypopi, WAM 83/360-2;<br />
IRSNB 10 hypopi; BMNlI 8 hypopi); 12 hypopi from various parts <str<strong>on</strong>g>of</str<strong>on</strong>g> body <str<strong>on</strong>g>of</str<strong>on</strong>g> dd (WAM<br />
82/1893, WADA TFH R15), 4.8 km S <str<strong>on</strong>g>of</str<strong>on</strong>g> Neale Juncti<strong>on</strong> (28°18'S, 125°49'E), 16.vii.1974,<br />
K.T. Richards (WAM 8 hypopi, WAM 83; 1RSNB 2 hypopi; B1\lNII 2 hypopi).<br />
Ctenocolletes nichols<strong>on</strong>i: 35 hypopi, same data as for holotype (WAM 27 hypopi, WA1\1<br />
83/363-8; 1RSNB 8 hypopi); 13 hypopi from 9, 16 km S <str<strong>on</strong>g>of</str<strong>on</strong>g> Nerren Nerren HS (27 0 08'S,<br />
114°38'E), 19.viii.1980, C.A. lIoward and T.F. Houst<strong>on</strong> 329-1 (WAM 8 hypopi, WAM<br />
83/379-82; lRSNB 5 hypopi); 40 hypopi ex tergal pouches <str<strong>on</strong>g>of</str<strong>on</strong>g> 9 (WAM 82/1881), 37 km NE <str<strong>on</strong>g>of</str<strong>on</strong>g><br />
Lavert<strong>on</strong>, 10-12.ix.1982, B. Hanich and '1'.1". Houst<strong>on</strong> (WAM 32 hypopi, WAM 83/369-73;<br />
IRSNB 6 hypopi; BMNII 2 hypopi).<br />
Diagnosis<br />
Distinguished from the two other speCies <str<strong>on</strong>g>of</str<strong>on</strong>g> the <str<strong>on</strong>g>genus</str<strong>on</strong>g> by the l<strong>on</strong>g, narrow<br />
shape <str<strong>on</strong>g>of</str<strong>on</strong>g> the rostrum.<br />
Descripti<strong>on</strong><br />
Holotype 412 pm l<strong>on</strong>g (including rostrum), 249 pm wide (maximum width).<br />
!\Ieasurements in 8 paratypes (in pm): 426 x 246; 420 x 248; 395 x 240; 390 x<br />
240; 380 x 242; 368 x 213; 350 x 210; 340 x 210. Specimens measuring less than<br />
380 pm arc less frequent than those exceeding this length. These small specimens<br />
corresp<strong>on</strong>d closely to the larger <strong>on</strong>es except for size <str<strong>on</strong>g>of</str<strong>on</strong>g> body and <str<strong>on</strong>g>of</str<strong>on</strong>g> some organs<br />
(setae, suckers, etc.). Dorsum: Rostrum 80 pm l<strong>on</strong>g and 27 pm wide in its middle.<br />
Cuticle covering the rostrum raised to form a crest. Setae v i arc thick spines, v e<br />
are smaller and thinner spines. All dorsal setae thin and short except Q :5 125 pm<br />
l<strong>on</strong>g. Venter: Palposoma 30 pm l<strong>on</strong>g, 19 pm wide; 0' 28-30 pm l<strong>on</strong>g. Sternum<br />
87 pm l<strong>on</strong>g. Pregenital sclerites fused anteriorIy with epimeres IV. Suctorial plate<br />
94 pm wide; anterior suckers 13-15 pm wide, posterior suckers 18-19 pm wide;<br />
they are preceded by a flat rounded area. Legs relatively thick and short. Length<br />
<str<strong>on</strong>g>of</str<strong>on</strong>g> tarsi I-IV (in pm): 24-24-27-27. Chaetotaxy <str<strong>on</strong>g>of</str<strong>on</strong>g> legs I-IV: Tarsi I-II with 2<br />
simple setae (<strong>on</strong>c <str<strong>on</strong>g>of</str<strong>on</strong>g> 50 pm and <strong>on</strong>c <str<strong>on</strong>g>of</str<strong>on</strong>g> 12-15 pm) and <strong>on</strong>c spinous seta.<br />
Holotype<br />
