FNZ 51 - Coccidae males (Insecta: Hemiptera: Coccoidea) - Adult males, morphology

Hodgson, CJ; Henderson, RC 2004. Coccidae (Insecta: Hemiptera: Coccoidea): adult males, pupae and prepupae of indigenous species. Fauna of New Zealand 51, 228 pages.
( ISSN 0111-5383 (print), ; no. 51. ISBN 0-478-09360-8 (print), ). Published 22 Jun 2004
ZooBank: http://zoobank.org/References/85A2C19D-CA28-4C43-9B7C-BABE3D5DFCE8

Adult males, morphology

Basic shape : typical insects, with a cylindrical body divided into 3 tagmata (head, thorax, and abdomen) and with well-developed forewings on the mesothorax and 3 pairs of long legs. The metathoracic wings have become modified into hamulohalteres (although these have been secondarily lost from most New Zealand species). Males also have long, filiform, 10-segmented antennae and a conspicuous, elongate, sclerotised, penial sheath which projects posteriorly from the abdomen.

Size : the adult males included in this study are relatively small, from around 1-2.3 mm, although coccid males from elsewhere can be up to about 3 mm long.

Chaetotaxy (Fig. 16, 18, 20): setae are of 2 main kinds, fleshy setae (fs) and hairlike setae (hs). Hairlike setae are, as their name suggests, hairlike or flagellate and have a small sclerotised ring or setal socket around the setal membrane at their base. Fleshy setae tend to be thicker and blunter and lack a sclerotised socket. However, the sclerotised ring can be hard to see and some fs are distinctly flagellate (as on Crystallotesta ornata (Maskell) and C. ornatella Henderson & Hodgson, Fig. 68, 69) and so differentiating between these two kinds of setae can be difficult. With the exception of Poropeza dacrydii (Maskell) (Fig. 84), hs are generally fewer and their positions tend to be more fixed and predictable. Thus, setae not in the predicted positions for hs are here generally referred to as fs unless the setal socket can be distinguished.

In addition, there are some setae that are referred to as bristles . These occur on antennal segments VIII, IX, and X (referred to as antennal bristles ). Giliomee (1967), Miller (1991), and Miller & Williams (2002) also found bristles on the anterior coxae of some coccid species, but these are absent from New Zealand species. Bristles are similar to fs but generally significantly stouter and larger.

Spurs , which are probably enlarged and thickened hairlike setae, are also present on the ventral surface of the distal end of most tibiae and tarsi. Most New Zealand species have single tibial and tarsal spurs, but both Pounamococcus species (Fig. 85, 86) have 2 tibial spurs per tibia, whilst spurs are absent on Inglisia patella Maskell (Fig. 75). Some of the setae near the distal end of the tibia are also generally spurlike, but are less clearly differentiated and smaller. Although not always clearly differentiated, the most posterior ventral seta near each claw is also generally spurlike, and is here referred to as a tarsal spur.

MOUNTED MATERIAL (Fig. 16-21)

We stress that the following descriptions are based on specimens that were stained and slide-mounted dorsoventrally in Canada balsam. Consequently, the body shape will have become distorted to some extent; in particular:

the head shape will have changed,
the sclerotised structures of the thorax have often been broken and distorted (they can be rather convex in life) and
the membranous abdomen appears broader in the figures than in real life.

Furthermore, marginal structures can be difficult to see; in particular, the basalare can rarely be seen and this sclerite may be important in separating higher taxa (Miller 1991).

In addition, it can be difficult to separate

the dorsospiracular setae on the metathorax from the antemetaspiracular setae, which lie more ventrally, and
the dorsopleural setae from the ventropleural setae.

The wing sclerites are also generally distorted and differences in their shape in the figures should not be taken as significant.

HEAD

The head capsule is rather oval to roundly quadrangular in shape, with either 2 or 4 pairs of simple eyes , with 1 or 2 pairs present anterodorsally and 1 or 2 pairs posteroventrally, the latter sometimes on a cone-shaped extension to the ventral surface of the head when seen from the side (as probably on Pounamococcus spp.). There is also a pair of smaller ocelli laterally. The antennae are long and 10-segmented, arising near the anterior margin of the head capsule. Mouthparts are entirely absent, but a non-functional mouth opening is present behind the ventral simple eyes.

Between the antennal bases is the median crest , which extends posteriorly on the dorsal surface to about half the length of the head: this is a slightly sclerotised and polygonally reticulated area, with fs and hs dorsal head setae ; it is generally rounded at the posterior end, where it may be slightly more heavily sclerotised. Rarely, vestiges of a sclerotised post-occipital ridge may be present at right angles to the median crest (cf. on P. dacrydii , Fig. 84 and Species A, Fig. 92) at its posterior end. The median crest may also have vestiges of a sclerotised dorsal section of the midcranial ridge (cf. Species A, Fig. 92, and Lecanochiton spp., Fig. 78, 79). Ventrally, the midcranial ridge is present as a strongly sclerotised, Y-shaped, longitudinal ridge, with the ventral part extending posteriorly from the median crest towards the ocular sclerite and with a pair of sclerotised arms at the anterior end ( lateral arms of midcranial ridge ), each arm extending towards a scape but not articulating with it. The ventral midcranial ridge may reach the ocular sclerite posteriorly or only extend part way; it is usually well developed but may be reduced (as possibly on Umbonichiton jubatus Henderson & Hodgson: Fig. 90). Laterad to the ventral midcranial ridge and posterior to each scape is a membranous area bounded laterally by the preocular ridge . The area immediately bordering the ventral midcranial ridge is usually polygonally reticulated, narrow anteriorly but widening posteriorly where it fuses with the ocular sclerite (but may be reticulated throughout, as on Pounamococcus cuneatus Henderson & Hodgson: Fig. 85); this area usually has a few fs and hs (here referred to as ventral midcranial ridge setae ).

