1.1.2.2 The adult male

Soft Scale Insects - Their Biology, Natural Enemies and Control

Y. Ben-Dov and C.J. Hodgson (Editors) 9 1997 Elsevier Science B.V. All rights reserved.

23

1 . 1 . 2 . 2 The Adult Male JAN H. GILIOMEE

INTRODUCTION The morphology of male Coccidae (Fig. 1.1.2.2.1) has been the subject of extensive studies by Giliomee (1967) and Miller (1991). These two authors have described males of about 50 species and expanded our knowledge of the male morphology and taxonomy of this family, which had been previously based mostly on the detailed morphological study of just one species (Theron, 1958). Using the terminology of Giliomee (1967), a few further species have been described by Gimpel et al. (1974), Ray & Williams (1980, 1983), Manawadu (1986), Farrel (1990) and Hodgson (1991, 1993). These studies on the male do not corroborate the classifications of the Coccidae suggested by workers such as Bodenheimer (1953) and Borchsenius (1957), who used only female characters. Giliomee (1967) proposed four species-groups, i.e. Eulecanium-, Eriopeltis-, Inglisia- and Coccus-groups. Later, Ray & Williams (1983) added the Toumeyella-group for the males of the genera Neolecanium, Pseudophilippia and Toumeyella studied by them. Miller (1991) in general supported these groupings. He redefined the Eulecanium-, Eriopeltis-, Coccus- and Toumeyella-groups and added the Philephedra-, Sphaerolecanium- and Protopulvinaria-groups. The f'mdings from studies on the male were ignored in the classification proposed by Tang Fang-teh et al. (1990) and Tang Fang-teh (1991). However, Hodgson (1994) made a courageous attempt to end the schism by taking the relationships indicated by the studies of the male into consideration when he proposed the division of the Coccidae into 10 subfamilies. The description that follows is mainly based on Giliomee (1967) and Miller (1991). The abbreviations refer to Fig. 1.1.2.2.2.

GENERAL APPEARANCE In contrast to the female, adult male Coccidae have a typical insect shape, with a clearly defined head, thorax and abdomen. The front wings and three pairs of legs are well developed. The antennae are long and filiform and the elongated penial sheath is conspicuous, forming the tip of the abdomen. A long wax filament is produced by a glandular pouch on each side of the VIII abdominal segment. The total length of the male varies between 1 and 3 mm, depending on the species.

HEAD The rounded head capsule is partly sclerotizexl. Dorsomedially, a reticulated median crest (me) is found, which extends anteriorly over the apex of the head. In some species, it carries vestiges of the midcranial ridge (mcr) dorsally or anteriorly. The large, reticulated ocular sclerite (oes) covers most of the ventral and part of the lateral surface of the head capsule. It is partly bounded anteriorly by the preocular ridge (procr), which bears a process for articulation with the scape, and posteriorly by the prominent postocular ridge (pocr). In some cases the latter surrounds or partly

Section 1.1.2.2 references, p. 30

24

Morphology

U Fig. 1.1.2.2.1. Dorsal view of an adult male soft scale insect.

surrounds the ocellus. The two ridges are sometimes joined together below the ocellus by an interocular ridge (ior). Ventrally, the ventral part of the midcranial ridge (vmer) extends anteriorly from the ocular sclerite and forks into the lateral arms of the midcranial ridge (liner), which may also bifurcate. Sometimes a linear vestigial ridge is present dorsally on the median crest, representing the dorsomedial part of the mideranial ridge (diner). On the posterior edge of the ocular sclerite, a narrow preoral ridge (WoO links the two postocular ridges and supports a scoop-like cranial apophysis (ca). Behind this structure a vestigial mouth opening (mo) is found, surrounded by two tendon-like apodemes (t), and two or four tentorial pits (atp, ptp). The ocular sclerite bears a pair of lateral ocelli (o) and a variable number of simple eyes. Some species only have large pairs of dorsal simple eyes (dse) and ventral simple eyes (vse), but others also possess 1 to 3 pairs of smaller lateral simple eyes

(Ise). Behind the postocular ridge, the gena (g) is found in the form of a reticulated bulge. The antennae are inserted on the anterolateral margin of the head. They are usually 10-segmented, but antennae with 9 or fewer segments are found in some groups. The scape (scp) is short and broad, the pedicel (pdc) subglobular and the flagellar segments (Fro-t,,) tubular.