Cte/locolletacanls gra/ldior sp. nov.<br />
Ficrures 8-9<br />
"<br />
WAM 83/170, hypopus <strong>on</strong> slide, ex tergal pouches <str<strong>on</strong>g>of</str<strong>on</strong>g> Ctenocolletes centralis 9 (WAM<br />
82/1890); 12.5 km ENE <str<strong>on</strong>g>of</str<strong>on</strong>g> AnketelllIS (28°02'S, 118°51'E), Western Australia, 6-7.ix.1981,<br />
'1'.1". Houst<strong>on</strong>.<br />
79
A New Genus <str<strong>on</strong>g>of</str<strong>on</strong>g> Mite Phoretic <strong>on</strong> Bees<br />
figure 1<br />
Ctenocolletacarus l<strong>on</strong>girostris Sp. novo Hypopus in ventral view.<br />
80
A. Fain<br />
2<br />
Figures 2-5<br />
Ctenocolletacarus l<strong>on</strong>g£rostris sp. novo Hypopus. Tarsus I in dorsal view (2) and<br />
ventral view (3). Tarsi HI (4) and IV (5) in lateral view.<br />
Paratypes<br />
Western Australia<br />
Ctenocolletes centralis: 11 hypopi from various parts <str<strong>on</strong>g>of</str<strong>on</strong>g> body <str<strong>on</strong>g>of</str<strong>on</strong>g> cl (WAM 82/1893); 4.8 km<br />
S <str<strong>on</strong>g>of</str<strong>on</strong>g> Neale Juncti<strong>on</strong> (28°18'S, 125°49'E), 16.vii.1974, K.T. Richards (WAM 6 hypopi, WAM<br />
83/357-9; IRSNB 3 hypopi; BMNH 2 hypopi).<br />
81
A N~w G~nus <str<strong>on</strong>g>of</str<strong>on</strong>g> Mit~ Phor~tic <strong>on</strong> B~es<br />
Ctenocolletes nichols<strong>on</strong>i: 1 hypopus from 9; 16 km 5 <str<strong>on</strong>g>of</str<strong>on</strong>g> Nerren Nerren HS (27°08'5,<br />
114 0 38'E), 19.viii.1980, C.A. Howard and T.F. Houst<strong>on</strong> 329-1 (WAM 83/356).<br />
Diagnosis<br />
Distinguished from C. l<strong>on</strong>girostris by the relatively wider and shorter rostrum,<br />
the much larger idiosoma and the relatively larger posterior suckers.<br />
Descripti<strong>on</strong><br />
Holotype 550 pm l<strong>on</strong>g (including rostrum), 345 pm wide. In 7 paratypes,<br />
measurements (in pm): 585 x 375; 572 x 360; 570 x 365; 569 x 350; 560 x 360;<br />
555 x 345; 548 x 325 (the smallest specimen <str<strong>on</strong>g>of</str<strong>on</strong>g> the collecti<strong>on</strong>). Postero-lateral<br />
margins <str<strong>on</strong>g>of</str<strong>on</strong>g> idiosoma membranous. Dorsum as in C. l<strong>on</strong>girostris. Rostrum<br />
relatively shorter but wider in its posterior half than in that species. Ventral surface<br />
Figure 6<br />
Figure 7<br />
Ctenocolletacarus l<strong>on</strong>girostris sp. novo Hypopus in dorsal view.<br />
Ctenocolletacarus brevirostris sp. novo Hypopus in dorsal view.<br />
82
A. Fain<br />
11<br />
sh<br />
9<br />
Figures 8-9<br />
Figure 10<br />
Figure 11<br />
Ctenocolletacarus grandior sp. novo Hypopns in ventral view (8). Rostrum and<br />
palposoma in ventral view (9).<br />
Ctenocolletacarus l<strong>on</strong>girostris sp. novo Hypopus. Rostrum and palposoma in<br />
ventral view.<br />
Ctenocolletacarus brevirostris sp. novo Hypopus. Rostrum and palposoma in<br />
ventral view.<br />
83
A New Genus <str<strong>on</strong>g>of</str<strong>on</strong>g> Mite Phoretic <strong>on</strong> Bees<br />