Articulating with the base of the antennae laterally is the preocular ridge , which is a heavily sclerotised ridge which extends both anteriorly and posteriorly from the antennal articulation and marks the antero-ventral margin of the ocular sclerite . On most mounted specimens, the anterior extension of the preocular ridge cannot be seen easily as it lies beneath the scape but is usually as long as the posterior part of the ridge, which varies in length from rather short (e.g., on I. patella , Fig. 75) to long, almost reaching the ventral midcranial ridge medially (as on Aphenochiton kamahi Henderson & Hodgson, Fig. 63) (on some non-indigenous species the ventral arms of the preocular ridge fuse medially). Making up most of the lateral part of the head is the ocular sclerite , which extends laterally from the dorsal simple eyes down between the preocular and postocular ridges to the ventral simple eyes , where they fuse. This sclerite is lightly sclerotised and polygonally reticulated throughout and has several to many fs and hs. The polygonal reticulations vary between species both in shape and size (i.e. each reticulation tends to be rather larger on Ctenochiton spp. than on most other indigenous species) and in the presence or absence of additional inner dermal ridges. These inner dermal ridges may take the form of 1 or 2 long, straightish microridges (as on A. kamahi , Fig. 63) or may be numerous, sinuous, divided or broken microridges (as on Crystallotesta fagi (Maskell), Fig. 67). Inner dermal microridges appear to be absent from Plumichiton flavus (Maskell) (Fig. 81). These differences appear to be fairly constant and are here considered to be of taxonomic significance. The setae on the ocular sclerite are referred to as ventral head setae and their number and distribution vary between taxa: they may be abundant and found throughout, including between the ventral eyes (as on Plumichiton flavus (Maskell), Fig. 81, and most Umbonichiton spp.), or they may be very few and restricted to just anterior to the ventral eyes (as on Umbonichiton jubatus Henderson & Hodgson, Fig. 90). Where these setae are also present posterior to the ventral eyes (as on P. flavus , Fig. 81), they are here referred to as ventral ocular setae . The area between the ocelli (or more properly the postocular ridge) and the dorsal simple eyes is part of the ocular sclerite and may bear setae, the dorsal ocular setae ; when present, these are generally fs. When the postocular ridge is short (i.e., does not nearly reach the ocelli, as on most New Zealand species), it can be difficult to identify the margin between the ocular sclerite and the genae, but this is usually identifiable by the change in shape of the polygonal reticulations (elongate near ocelli, squarish on gena); setae laterad to or just posterior to the ocelli (i.e., within the area of elongate reticulations) are considered to be dorsal ocular setae, genal setae being considered to be those on the area with non-elongate reticulations.

Marking the posterolateral margins of the ocular sclerite is a long, strongly sclerotised ridge, the postocular ridge ; although most of this ridge runs along the lateral margins of the head, on mounted specimens it generally appears to lie lateroventrally and this is how it is illustrated; it is shown most naturally in the enlarged drawings for Lecanochiton spp. (Fig. 78, 79). The postocular ridge separates the ocular sclerite from the genae dorsally; anterodorsally, the postocular ridge may almost meet the median crest (as on C. ornata and C. ornatella , Fig. 69, 70) but is usually much shorter; when long dorsally, it runs just ventral to each ocellus, where it may divide, with a short arm running around the dorsal margin of each ocellus (common on many non-indigenous species; typical of C. ornata and C. ornatella , Fig. 69, 70); ventrally, the postocular ridge extends round posterior to the ventral simple eyes where it meets the preoral ridge . The interocular ridge , which runs between the preocular and postocular ridges on some soft scales species from elsewhere, is absent from all known indigenous New Zealand species.

The large area posterior to the postocular ridge is the gena , which is membranous and generally polygonally reticulated throughout. As on the ocular sclerite, these reticulations vary in structure: on C. ornata and C. ornatella and several other species, they are barely visible and appear to be represented by numerous raised spots (presumably very short microridges), some of which can form a faint reticulate pattern. On other species, the anterior reticulations are well developed whilst those more posteriorly become faint or absent (e.g., Epelidochiton piperis (Maskell), Fig. 74). They may also have extra inner microridges, which may be similar or dissimilar to those on the ocular sclerite. Genal setae are generally present and are usually fs, but a few hs may also be present; genal setae may be absent (as on Aphenochiton spp., Fig. 62-66).

