Matsucoccus josephi Bodenheimer & Harpaz, 1955

Hodgson, Chris & Foldi, Imre, 2006, A review of the Margarodidae sensu Morrison (Hemiptera: Coccoidea) and some related taxa based on the morphology of adult males, Zootaxa 1263 (1), pp. 1-250 : 1-250

Matsucoccus josephi Bodenheimer & Harpaz

( Fig. 7 )

Matsucoccus josephi Bodenheimer & Harpaz, 1955: 12 .

Material studied

ISRAEL, Bet Dagan, on Pinus halepensis , 2002, Y. Ben­Dov ( BMNH): 18/18ad ♂♂ (described mainly from four specimens in fair to good condition but most mounted on their sides, plus another 14 in good to poor condition, also mounted on their sides but including 2 pharate adults mounted dorsoventrally) .

Mounted material

Fairly small, 1.6–1.75 mm long; width very narrow when viewed dorsally or ventrally, only 300–320 m wide across prealare; body deep and very difficult to mount dorsoventrally; basisternum unusually elongate, with anterior margin lying close to point below anterior margin of prescutum. Body with very few setae, these all hair­like (hs) with well­developed, shallow basal sockets; most about 8–10 m long; hairs (hrs), collared setae (cs), loculate pores (lp) and convex pores (cp) absent. Antennae long, flagellar segments all quite narrow, parallel­sided, becoming slightly shorter near apex; antennae with fleshy setae (fs) but without satellite setae (sats); most segments with capitate setae (caps) and hs, and terminal six segments with antennal bristles (ab). Sclerotised areas without nodulations or reticulations. Legs well developed and setose, with many (apparently) fleshy setae (fs) but no bifurcated setae (bs); tarsi 1 segmented; claws with a strong denticle proximally; claw digitules clavate. Abdomen with segment I visible ventrally; without lateral caudal extensions; terga of abdominal segment VII with a group of large tubular ducts (tdc), each internally ridged; tubular ducts absent from segment VI. Penial sheath short, more or less terminal, with a long narrow aedeagus but without an eversible endophallus.

Head

Short but very wide in dorsal view, length 130–160 m, width across compound eyes 380– 420 m. Dorsally: dorsomedial part of epicranium (dmep) mildly sclerotised across posterior half, with a narrow postoccipital suture (pos) posteriorly, extending full width of head; postocciput (poc) probably absent. Midcranial ridge (mcr) narrow, extending full length of dorsal part of epicranium (dmep), fusing with pos posteriorly. Preocular ridge (procr) very short or absent dorsally. Dorsal epicranium (dmep) with 2 or 3 short hs on each side. Laterally with a pair of large compound eyes (cde), each about 200–230 m long, with about 150–170 ommatidia, each ommatidium about 15 m wide. Each cde with a narrow, sclerotised ocular sclerite (ocs) along posterior margin, each with a single ocellus (o) protruding laterally; width of each ocellus 25–30 m; ocular sclerites without a distinct postocular ridge (pocr) along dorso­lateral margins, but with a short sclerotised ventral projection (p) extending postero­medially to articulate with cervical sclerite (cv). Ventrally: ventromedial part of epicranium (vmep) with a large, mildly sclerotised area medially; vmep membranous anteriorly and laterally, with a short, heavily sclerotised ventral extension of midcranial ridge (vmcr) but also with a lightly sclerotised extension posteriorly on some specimens; without lateral arms to mid­cranial ridge (lmcr); preocular ridges (procr) absent; preoral ridge (pror) suggested by a slightly more heavily­sclerotised transverse ridge about half­way along vmep, which extends almost to (but not actually reaching) compound eyes laterally. Setae: with 1–5 short hs on each side of vmep near anterior margin. Cranial apophysis (ca) absent. Rudimentary mouth (m) present medially, almost in neck region, surrounded by a mildly sclerotised ventral plate (vp); and with another narrow plate (post­oral plate (pop)) posterior to ventral plate. Anterior tentorial arms (ata) arise just postero­laterally to preoral ridge (pror), with tentorial arms extending posteriorly; posterior tentorial arms (pta) possibly present on either side of posterior margin of post­oral plate (pop); tentorial bridge (tb) not detected.