Figures 12-13 Ctenocolletacarus brevirostris sp. novo Hypopus in ventral view (12). Tarsus I in<br />
dorsal view (13).<br />
84
A. Fain<br />
as in C. l<strong>on</strong>girostris except the suctorial plate which is relatively much larger<br />
(width 155 flm) and bears much larger suckers. Anterior suckers 18-20 flm wide,<br />
posterior suckers 33-36 flm wide (excepti<strong>on</strong>ally 38 flm). Lateral c<strong>on</strong>oids slightly<br />
more anterior than posterior <strong>on</strong>es and preceded by large oval flat area. Legs l<strong>on</strong>ger<br />
than in C. l<strong>on</strong>girostris; length <str<strong>on</strong>g>of</str<strong>on</strong>g> tarsi (in flm): 31-31-3333.<br />
Ctenocolletacarus brevirostris sp. nov.<br />
Figures 7, 11-13<br />
Holotype<br />
WAM 83/171, hypopus <strong>on</strong> slide, ex tergal pouches <str<strong>on</strong>g>of</str<strong>on</strong>g> Ctenocolletes albomarginatus 9 (WAM<br />
47/1232); Merredin, Western Australia, presumably in 1947, A.M. Douglas.<br />
Paratypes<br />
Western Australia<br />
Ctenocolletes albomarginatus: 20 hypopi, same data as for holotype (WAM 16 hypopi, WAM<br />
83/353-5; IRSNB 4 hypopi); 38 hypopi ex tergal pouches <str<strong>on</strong>g>of</str<strong>on</strong>g> 9 (WAM 82/1879); 37 km NE <str<strong>on</strong>g>of</str<strong>on</strong>g><br />
Lavert<strong>on</strong> (28°21'S, 122°37'E), 10-12.ix.1982, B. Hanich and T.F. Houst<strong>on</strong> (WAM 29 hypopi,<br />
WAM 83/350-2; IRSNB 5 hypopi; BMNH 4 hypopi); 9 hypopi from body <str<strong>on</strong>g>of</str<strong>on</strong>g> d (WAM 82/122),<br />
21 km N <str<strong>on</strong>g>of</str<strong>on</strong>g> Geraldt<strong>on</strong>, 19.viii.l971, T.F. Houst<strong>on</strong> (WAM 5 hypopi, WAM 83/348-9; IRSNB<br />
2 hypopi; BMNH 2 hypopi).<br />
Diagnosis<br />
Differs from the two other species in the <str<strong>on</strong>g>genus</str<strong>on</strong>g> by the much shorter rostrum,<br />
the spinose dorsal chaetotaxy and the network formed by the dorsal striati<strong>on</strong>.<br />
Descripti<strong>on</strong><br />
Holotype 450 flm l<strong>on</strong>g, 240 flm wide. In 8 paratypes (in flm): 460 x 265 (largest<br />
specimen); 435 x 250;423 x 237;405 x 238;390x240;363x225;340x220;<br />
300 x 180 (smallest specimen). Posterior margin <str<strong>on</strong>g>of</str<strong>on</strong>g> body with very narrow<br />
membrane. Dorsum as in C. l<strong>on</strong>girostris but transverse lines are c<strong>on</strong>nected to each<br />
other and form an irregular network. Dorsal setae are short thick spines, se i<br />
situated distinctly in fr<strong>on</strong>t <str<strong>on</strong>g>of</str<strong>on</strong>g> se e. Rostrum short and with rounded apex, bearing<br />
small v i setae. Venter: Palposoma slightly l<strong>on</strong>ger than wide. Epimeres and<br />
suctorial plate as in C. l<strong>on</strong>girostris, the plate is 93 flm wide, anterior suckers 15 flm,<br />
wide, posterior suckers 21 flm wide. Lateral c<strong>on</strong>oids either <strong>on</strong> the same line as<br />
posterior suckers or slightly behind this line. Legs thicker than in C. l<strong>on</strong>girostris<br />