The simple eyes consist of a dorsal pair and a ventral pair and (in New Zealand species) 0 or 2 pairs of lateral eyes (in non-indigenous species, there can be up to 3 pairs of lateral eyes). In general, the lateral eyes are clearly smaller than the dorsal and ventral pairs but occasionally may be similar in size. The dorsal eyes lie some distance apart (with the median crest between them), dorsolateral to the antennae, whilst the ventral simple eyes tend to be close together on a medioventral bulge. On a few species, such as I. patella , P. cuneatus, and Species A (Fig. 75, 85, 92), the lateral eyes are absent. Posterior to each dorsal lateral simple eye and anterior to the dorsal end of each postocular ridge is an ocellus . These are lightly sclerotised spots, slightly larger than the polygonal reticulations, and can be quite obvious on some species (e.g., on Lecanochiton spp., Fig. 78, 79) but much less so on others (e.g., on C. fagi , Fig. 67). They may be absent on Umbonichiton pellaspis Henderson & Hodgson (Fig. 91).

The preoral ridge is a weak inverted U-shaped structure (not shown in figures), each arm of which meets with a postocular ridge. Arising from the preoral ridge is the cranial apophysis , which is a strong scoop-like apodeme that extends anteriorly within the head (Fig. 16). Its apex is usually bifurcate but may be trifurcate (as on Pounamococcus spp., Fig. 85, 86, and I. patella , Fig. 75).

Also lying internally, dorsad to the cranial apophysis, is a strong, inverted U-shaped structure, the tentorial bridge (Fig. 16). This structure is usually present but appears to be absent on C. ornata and C. ornatella (Fig. 69, 70).

Two kinds of pore are known from New Zealand soft scales. These pores are rare on the males of other, non-indigenous, coccids. I. patella (Fig. 75) has a small group of pores just laterad to the median crest near each scape. These pores are convex, of variable size and shape and are probably not homologous with the loculate pores in the same position on Pseudococcidae and Eriococcidae (Afifi 1968). The second kind of pore, the dorsal pore, is found on P. dacrydii (Fig. 27). It is more uniform in shape, small, round, and convex and is frequent not only on the head but also throughout the membranous areas of the dorsum, including the thorax and abdomen; it is also present on the pleurites but absent from the venter. Giliomee (1967) found small circular pores on Ctenochiton (probably = Avricus ) sp., Genus B and on Luzulaspis luzulae (Dufour) (dorsally on prothorax or abdominal segment VIII or both). Miller & Williams (1995) noted them on Toumeyella virginiana Williams & Kosztarab, and Miller & Williams (2002) mention pores on the scutellum of Philephedra floridana Nakahara & Gill. Because none of these pores were noted on the head, they may not be homologous with the pores found on either New Zealand species.

Antennae : situated on the anterolateral margin of the head but appear ventral on most mounted specimens. They are filiform, 10-segmented (although fewer on some non-New Zealand species due to fusion of certain segments (Giliomee 1967)). They are usually about 0. 6´ total body length but may be as long as 0. 8´ (as on Pounamococcus sp.) or as little as 0. 5´ ( Aphenochiton subtilis Henderson & Hodgson). The scape is short and broad, with (usually) 3 hairlike setae, 1 on the ventral surface and 2 near the inner margins on the dorsal surface (occasionally there are many setae, as on P. dacrydii , Fig. 84). The pedicel is roundish, with a few polygonal reticulations and has both hs and fs; when these are few, they appear to be restricted to the ventral surface but can be found throughout when setae are abundant. The remaining flagellar segments (III-X) are elongate, mostly parallel-sided, with many fs and no, or very few, hs (many on P. dacrydii , Fig. 84), each fs generally about twice as long as the width of the flagellar segments but may be much shorter (as on P. dacrydii , Fig. 84). Segment III is sometimes club-shaped, with a few hs and fs and 1-3 sensilla basiconica ; segments V-VI are usually the longest; segments VIII-IX (Fig. 16) usually each has a single antennal bristle but these can be hard to separate from fs on many species. Segment X (Fig. 16) is short and constricted near the apex on some species, with 3 capitate setae (only 2 on I. patella , Fig. 75), usually 3 long and 2 short antennal bristles near apex, some fs and a sensillum basiconicum on apex and another more proximally between the bases of 2 bristles.

THORAX

This is separated from the head by a deep cervical constriction and consists of a more or less membranous prothorax, a large and mainly sclerotised mesothorax, and a mainly membranous metathorax. Each segment has a pair of long, 5-segmented legs, which are subequal in length. From the mesothorax arise the fore wings, which are large and well developed, although the venation is much reduced. The hind wings are either absent or are represented by a pair of hamulohalteres , as on Pounamococcus spp. (Fig. 85, 86).

Prothorax : mostly membranous; with a strongly sclerotised ridge immediately posterior to the neck region on the dorsal surface, the pronotal ridge , which is usually in 2 lateral parts, with a broad gap ventrally where it articulates with the proepisternum + cervical sclerite and a narrow gap dorsally although, on some non-indigenous species, the 2 halves may be fused. Running along the lateral margins of the pronotal ridge is a sclerite, the lateral pronotal sclerite , which is either striated or polygonally reticulated; on New Zealand species, it is usually well developed and broad. Lateral pronotal setae are associated with the pronotal sclerite. These may be present or absent; when present they are usually represented by a pair of hs but, on P. flavus (Fig. 81), there are up to perhaps 7 fs on each side, although some could be lateral prothoracic setae .