Antennae: 10­segmented; length 1.4 mm (ratio of total­body length to antennal length 1:0.84). Scape (scp) 53–70 m long, 75–80 m wide, sclerotised, with a sclerotised articulatory socket with head; with, on dorsal surface, 8 or 9 very short setae, each about 3–5 m long, plus 1–3 slightly longer setae (each about 8 m long). Pedicel (pdc) 27–40 m long, 50–55 m wide; with 4 or 5 short setae plus one much longer seta on ventral surface, about 50 m long, plus 3 campaniform sensilla (camp) on dorsal surface. Segments III–IX of flagellum parallel­sided or each widening slightly distally; each about 25–35 m wide; surface of segments with transverse ridges; each segment with many fleshy setae (fs), each 35–45 m long, but without satellite setae (sats). Segment lengths (m): III 150–160, IV 185–190, V 170–180, VI 185–195, VII 170–185, VIII 155–160, IX 135–145 and X 115–120. Setal distribution: III 18–20 fs, IV 21–26 fs, 2 hs + 2 capitate setae (caps); V 42 or 43 fs, 2 hs + 2 caps; VI 33–35 fs, 2 hs, 1 antennal bristle (ab) + 1 or 2 caps; VII 36–39 fs, 1 hs, 2 ab, 2 or 3 caps + 1 coeloconic sensilla (cos); VIII 38 or 39 fs, 1 hs, 2 ab, 2 or 3 caps + 1 coeloconic sensilla (cos); IX 31 fs, 0 hs, 2 ab + 2 or 3 caps, and X 17–21 fs, 0 hs, 2 ab + 5 caps (when present, hs, ab and caps restricted to distal end of each segment).

Thorax

Prothorax: separated from head by a distinct, shallow neck. Dorsally without a pronotal ridge, lateral pronotal sclerites or other signs of pronotum (prn); also without lateral pronotal setae but with a line of 35–45 median pronotal setae (mpns) immediately anterior to prescutum between post­tergites. Post­tergites (pt) longitudinal, lightly sclerotised and elongate, each about 110 m long. Ventrally with a pair of strong cervical sclerites (cv) which possibly articulate anteriorly with a ventral projection (p) from ocular sclerite. Pleural ridge (plr 1) well developed and quite long, extending dorsally from articulation with coxa and fusing with cv posteriorly. Proepisternum (eps 1) well developed between cervical sclerite (cv) and pleural ridge (plr 1). Propleural ridge (plr 1) rather long, extending ventrally. Prosternum (stn 1) with a lightly sclerotised median ridge about 270 m long, which becomes slightly more sclerotised anteriorly; with a single sternal apophysis (stn 1 a) medially; without a transverse ridge. Ventrally without setae or pores.

Mesothorax: dorsally: prescutum (prsc) roundly diamond­shaped (length 225–275 m, width 260 m); mesoprephragma (phr 1) narrow; prescutal ridges (pscr) extending anteriorly to mesoprephragma (phr 1); prescutal sutures (pscs) well sclerotised; without prescutal setae (prscs). Scutum (sct) unusually narrow, sclerotised, without nodulations and without a median membranous area; distance from prescutum to scutellum 125–190 m; scutal setae (scts) absent. Scutellum (scl) almost diamond­shaped although lateral margins occasionally nearly parallel, separated from scutum by weak but distinct scutoscutellar sutures (scuts), which curve posteriorly, fusing with a strong ridge (rd) along posterior margin; scutellum quite deep, posterior margin curving ventrally; scl without either membranous areas laterally or scutellar setae (scls) or pores. Mesopostnotum (pn 2) lying almost vertical, appearing narrow in dorso­ventral view but obviously quite broad when viewed laterally, extending around laterad to scutellum, forming a deep mesopostphragma (phr 2) posteriorly; without mesopostnotal apophyses (pn 2 a); membranous area posterior to scutellum narrow. Laterally: prealare (pra) elongate and broad. Tegula (teg) large and sclerotised but without tegular setae (tegs). Mesopleural ridge (plr 2) well developed, extending vertically from mesocoxal articulation; pleural apophysis (pla 2) apparently absent. Mesepisternum (eps 2) not nodulated. Mesothoracic spiracles (sp 2) very small and only lightly sclerotised, peritremes 17–25 m broad; each muscle plate rather poorly sclerotised, narrow, about 45 m long. Ventrally: basisternum (stn 2) unusually elongate and narrow, anterior margin approximately level with anterior margin of prealare; length 340–360 m, width uncertain but slightly greater than width of prescutum; without a sclerotised median ridge (mdr) and without basisternal setae (stn 2 s); some specimens with derm quite distinctly nodulated; bounded anteriorly by a fairly welldeveloped marginal ridge (mr) which extends round posteriorly along lateral margins, where sides almost parallel; bounded posteriorly by well­developed precoxal ridges (pcr 2); furca (f) very narrow ventrally, with rather short arms, which diverge strongly; lateropleurite (lpl) distinct, without an extension from marginal ridge; subepisternal ridges (ser) only present between each lateropleurite and mesepisternum; without lateropleurite setae (lpls) in this area. Postmesospiracular setae (pm 2 s) absent. Wing sclerites: apparently similar to other Coccoidea : costal complex of veins (ccx) distinct; axillary sclerites showing nothing distinctive.