THORAX Prothorax. A deep cervical constriction separates the head from the largely membranous prothorax. Just posterior to the constriction, the collar-like pronotal ridge (prnr), with the associated pronotal sderite (prn) on each side, is very distinct. Posterolaterally a small, weakly sclerotized, sometimes striated posttergite (pt) is found. Pleurally, a ridge-like pro-episternum + cervical sclerite (pepcv) stretches from the postocular ridge anteriorly to the short propleural ridge (plr~), which articulates with the coxa of the front leg; posteriorly, an invagination of the pleural ridge forms a small propleural apophysis (plal). The prosternum (stnl) consists of a triangular or oval

The adult male

25

sclerite, bounded posteriorly by a transverse ridge and traversed medially by a longitudinal ridge which shows varying degrees of inter- and intraspecific development. Mesothorax. As principal wing-bearing segment, this part of the body is heavily sclerotized, with strong ridges and sutures. The notum is widely separated from the postnotum by a membranous area and is subdivided into a prescutum, scutum and scutellum. The prescutum (prsc) has the shape of a subrectangular, reticulated bulge, which bears an internal mesoprephragma anteriorly. Laterally and posteriorly it is separated from the scutum by strong prescutal ridges (pscr) and a prescutal suture (pscs) respectively. The scutum (set) is characterized by a large, median, membranous area and two lateral parts which extend along the prescutum anteriorly and the scutellum posteriorly. Anterolateral to the scutum, a semitubular prealare (pra) and convex triangular plate (tp) extend to the epistemum. Posteriorly, along the lateral margin of the scutum, an anterior notal wing process (anp) and a posterior notai wing process (pnp) can be discerned. The scuteilum (scl) has a transversely rectangular shape and is usually semi tubular with an internal scuteilar foramen (sclf). Behind the scutellum a large membranous area separates it from the mesopostnotum (pnz), which curves deeply into the metathoracic cavity. The postnotum bears a mesopostphragma on its posterior margin and a deep postnotal apophysis (pna) on each side. Anterolaterally it is produced into a strong postalare (pa) which articulates with the mesopleural ridge and bears the anterior and posterior postalar ridges (apar, ppar) on its margins. Pleurally, the large episternum (epsz) is divided into dorsal and ventral parts by a membranous cleft. The dorsal part is convex and bounded anteriorly by a subepisternal ridge (ser) which extends across the cleft. The ventral part is a narrow sclerite which extends from the mesopleural ridge (plr2) posteriorly to the lateropleurite (lpi) anteriorly. The mesopleural ridge (plrz) is a broadly sclerotized ridge which stretches obliquely across the pleuron from the large, rounded mesopleural wing process (pwpz) dorsally to the coxa ventrally. The mesopleural apophysis (plaz) is invaginated, where it is overlapped by the postalare. On the lower anterior margin, a small, tendon-like apodeme (t) is found and posterodorsally a small subalare (sa). The pleural wing process is usually connected with the epistemum by means of a narrow sclerite, the basalare (bas); in some species, such as Coccus hesperidum L., it is vestigial. Posterior to the lower extremity of the pleural wing process, a small sclerite represents the mesepimeron (epmz). The mesothoracic spiracle (sp,.), supported by a peritreme (ptrz), is situated behind the pleural apophysis of the prothorax. The mesosternum consists almost entirely of the large basisternum (stnz), of which the anterior part is bounded by a short marginal ridge (mr) and the posterior part by a precoxal ridge (pcrz). A longitudinal median ridge (mdr), which is sometimes interrupted or vestigial, divides the basistemum into two parts. On the posterior margin of the basistemum, a transverse furcal pit (fp) with a large internal furca (f) is situated.

Metathorax. The metathorax is largely membranous. However, a well developed, invaginated metanotum closely overlaps the mesopostnotum, with its posterior margin usually forming a ridge-like structure dorsally. On each side, a suspensorial sclerite (ss) anchors the haltere when the latter is present, while further posteriorly a small transverse sclerite represents the metapostnotum (pn3). Pleurally, the metapleural ridge (pir 3) extends dorsally from the coxal articulation towards the base of the haltere where it expands into a small metapleurai wing process (pwp3). About half-way from the coxa, the metapleurai apophysis (pla3) is represented by a shallow depression. When the halteres are absent, the ridge only extends for a short distance above the coxal articulation. From the ridge, the metepisternurn (epss) extends anteroventrally and the metepimeron (epm3) posteriorly. A vestigial precoxal ridge (pcr3) sometimes extends anteriorly from the pleural ridge along the margin of the episternum.