On many Coccoidea, an area posterolaterally on each side of the dorsum is sometimes sclerotised and represents the post-tergites . These are absent from all New Zealand soft scales except I. patella (Fig. 75), although C. ornata (Fig. 69) has a few fs post-tergital setae , whilst P. dacrydii has a group of pores in this position. In addition, P. dacrydii (Fig. 84) has 2 groups of setae and pores laterally on each side of the prothorax and these represent the lateral prothoracic setae .

Ventrally, a strongly sclerotised ridge - the proepisternum + cervical sclerite - runs posteriorly from each postoccipital ridge and fuses with a short pleural ridge , which articulates with the procoxae. An apophysis, the propleural apophysis , is probably present posteriorly but is generally not distinguishable.

Medially on the venter is the prosternum . This is usually somewhat reduced on New Zealand species but, in its most complete state, consists of a large triangular sternite, which is often ridged or even polygonally reticulated, bordered posteriorly by a strongly sclerotised transverse ridge, from which a longitudinal median ridge arises and extends anteriorly down the centre of the sclerite. On New Zealand species, the transverse ridge is always present (and remnants of the prosternal apophyses can sometimes be found laterally) and the sclerite is generally more or less present but the median ridge is absent or poorly developed (present on both Pounamococcus spp., Fig. 85, 86). Almost invariably there is an hs on either side of the sclerite; frequently there are also other hs and some fs; these are the prosternal setae ; they do not usually extend anterior to the procoxae. On other non-indigenous species, these prosternal setae may spread laterally to anterior to the mesothoracic spiracle; on New Zealand species, when there are setae present anterior to the anterior spiracle (as on C. fagi and C. ornata , Fig. 67, 69), they form a distinct group and are here referred to as the antemesospiracular setae . Another group (of generally fs) are sometimes present between the prosternum and the mouth, the anteprosternal setae .

The pores found on P. dacrydii (Fig. 84) are located in groups approximately where the median pronotal setae and post-tergital setae would be and also in a diffuse group medially on the dorsum.

Mesothorax : this has numerous sclerites, many of them bounded by strong ridges. The shape and position of these sclerites varies little within the family.

Anteromedially on the dorsum is the strongly sclerotised prescutum . On unmounted specimens, the prescutum bulges over the prothorax and is quite convex, but on dorsoventrally mounted specimens this is not obvious. The prescutum is rectangular or almost square in shape, bordered anteriorly by the mesoprephragma (which extends internally beneath the dorsum of the prothorax), laterally by the prescutal ridges, and posteriorly by the prescutal suture . The surface of the prescutum is generally fairly smooth but, on a few species, it is clearly polygonally reticulated (e.g., on I. patella and P. dacrydii , Fig. 75, 84); a few species show only very faint reticulations (e.g., E. piperis , Fig. 74). Rarely, it is divided longitudinally by a medial ridge, as on C. ornata (Fig. 69).

The scutum is more or less H-shaped, the central bar being represented by a membranous area between the prescutum and the scutellum, and the 2 upright arms being the sclerotised area laterad to the prescutum, membranous area of the scutum, and the scutellum. The membranous area is bounded anteriorly by the prescutal suture , laterally by a posterior extension of the prescutal ridge, and posteriorly by the scutellum. This membranous area varies somewhat in shape (although this might depend on how the specimen has been mounted): on Lecanochiton spp. and Species A (Fig. 78, 79, 92), it is about 6- 8´ wider than long, whilst on Ctenochiton spp. (Fig. 71-73), it is much longer, being only about 2´ as wide as long. In addition, the membranous area may bear fs and hs scutal setae : usually there is at least 1 pair of hs, as on Species A and Lecanochiton spp. (Fig. 78, 79, 92), although it is possible that even these may occasionally be absent; on other species, setae may be extremely abundant, as on P. dacrydii (Fig. 84), which has about 60 setae, and some Plumichiton spp. (e.g. P. flavus , Fig. 81). Extending laterally from each anterolateral margin of the scutum is a pair of semitubular plates, the prealare ; each end laterally in a heavily sclerotised triangular plate , which extends posteriorly to the mesepisternum . Posterior to each prealare, and bordered laterally by the anterior notal wing process and the mesepisternum is a membranous area, within which is a small, sclerotised plate, the tegula ; usually associated with the tegula are tegular setae ; these are usually hs but a few fs were also noted on some Plumichiton spp. and on P. dacrydii (Fig. 81, 82, 84); tegular setae are absent on some New Zealand species (e.g., Kalasiris perforata (Maskell), Fig. 27).

The anterior part of the scutum is rarely reticulated on New Zealand species (very lightly on Ctenochiton chelyon , Fig. 71, and Inglisia patella , Fig. 75) but the posterior part laterad to the scutellum is clearly reticulated on a few species ( Crystallotesta leptospermi (Maskell), C. ornatella , K. perforata and both Pounamococcus species, Fig. 68, 70, 77, 85, 86) and this is considered to be of taxonomic importance.

The scutellum appears rectangular in dorsal view but, in fact, is usually more or less tubular, with the anterior and posterior margins having turned inwards and fused, leaving a medial foramen, which may be small (as on K. perforata (Fig. 77) and some Ctenochiton spp.), quite large (as apparently on some Umbonichiton spp.), or the foramen may be absent, the margins actually fusing, as perhaps on Lecanochiton and Pounamococcus spp. Extending posterolaterally from each side of the scutellum is a strongly sclerotised ridge, the posterior notal wing process ; this is particularly pronounced on Lecanochiton species (Fig. 78, 79).