Metathorax: dorsally: metapostnotum (pn 3) present as 2, fairly small, mediolateral areas of sclerotisation; with a band of 5–11 metatergal setae (mts) on each side. Laterally: suspensorial sclerites (ss) present. Pleural ridge (plr 3) well developed; precoxal ridge (pcr 3) well developed and extending about 140–155 m medio­ventrally; with a well­developed pleural apophysis (pla 3). Metepisternum (eps 3) lightly sclerotised; postmetaspiracular setae (eps 3 s) absent; metepimeron (epm 3) represented by a strong sclerotisation extending dorsoposteriorly around metacoxae. Posterior spiracles (sp 3) similar in structure and size to anterior spiracles. Ventrally: metasternum (stn 3) quite well developed, with a single, media, circular metasternal apophysis (stn 3 a) and 2 well­developed metafurca. Anterior and posterior metasternal setae absent. All other setae and pores absent.

Wings: quite large and well developed; each 1.6–1.7 mm long, 0.6–0.7 mm wide (ratio of length to width 1:0.39; ratio of wing length to body length about 1:1). Wings with well­developed subcostal thickening (sclt), radius (rad) and media veins (med) but anal fold vein (af) very faint. Surface of wing with numerous fine parallel folds, mostly running radially from wing margin, where they form a discrete reticulated pattern. Wing anterior to radial vein (rad) also with fine folds at right angles to margin but also well sclerotised proximally, becoming less so towards wing tip; with a distinct group of 7–9 circular sensoria (sens) near base of subcostal thickening; other alar pores and alar setae (als) absent. Alar lobe (al) very narrow, represented by a fold along proximal posterior margin. Hamulohalteres (h) mainly sclerotised, long and narrow, broadening abruptly distally; length 170–210 m, greatest width 40–45 m; with 3 or 4 hamuli (ham) along anterior margin of swollen distal end, each quite long and highly curved, with a large basal socket and a clavate apex.

Legs: prothoracic legs longest. Coxae (cx): I 165–190; II 145–160; III 135–145 m; each with 8–10 very short setae (similar to those on scape) near base and 22–27 fs + 5–7 longer hs more distally. Trochanter (tr) + femur (fm): I 320–345; II 275–295; III 315–325 m; trochanter with 8–14 fs + 2 hs; long trochanter seta not differentiated but seta in this position perhaps 26–33 m long; each trochanter with a group of 4 oval campaniform sensilla (camp) on each side, most pores lying within a groove but usually with 1 present elsewhere; trochanterofemur articulation clear and diagonal; femur with about 30–40 fs + 3–5 hs. Tibia (ti): I 405–420; II 365–370; III 375–390 m; each with 39–50 setae, those on distal 2/3rds spur­like laterally and ventrally, these replaced by 3–6 longish flagellate setae dorsally; with about 3–5 tibial spurs (tibs) along distal end, longest about 20–25 m long; bifurcated setae (bs) absent. Tarsi (ta) 1 segmented; length: I 140–145; II 125–130; III 125–130 m; each with a series of shallow transverse ridges; without tarsal campaniform sensilla (camp); without bifurcated setae (bs), but with 14–18 spinose setae along ventral and lateral margins and 1 or 2 hs laterally and dorsally; tarsal digitules (tdgt) rather short (11–15 m long) and setose. Claws (c) broad, with a strong pointed denticle (cd) proximally near base; claw III about 28–30 m long, with 2 strong, clavate digitules (cdgt), each longer than claw (about 35 m long), arising very close to base of tarsus.