Section 1.1.2.2 references, p. 30

26

Morphology

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27

The adult male

Abbreviations used in Fig. 1.1.2.2.2 aas ab ads

aed al ams amss an anp

apar as asc ase

astnls at atp ax 1

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cb CCX ce

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fp fs g gls gP gs gts h hs ior lmcr lpl lpns lse mc mdr med mo

mpns nlr mt8 o ocs

pa per:

ante-anal setae antennal bristles abdominal dorsal setae aedeagus alar lobe antemetaspiracular setae anterior metasternal setae anus anterior notal wing process anterior postalar ridge abdominal sternite additional sclerite apical seta anteprosternal seta abdominal tergite anterior tentorial pit first axillary wing sclerite second axillary wing sclerite third axillary wing sclerite basal rod of aedeagus basalare basal membranous area sensilla basiconica cicatrix cranial apophysis coxal bristle(s) costal complex of wing veins caudal extension claw coxa dorsal head setae dorsomedial part of midcranial ridge dorsal ocular setae dorsopleural setae dorsal simple eye dorsospiracular setae mesepimeron metepimeron mesepisternum metepisternum postmetaspiracular setae furca segments of flagellum, 3rd to 10th femur furcal pit fleshy seta gena seta of glandular pouch glandular pouch genal setae setae of genital segment haltere hair-like seta interocular ridge lateral branch of midcranial ridge lateropleurite lateral pronotal setae lateral simple eye median crest median ridge media mouth opening medial pronotal setae marginal ridge metatergal setae ocellus ocular sclerite postalare precoxal ridge of mesothorax

pcq pdc pepcv plat plaz pla3 plr~ plr2 plrs pms pmss pth pn3 pna pnp pnr pocr ppar pra prn prnr procr pror prsc ps pscr pscs pt pta ptp ptr2 ptr3 pts pwp2 pwp3 rad sa set.scla scl self scls scp set sctse ser

sp2 sP3 spl ss stn 1

stn2 stn3 stnls stn2s t

tar tdgt teg tegs tib tibs tp tr udgt vhs vmcr vps vs vse

vestigial precoxal ridge of metathorax pedicel proepisternum + cervical sclerite propleural apophysis mesopleural apophysis vestigial metapleural apophysis propleural ridge mesopleural ridge metapleural ridge postmesospiracular setae posterior metasternal setae mesopostnotum metapostnotum postnotal apophysis posterior notal wing process pronotal ridge postocular ridge posterior postalar ridge prealare lateral pronotal sclerite pronotal ridge preocular ridge preoral ridge prescutum penial sheath prescutal ridge prescutal suture posttergite posterior tentorial arm posterior tentorial pit peritreme of mesothoracic spiracle peritreme of metathoracic spiracle posttergital setae mesopleural wing process vestigial metapleural wing process radius subalare subapical seta scutellum scutellar foramen scutellar setae scape scutum scutal setae subepisternal ridge mesothoracic spiracle metathoracic spiracle sensillum placodeum suspensorial sclerite prosternum basisternum of mesosternum metasternum prosternal setae basisternal setae tendon-like apodeme tarsus tarsal digitule tegula tegular setae tibia tibial spur triangular plate trochanter ungual digitule ventral head setae ventral part of midcranial ridge ventropleural setae ventral sclerite ventral simple eye

Morphology

28

Anterior to the epistemum, the metathoracic spiracle (spa) and its supporting peritreme (ptr3) are located. Ventrally a median plate or irregular sclerotization represents the metasternum (Stna).

Wings The semitransparent fore-wings are about as long as the body (excluding the genital segment) and two to three times longer than wide. They have a narrow base and a broadly rounded apex. When halteres are present, a small alar lobe (ai) is found near the base on the hind-margin of the wing; the apical setae of the haltere hook onto an invagination of the lobe. Two distinct wing veins are present, the radius (rad) and media (reed), while an elongate sclerite near the anterior margin of the wing forms the costal complex of wing veins (ccx) which articulates with the anterior notal wing process. Other sclerites involved in the articulation of the wing are the tegula (teg), three axillary sclerites (axl, ax2 and ax a) the additional sclerite (asc). The hind wings are either absent or reduced to halteres (h), also called hamulohalteres. They usually bear one hooked seta, but in some species there may be three or four.

Legs All three pairs of legs are usually long and slender. The eoxa (cx) is short and broad, and articulates distally with the short trochanter (tr) which is narrow basally and broad distally, and is separated from the femur (fro) by a narrow strip of membrane. The femur is long and broad, the tibia (tib) long and slender. The elongate tarsus (tar) is one-segmented and articulates with a single claw (el).