Immediately posterior to the scutellum is a large more or less triangular membranous area, bordered posteriorly by the sclerotised mesopostnotum . The mesopostnotum is almost U-shaped, each arm extending anteriorly, where it becomes the postalare . Posteriorly, the mesopostnotum extends internally under the metathoracic metapostnotum , forming the mesopostphragma . Anterolaterally are a pair of strongly sclerotised mesopostnotal apophyses . Sometimes the whole of the mesopostnotum is mildly reticulated (e.g., C. ornata , Fig. 69, and (rather faintly) I. patella , Fig. 75). The postalare also extends anteriorly to articulate with the mesopleural ridge; anterolaterally, the postalare is often densely reticulated but these can sometimes take the form of punctations (e.g., on I. patella , Fig. 75) or the reticulations, etc., may be absent, as apparently on Lecanochiton actites Henderson & Hodgson (Fig. 78). Rarely setae, the postalare setae , are present on the anterior margins of the postalare and in the general area of the mesopleural apophyses (e.g., on Plumichiton spp., Fig. 80-83).

Ventrally, the main structure on the mesothorax is the meso - or basisternum , which is large and extends across the full width of the segment. Anteriorly, it is demarcated by the marginal ridge and posterolaterally by the precoxal ridges ; the marginal and precoxal ridges fuse laterally and then join the mesopleural ridge just before it articulates with the mesothoracic coxae. The basisternum is usually divided down the middle by a strongly sclerotised median ridge , although this can be poorly defined or even absent (e.g., on I. patella ). Basisternal setae are usually absent on New Zealand males, but are present at the base of the median ridge on C. ornatella (Fig. 70). Laterad to the marginal ridge is a narrow sclerite, the anterior part of which is the lateropleurite , which may be quite narrow or fairly broad and may or may not have a thin extension from the marginal ridge along its anterior margin; it is also occasionally reticulated (e.g., Plumichiton nikau Henderson & Hodgson, Fig. 82). The posterior margin of the basisternum is invaginated, forming the furca ; from this 2 further invaginated arms extend anteriorly; these arms are fairly narrow, diverge strongly, and can vary in length between species (rather short on I. patella , Fig. 75, and long on Lecanochiton scutellaris Henderson & Hodgson and P. flavus : Fig. 79, 81). The base of the furca is rather broad on most species of Coccidae and has a "waist" between the base and the lateral arms.

Laterally on the mesothorax are a number of structures, many of which are hard to discern on dorsoventrally mounted specimens. The mesopleural ridge arises from near the postalare and passes ventrally to articulate with the mesothoracic legs, being joined near its ventral end by the fused marginal and precoxal ridges of the basisternum. Also laterally, more or less at the anterior end of the mesopleural ridge is the episternum , which is a large sclerite just anterior to the wing articulations. It is divided into 2 parts, a dorsal part that is strongly sclerotised and sometimes reticulated (as on P. dacrydii , Fig. 84) and a ventral part that is attached laterally to the marginal ridge on the basisternum, fusing with the lateropleurite anteriorly. The dorsal part has a strong sclerotised ridge running along its anteroventral margin, the subepisternal ridge , which extends more or less to the lateropleurite ventrally.

The mesothoracic spiracles are placed just posterior to each procoxa and laterad to the prosternum. Lying between the basisternum and the prosternum is a narrow membranous area which bears the postmesospiracular setae ; these are mainly fs and, when abundant, they extend the full width of the segment, as on most Plumichiton spp. They may be absent, however, as on C. ornata and I. patella (Fig. 69, 75), very few (as on Lecanochiton spp.) or they may be more or less restricted to just posterior to each spiracle (as on U. jubatus , Fig. 90).

In addition, dorsolaterally, there are 3 or 4 sclerites, the axillary sclerites , which form the articulation for the wings, but these are small and generally difficult to see properly but appear to show little variation between species.

The pores on P. dacrydii (Fig. 84) are present on the membranous area of the scutum, within the groups of tegular setae and on the membranous area between the scutellum and mesopostnotum.

Metathorax : due to the reduction or absence of the hindwings, the metathorax is largely membranous. Dorsally, on many Coccoidea, the metapostnotum is represented by a pair of small sclerites which largely overlie the mesopostnotum, but these are absent on most New Zealand Coccidae. On each metapostnotum, there is 1 or more setae, the metatergal setae ; these are usually represented by a single pair of hs but, on P. dacrydii (Fig. 84), there is a group of hs and, on some Plumichiton spp., there are also groups of fs. Metatergal setae are apparently absent on some species (e.g., Aphenochiton matai Henderson & Hodgson, Fig. 63).

Laterally, the structure depends on whether hamulohalteres are present or not. On all species, the metapleural ridge extends from the metacoxal articulation anterodorsally; this ridge is short on species lacking the hamulohaltere (most New Zealand species) but is long on the 2 Pounamococcus spp. (the only New Zealand species on which hamulohalteres are present, Fig. 85, 86) and extend to the base of the hamulohaltere (Fig. 17). Just dorsad to this wing process is another small sclerite, the suspensorial sclerite , which is connected to the hamulohaltere. Near the metacoxal articulation, the metapleural ridge may have 2 areas of sclerotisation: (a) one extending ventrally from the pleural ridge, the metepisternum , which is usually approximately triangular in shape (but not always sclerotised), and (b) another extending posteriorly just dorsad to the dorsal margin of the metacoxa, which is the metepimeron . The episternum usually carries some postmetaspiracular setae , which are mainly fs but may include an occasional hs. The epimeron may occasionally also have fs setae, the metepimeron setae , as on U. bullatus (Fig. 88).