and bearing much thicker spines <strong>on</strong> tibiae I-IV and <strong>on</strong> genua I-I!. Tarsi I-IV<br />
29-29-30 and 30 flm l<strong>on</strong>g.<br />
Discussi<strong>on</strong><br />
The material examined was obtained from a relatively small number <str<strong>on</strong>g>of</str<strong>on</strong>g> host<br />
specimens: C. l<strong>on</strong>girostris from 3 speCimens each <str<strong>on</strong>g>of</str<strong>on</strong>g> Ctenocolletes centralis and<br />
85
A New Genus <str<strong>on</strong>g>of</str<strong>on</strong>g> Mite Phoretic <strong>on</strong> Bees<br />
C. nichols<strong>on</strong>i; C. grandior from 2 Ctenocolletes centralis and 1 C. nichols<strong>on</strong>i; and<br />
C. brevirostris from 3 Ctenocolletes albomarginatus. However, <strong>on</strong> the basis <str<strong>on</strong>g>of</str<strong>on</strong>g><br />
these records it appears that C. l<strong>on</strong>girostris and C. grandior share (and are<br />
c<strong>on</strong>fined to) two bee hosts (Ctenocolletes centralis and C. nichols<strong>on</strong>i) while<br />
C. brevirostris is c<strong>on</strong>fined to a third (C. albomarginatus). This situati<strong>on</strong> reflects<br />
the affinities <str<strong>on</strong>g>of</str<strong>on</strong>g> the host <strong>bees</strong> where centralis and nichols<strong>on</strong>i are very closely<br />
related to each other and more distantly related to albomarginatus (Houst<strong>on</strong><br />
1983).<br />
Acknowledgements<br />
I am grateful to Dr T.F. Houst<strong>on</strong> for his c<strong>on</strong>structive criticism <str<strong>on</strong>g>of</str<strong>on</strong>g> a draft <str<strong>on</strong>g>of</str<strong>on</strong>g> this<br />
paper.<br />
References<br />
Berlese, A. (1923). Centuria sesta di Acari nuovi. Redia, 15: 262.<br />
Fain, A. (1973). Notes sur les hypopes des Saproglyphidae (Acarina: Sarcoptif<strong>on</strong>nes) HI. Le<br />
genre Crabrovidia Zachvatkin, 1841. Descripti<strong>on</strong> de huit especes nouvelles symphoretiques<br />
sur les Sphecidae (Hymenopteres). Bull. Ann. Soc. r. belg. Ent. 109: 153-189.<br />
Fain, A. and Beaucournu,j.C. (1973). Deuxnouveauxhypopes vivant en associati<strong>on</strong> ph<strong>on</strong>!tique<br />
sur des puces de carnivores (Acarina: Sarcoptiformes). Acarologia 15: 138-143.<br />
Houst<strong>on</strong>, T.F. (1983). A revisi<strong>on</strong> <str<strong>on</strong>g>of</str<strong>on</strong>g> the bee <str<strong>on</strong>g>genus</str<strong>on</strong>g> Ctenocolletes (Hymenoptera: Stenotritidae).<br />
Rec. West. Aust. Mus. 10 (3): 269-306.<br />
Oudemans, A.C. (1916). Acari verzameld bij B<strong>on</strong>n. Ent. Ber., Ams IV, no. 88: 261-266.<br />
Zachvatkin, A. (1941). Fauna <str<strong>on</strong>g>of</str<strong>on</strong>g> U.S.S.R. Arachnoidea. Vol. VI, no. 1. Tyroglyphoidea (Acari).<br />
Zool. Inst. Acad. Sci. U.S.S.R. <str<strong>on</strong>g>new</str<strong>on</strong>g> ser. 28 (English translati<strong>on</strong>, 1959: 1-573).<br />
Received 7 April 1983 Aecepted 1 June 1983<br />
Published 19 April 1984<br />
86