ABDOMEN The abdomen is composed of eight pregenital segments and a 9th or genital segment; the first segment is not developed ventrally. The segmentation is indicated by shallow transverse grooves. Most of the abdomen is membranous, but tergal and sternal plates may be present in some species. Where present, the abdominal tergites (at) are usually situated laterally in the more anterior segments and medially in the posterior segments. The abdominal sternites (as) are large, transverse plates which may be absent or medially interrupted in the intermediate segments. Pleural sclerotization may be present on the caudal extensions of segment VII and VIII and occasionally more anteriorly. Caudal extensions (ce) are found on the VII and VIII segments of many species. Their shape and size vary interspecifically. Those of the VII segment are usually lobiform, but they may be tapering and finger-like. On the VIII segment, they may be lobiform, cylindrical, mammillate or various other shapes. In the Coccus-group of species, a circular, membranous, weakly-reticulated cicatrix (c) is found on the distal part of caudal extension VIII. Near the posterior rim of the VIII segment, a funnel-shaped glandular pouch (gp) with two long setae is usually present on each side. These setae give support to the waxy filament produced by the multilocular pores in the pouch. The filaments are very conspicuous in the living males but their function is uncertain. The pouch and pores are absent in some species. The IX or genital segment is elongate and forms a long, tubular style which is comprised of the penial sheath (ps) and the aedeagus (aed). The penial sheath is composed of sternum IX and sclerotized laterally. Anteroventrally, a basal membranous area (bma) is found and, posterior to this, a narrow median ridge represents the basal rod (bra). The latter supports the tubular aedeagus which is situated in a slit of the penial sheath. Dorsally, a small anus (an) is present in the membrane at the base of the segment. Anterior to this a small 9th tergite (atg) can sometimes be seen.

The adult male

29

CHAETOTAXY The two basic types of setae of male Coccidae are the fleshy setae (fs) and hair-like (hs) setae. The former are more thick-set, with a blunt apex and the setal membrane is not surrounded by a distinct basal ring; the latter have an acute apex and a distinct basal ring. Bristles, larger than the fleshy setae, are present on the distal segments of the antennae and on the front coxae. The setae of the head can be divided into four groups, i.e. dorsal head setae (dhs), dorsal ocular setae (dos), ventral head setae (vhs) and genal setae (gs), the latter being only present in some groups of species. When the setae are predominantly of the fleshy type, as in C. hesperidum, the head appears distinctly hairy. The antennae also appear hairy, carrying numerous fleshy and a few hair-like setae. Large antennal bristles (ab) are present on the distal segments. On the terminal segment, long, capitate subapical setae (set. scla) are found. The latter are usually three in number, but may vary from zero to six. On the prothorax, several groups of fleshy and hair-like setae are found, i.e. the lateral pronotal setae (lpns), medial pronotal setae (mpns), posttergital setae (pts), anteprosternal setae (astnis) and prosternai setae (stnis). However, the full complement is only present on some species. On the mesothorax, one may find scutai setae (sctse), scutellar setae, (scls), tegular setae (tegs), postmesospiracular setae (pms) and basisternal setae (stn=,s). Metathoracic setae that may be present are the metatergai setae (mts), dorsospiracular setae (dss), antemetaspiracular setae (ams), postmetaspiracular setae (eps~s), anterior metasternal setae (amss) and posterior metasternal setae (press). The wing surface is covered with microtrichia and, in addition, a small number of hair-like alar setae (als) may be present on the anterior part of the base of the wing. On the legs, fleshy and hair-like setae occur abundantly on all the segments except the claw. In some species, the anterior coxae also carry long coxal bristles (cb) which are sometimes capitate. Ventrally, near the apex of the coxa and the trochanter, one or two long, hair-like apical setae (ase) are found, while an apical spur (tibs) occurs in this position on the tibia. A minute seta occurs anteriorly and posteriorly in the articular membrane between the coxa and trochanter. A pair of capitate tarsal digitules (tdgt) are present near the dorsal apex of the tarsus and a pair of claw or ungual digitules (udgt) occur on the claw. The abdomen usually has a single pair of hair-like dorsal setae (ads) on segments IV to VII, but in some species they are present on all segments; dorsal fleshy setae are present on some species. Pleural setae occur in groups of dorsopleural setae (dps) and ventropleural setae (vps) which sometimes coalesce, while ventral setae (avs) occur medially on the segments. The position and number of the hair-like setae are more constant than those of the fleshy setae. In addition to these setae, three hair-like setae and occasionally a fleshy seta occur lateral to the glandular pouch on the posterior margin of the VIII segment. In the region anterior to the anus, two long, two short and a few fleshy ante-anal setae (aas) may be present. On the penial sheath a number of small, scattered genital setae (gts), which are probably tactile receptors, can be seen. Fleshy setae are found on the penial sheath of Ceroplastes ceriferus (Fabricius) (Miller, 1991).