Also laterally, between the metatergal setae and the wing sclerites near the margin of the metapostnotum, are the dorsospiracular setae ; these are more or less in line with the dorsopleural setae laterally on the abdomen. When present, they are generally rather few and fs, but, on these dorsoventrally mounted specimens, can be very hard to separate from the antemetaspiracular setae which, when present, are just anterior to each metathoracic spiracle and are also apparently always fs. Dorsospiracular setae are considered to be present on most New Zealand males.

The metathoracic spiracles are found ventrally, apparently lying beneath the mesocoxae; they are similar in size and shape to the mesothoracic spiracles.

The metasternum is rarely sclerotised on New Zealand species, and only lightly when present, and there are no metasternal apophyses. The metasternum is divided into an anterior and posterior half, each of which usually bears a group of setae: the anterior metasternal setae and the posterior metasternal setae , both usually fs. However, setae may be missing from both areas (as on I. patella and Pounamococcus spp., Fig. 75, 85, 86, although these species can have hs instead); occasionally only the posterior metasternal setae are absent (as on C. fagi and Species A, Fig. 67, 92).

The dorsal pores on P. dacrydii (Fig. 84) are found along the margin of the metapostnotum , associated with the metatergal and dorsospiracular setae .

Wings : the fore wings are large and quite broad although narrow basally, with a broadly rounded apex. They are usually about 2´ as long as broad and about 4/5 of the length of the body, but are longer on some species (c.f. Pounamococcus tubulus Henderson & Hodgson) and shorter on others (e.g., C. fagi and C. ornatella ). On Pounamococcus species, on which hamulohalteres are present, a small lobe, the alar lobe is present on the hind margin, and this provides a structure for the hamulus or haltere seta to hook onto; when the hamulohaltere is absent, this lobe is lacking. The surface of the wing is covered in small hairlike microtrichia . Only 2 distinct veins are present, the radius , which runs along parallel to the anterior margin, and the media , which runs more posteroventrally. Alar setae are present on some non-indigenous soft scale species but appear to be absent from all New Zealand males.

The hamulohalteres of P. cuneatus and P. tubulus (Fig. 85, 86) articulate with the suspensorial sclerite (Fig. 17); the hamulohalteres are mildly sclerotised, with a single hamulus or hooked haltere seta apically.

Legs : the legs are normal insect legs but with a single tarsal segment on most species (2 segments on Pounamococcus spp., Fig. 17) and a single claw. On New Zealand species, the anterior pair of legs is sometimes marginally the longest. All legs are covered in fs and hs setae, the former usually being the most frequent but, on P. dacrydii (Fig. 84), hs are more frequent. On most species, the coxae have 1 or 2 elongate setae on or near their apex on the inner margin that are here referred to as long coxal setae . A long hs is also present on each trochanter near the segmental membrane separating it from the coxa and this is here referred to as the long trochanter seta ; these can vary considerably in length between species. Also on each trochanter is a ring of 6 campaniform sensilla .

The relative length of the tibia and tarsus varies between species, some species have the tibia about 1. 6´ the length of the tarsus, others up to 2.5 ´ . Both the tibia and tarsus are covered in fs and hs but the setae tend to become spurlike more distally; in addition, with the exception of Pounamococcus spp. which have 2 (Fig. 85, 86), each tibia of New Zealand species has 1 strong tibial spur on its inner margin near the tarsus (Fig. 17). Similarly, on the tarsus, one of the setae on the inner margin next to the claw is often particularly long, strong, and spurlike, and is here referred to as the tarsal spur (Fig. 17). In addition, on the outer margin at the proximal end of each tarsus on Pounamococcus spp. (Fig. 17, 85, 86), there is a tarsal campaniform pore ; these are absent on all other known coccid males. On the distal end of each tarsus is a pair of capitate tarsal digitules (Fig. 17), which are usually slightly shorter than the length of the claw.

The single claw is usually held more or less at right angles to the tarsus, slightly curved and pointed; on New Zealand males, the small denticle near the apex, typical of many other coccids, appears to be absent or very indistinct. Each claw has a pair of capitate claw digitules , which are slightly longer than the claw (Fig. 17).

ABDOMEN

The abdomen is elongate, more or less parallel-sided and slightly tapering, consisting of 8 membranous segments and the genital segment (IX), which is elongate and narrow, partly sclerotised, and carries the genital organs. The segmentation is usually reasonably clear and most segments have a broad band of microtrichia on both the dorsal and ventral surfaces. Segments VII and VIII can be produced laterally to form well developed lobe-like caudal extensions on some male coccids, but these are short or absent on most New Zealand coccids. Also, segment VIII often has a pair of invaginated glandular pouches , each with 2 long glandular pouch setae , which support the long wax filaments found posteriorly on many live males. The glandular pouch and setae may be present or absent on New Zealand species, even within the same genus.

Segments I-VII : segment I is developed dorsally and pleurally but not ventrally. Segments I-VI are more or less membranous on New Zealand species, but segment VII usually shows some sclerotisation on the tergite and more particularly the sternite . On P. cuneatus (Fig. 85), all segments show some degree of reticulation; this is much less obvious on P. tubulus (Fig. 86). No pleural sclerites are present on segments I-VI on indigenous species. The caudal extension of segment VII is usually rounded, and could even be described as absent on most species, but is obvious on C. ornata , C. ornatella , E. piperis , P. tubulus , P. dacrydii, and Species A. The caudal extension on segment VII is lightly sclerotised on Pounamococcus species (Fig. 85, 86).

Each segment usually bears some dorsal abdominal setae , ventral abdominal setae , dorsopleural setae, and ventropleural setae . The dorsal abdominal setae usually consist of 1-3 pairs of hs on each segment, but many species also have some fs, particularly C. ornata and C. ornatella (Fig. 69, 70) and most Plumichiton spp. Ventral abdominal setae occur in a transverse line or band across each segment; when least abundant, there may be only 1 pair of hs but, when abundant (as on Plumichiton spp.), there are many fs and then there are always more fs on the ventral than on the dorsal surface. The dorsal and ventral pleural setae can be hard to differentiate on mounted specimens and can, in any case, run into each other when abundant. The dorsopleural setae may be all hs or varying combinations of fs and hs; the ventropleural setae are generally fewer and are mainly hs.

On I. patella, medially on the tergite of segment IV, are a pair of round, membranous cicatrices (Fig. 75). Cicatrices have been noted on a number of species on or near the apex of the caudal extension of segment VIII (see Giliomee 1967; Miller 1991) but not previously from elsewhere on the body.

Segment VIII : is usually similar in length to the preceding segments but is noticeably longer on C. ornata and C. ornatella (Fig. 69, 70). Both the tergum and the sternum of segment VIII are clearly sclerotised; the caudal extension is usually rounded and, like that for VII, could be described as absent on some species, but is most obvious (although still small) on I. patella and Plumichiton elaeocarpi (Maskell) (Fig. 75, 80); it is mildly sclerotised on Aphenochiton subtilis Henderson & Hodgson (Fig. 66). The tergite bears the ante-anal setae: on many species, these are represented by a single pair of hs, but on others (e.g., C. ornata , C. ornatella, Fig. 69, 70, and most Plumichiton spp.), there are many fs, and these can appear to be in 2 groups, one group on the anterior half and the other on the posterior half of the tergite, just anterior to the anus. When abundant, these may merge with the pleural setae laterally on the caudal extension.

Segment VIII also carries the glandular pouch and glandular pouch setae , when these are present. The glandular pouch is usually quite deep and contains 2 kinds of pore : those at the base of the pouch are not loculate, appearing like small cones (e.g., on Fig. 70, 80), whereas those near the opening (and sometimes outside the pouch) are loculate. Both pore kinds appear to be openings to small tubular ducts (cf. Fig. 70, C. ornatella ); these ducts were also described by Sulc (1931). The frequency of each kind of pore varies between species. The glandular pouch setae are setose on New Zealand species (although they can be capitate on other species) and one is generally noticeably shorter than the other. Even within a genus of what appear to be closely related species (e.g., Plumichiton spp.), glandular pouches may be present on some species but absent on others, and thus the presence or absence of glandular pouches appear not to be an apomorphic character for the genus.

Ventrally, the sternite of VIII may or may not carry ventral abdominal setae ; when present these are generally fs. In addition, there is usually a group of 3 hs just laterad to the glandular pouch; on many species, there may be additional fs pleural setae. When the glandular pouch is absent, 1 or more of these setae may be quite long (e.g., on A. subtilis , Fig. 66).

Genital segment : this is considered to represent segment IX. This consists of a (generally) long tubular structure that extends posteriorly and is referred to as the penial sheath , while dorsomedially, where the penial sheath joins segment VIII, is the anus although this is not usually visible. The penial sheath emerges from beneath segment VIII and varies somewhat in length, but is generally between 1/3 and 1/6 of total body length. It is broadest anteriorly, tapering posteriorly more or less to a point, although the apex is very blunt on some species (e.g., Lecanochiton spp., Fig. 78, 79); on Plumichiton spp. the penial sheath is distinctly constricted near the apex. The penial sheath is well sclerotised laterally but membranous ventromedially (Giliomee, 1967), with a slit ventrally towards the apex, through which the aedeagus emerges. The slit widens at the anterior end into a large membranous area, which is referred to as the basal membranous area ; this is approximately triangular and does not vary much in size or shape. Between the 2 sclerotised margins of the penial sheath just posterior to the basal membranous area, is a sclerotised ridge, the basal rod ; this may be very short or rather long and may or may not reach the basal membranous area anteriorly; the basal rod of Lecanochiton spp. (Fig. 78, 79) appears to be made of parallel ridges. Posteriorly, the basal rod is attached to the anterior end of the aedeagus , which lies in a groove in the ventral wall of the penial sheath; the penial sheath generally ends in a blunt apex some distance from the end of the penial sheath but, on Species A, it appears to have a distinct "shovel-shaped" apex which extends posteriorly past the end of the sheath (Fig. 92). The length of the aedeagus is very variable between species.