OTHER CUTICuLAR STRUCTURES Sensilla and pores occur on various parts of the body. Some of the sensilla are very constant in position and occur in the same position on all the species studied. Examples are the sensillum placodeum (spi), which occurs distally on the dorsolateral surface of the pedicel, the two sensilla basiconica (bs) on the ventral surface of the terminal antennal segment and the ring of six (sometimes eight) oval, campaniform sensilla in the

Section 1.1.2.2 references, p. 30

30

Morphology

basal half of the trochanter. A variable number of sensilla basiconica may occur on the 3rd antennal segment and what are possibly campaniform sensilla on the apex of the style. Very few pores are found on the body. In addition to the multilocular disc-pores of the glandular pouch on the VIII abdominal segment, a variable number of circular pores that resemble seta sockets, from which the seta was detached, occur dorsally behind the pronotal ridge in some species. Minute dermal denticulations are present on the dorsal and ventral surfaces of the median part of each abdominal segment.

REFERENCES Bodenheimer, F.S., 1953. The Coccoidea of Turkey. III. Revue de la Facult6 des Sciences de l'Universit6 d'instanbul (Series B), 18: 91-164. Borchsenius, N.S., 1957. Sucking Insects, Vol. IX, Suborder mealybugs and scale insects (Coccoidea). Family cushion and false scale insects (Coccidae). Fauna SSSR. Zoologicheskii Institut Academii Nauk SSSR, Novaya seriya, 66: 1-493. (In Russian). Farrel, G.S., 1990. Redescription of Cryptes baccatus (Maskell) (Coccoidea: Coccidae), an Australian species of soft scale. Memoirs of the Museum of Victoria, 51: 65-82. Giliomee, J.H., 1967. Morphology and taxonomy of adult males of the family Coccidae (Homoptera: Coccidae). Bulletin of the British Museum (Natural History), Entomology Supplement 7: 1-168. Gimpel, W.F., Miller, D.R. and Davidson, J.A., 1974. A systematic revision of the wax scales, genus Ceroplastes, in the United States (Homoptera: Coccoidea: Coccidae). Miscellaneous Publication Agricultural Experiment Station, University of Maryland, 841 : 1-85. Hodgson, C. J., 1991. A redescription of Pseudopulvinaria silda'mensis Atkinson (Homoptera, Coccoidea), with a discussion of its affinities. Journal of Natural History, 25:1513-1529. Hodgson, C. J., 1993. The immature instars and adult male of Etiennea (Homoptera: Coccidae) with a discussion of its affinities. Journal of African Zoology, 107: 193-215. Hodgson, C. J., 1994. The Scale Insect Family Coccidae: an Identification Manual to Genera. CAB International, Wallingford, UK, 639 pp. Manawadu, D., 1986. A new species ofEriopeltis Signoret (Homoptera: Coccidae) from Britain. Systematic Entomology, 11 : 317-326. Miller, G.L., 1991. Morphology and Systematics of the Male Tests and Adult Males of the Family Coccidae (Homoptera: Coccoidea) from America North of Mexico. Ph.D. Thesis, Auburn University, Auburn, USA. Ray, C.H. and Williams, M.L., 1980. Description of the immature stages and adult male of Pseudophilippia quaintancii (Homoptera: Coccoidea: Coccidae). Annals of the Entomological Society of America, 73: 437-447. Ray, C.H. and Williams, M.L., 1983. Description of the immature stages and adult male of Neolecanium cornuparvum (Homoptera: Coccidae). Proceedings of the Entomological Society of Washington, 85: 161-173. Tang, Fang-teh, 1991. The Coccidae of China. Shanxi United Universities Press, China, 377 pp. (In Chinese, English summary). Tang, Fang-teh, Hao, J., Xie, Y. and Tang, Y., 1990. Family group classification of Asiatic Coccidae (Homoptera, Coccoidea, Coccidae). Proceedings of the Vlth International Symposium of Scale Insect Studies, Cracow, August 6-12th, 1990. Part II: 75-77. Theron, J.G., 1958. Comparative studies on the morphology of male scale insects (Hemiptera: Coccoidea). Annals of the University of Stellenbosch, 34(A): 1-71.