Along the sides and near the apex of the penial sheath are a number of small setae, the penial sheath setae . In addition, towards the apex of the sheath are a group of minute pores, the penial sheath pores , which may be campaniform sensilla.

Comment . The taxonomic significance of male characters for defining higher taxa is becoming clearer. Some characters that had appeared likely to be of considerable importance are now considered of lesser significance. Two good examples of such characters are the number of simple eyes and the presence or absence of glandular pouches and glandular pouch setae. With regard to the number of simple eyes, Pounamococcus cuneatus has 2 (Fig. 85) and P. tubulus has 4 (Fig. 86). As there are numerous other significant characters shared by these two species, they are clearly congeneric. And with regard to glandular pouches, there are several genera where some species have them whilst others do not, i.e., Aphenochiton , Plumichiton, and Umbonichiton . Nonetheless, there are many characters that appear to be good for defining groupings here considered to represent genera, some apparently relatively insignificant. Thus, the males of three Ctenochiton species described here (Fig. 71-73) all have 1 or more setae on the metathorax immediately posterior to where the marginal ridge of the basisternum joins the precoxal ridge; these setae are absent from almost all other known male Coccidae (only otherwise known on U. hymenantherae , Fig. 89 and Kalasiris depressa , Fig. 76). Equally, all the known males of Plumichiton species have a trochantofemur (Fig. 80-83), where the segmentation between the trochanter and femur is poorly defined or entirely absent; in addition, Plumichiton species also have a distinct constriction towards the apex of the penial sheath. Pounamococcus species have tarsal campaniform pores, 2-segmented tibiae, 2 tibial spurs per leg, and a tripartite cranial apophysis; male Lecanochiton have a poorly defined basal rod (Fig. 78, 79); and the ornata -group of Crystallotesta have no tentorial bridge (Fig. 69, 70), an unusually long abdominal segment VIII, and the postocular sclerite extends around the ocelli (although this is shared with P. cuneatus , Fig. 85).

With regard to useful characters that might help in identifying species rather than higher taxa, the main new character used here is the structure of the dermal reticulations on the head. It was noted that the size and, more particularly, the structure of the inner microridges within each reticulation seemed to be highly variable between species but relatively uniform within a species; Fig. 62-92 show this structure for the gena and ocular sclerite for each species. Again, even within the genera as defined here, these characters can be reasonably constant: thus, Ctenochiton species (Fig. 71-73) tend to have large genal reticulations with very few or no inner microridges. In addition, the frequency of fleshy and hairlike setae on the metasternum, ventrally on the abdomen, and on the membranous area of the scutum appears to be important, particularly at the species level. Pores are only known from 2 species in New Zealand, I. patella (on the head only, Fig. 75) and P. dacrydii (all over the dorsum, Fig. 84), but the structure of these pores is quite different on the two species.

Important taxonomic characters

Characters that appear to be of help in diagnosing species and higher taxa of New Zealand Coccidae are therefore the following (where p/a = present/absent).

General :

overall size.

Head:

p/a dorsal midcranial ridge;
p/a postoccipital ridge;
p/a fleshy setae;
p/a dorsal ocular setae;
p/a genal setae;
p/a ventral head setae between or posterior to ventral eyes;
whether postocular ridge reaches ocelli;
form of genal and ocular sclerite reticulations;
p/a tentorial bridge;
p/a pores;
shape of cranial apophysis;
number of simple eyes.

Antennae:

overall length;
number of setae on scape;
length and frequency of fleshy setae;
p/a of hairlike setae on segments III-IX;
number of capitate setae;
relative lengths of each segment.

Prothorax:

p/a fleshy setae;
p/a lateral pronotal setae;
p/a pores;
p/a median ridge of prosternum;
frequency and kind of prosternal setae;
p/a anteprosternal setae;
p/a antemesospiracular setae;
p/a post-tergite.

Mesothorax:

shape of membranous area of scutum;
frequency and kind of setae on membranous area;
p/a reticulations on scutum laterad to scutellum;
p/a tegular setae;
frequency and distribution of postmesospiracular setae;
p/a reticulation on mesepisternum;
p/a median ridge on basisternum;
p/a basisternal setae;
p/a setae on/near lateropleurite and posterior part of mesepisternum;
p/a postalare setae.

Metathorax:

frequency and kinds of metatergal setae;
p/a dorsospiracular setae;
p/a anterior and posterior metasternal setae;
p/a postmetaspiracular setae;
p/a setae just posterior to where marginal ridge of basisternum and precoxal ridge meet;
p/a dorsal part of pleural ridge.

Wings:

p/a hamulohalteres;
ratio of wing length to width.

Legs:

whether tarsi 1- or 2-segmented;
p/a trochantofemur;
number of tibial spurs;
frequency of both hs and fs;
p/a long hairlike seta on trochanter;
p/a tibiotarsal articulation;
p/a tarsal campaniform pores.

Abdomen :

frequency and kind of setae;
p/a tergites and sternites;
p/a pleurites;
p/a cicatrix;
p/a fleshy ante-anal setae;
p/a glandular pouch;
size of caudal extensions on segments VII and VIII;
length of segment VIII.

Genital segment:

shape of penial sheath;
relative length of penial sheath to total body length;
shape of aedeagus;
whether basal rod reaches basal membranous area;
length of basal rod;
shape of aedeagus apex.

Purchase this publication

Purchasing details