Abdomen

Almost entirely membranous; segment I present ventrally; without caudal extensions (ce), margins of segment VII and VIII rounded. Tergites (at) present on all segments, represented by pairs of sclerotisations mediolaterally, these becoming smaller posteriorly; sternites (as) and pleurites absent. Dorsal abdominal setae (ads) short, in 2 groups medially on each segment; mostly 5–8 m long; with (hs) on segment I 12–22; II 4–12; III–VI 1–4; VII 3–6 and VIII 0–2. Loculate pores (lp) absent. Pleural setae not clearly divided into dorsal (dps) and ventral pleural groups (vps), with perhaps a total of 2 or 3 hs on segments IV–VIII. Ventral abdominal setae (avs) similar to those on dorsum but perhaps a little longer (8–15 m long), with (totals per segment) I–III 0; IV–VII 1 or 2 and VIII 4 or 5. Dorsally with a plate­like tergite medially on segment VII with a group of 13–18 large tubular ducts (tdc) (each about 6–8 m wide), each pore conical, with a broad inner ductule 25–27 m long with about 8 longitudinal ridges, giving it a corrugated appearance. Abdominal spiracles (asp) present on segments II–VIII, apparently very small and unsclerotised, and placed rather ventrally.

Genital segment: anus (an) hard to discern but present medially over penial sheath.

Penial sheath (ps) elongate triangular, more or less on apex of abdomen; about

215–240 m long and 125 m wide across base, narrowing to a short, blunt apex posteriorly; mainly sclerotised laterally; ventrally with a group of 19–23 small setae medially (each about 8–10 m long) anterior to base of aedeagus (aed), but setae absent or minute more posteriorly; with a few small sensoria (psp) at apex. Aedeagus (aed) narrow, 210–240 m long, longer than penial sheath, sclerotised and bent ventrally; with a short basal rod (bra) at anterior end of aedeagus, length perhaps 45 m; eversible endophallus (eph) absent.

Comment

The adult male of M. bisetosus Morrison was redescribed by Beardsley (1968) and the above description is very similar. Beardsley provided a long discussion in which he compared the structure of M. bisetosus with the descriptions of adult male Margarodes , Pseudaspidoproctus and Steingelia given by Theron (1958). Beardsley’s discussion is excellent and the points that he made will not be repeated here. In addition to the structures he covered, there are a few others that seem to be worthy of comment.

Firstly, the antennae have fleshy setae, hair­like setae, capitate setae and antennal bristles but no satellite setae. This combination appears to be absent from all margarodoids (and aphids) but is present on many neococcoids suggesting, perhaps, a close link with the neococcoids.

Secondly, the basisternum on Matsucoccus appears to be very long, extending anteriorly to a point below the prescutum whereas, on all other male coccoids, it only extends anteriorly to about the middle of the scutum. This anterior extension of the basisternum closely resembles the condition on aphids (see Fig. 1 ), where the anterior border of the prescutum overlies that of the basisternum. This is surely, therefore, a primitive condition.

In addition, the mesopostnotum on Matsucoccus appears to be very short when viewed from above but actually is quite long, curving down at a sharp angle posteriorly to form the mesopostphragma. Whilst a mesopostphragma is present on all coccoids, the mesopostnotum usually appears rather broad, only extending below the metatergite at some distance posterior to the scutellum. In appearing narrow, and lying rather vertically, the mesopostnotum on Matsucoccus more closely resembles that on the Aphididae than other coccoids.

Thirdly, one character­state which appears to be constant for male Margarodidae , Xylococcidae and Neosteingelia sp. but which is absent from the Monophlebidae and Coelostomidiidae is the group of very small setae on the dorsal surface of the scape and on the base of all the coxae. These groups of setae are also present on Matsucoccus , suggesting a closer relationship to the first two families than to the Monophlebinae and Coelostomidiinae .

Other characters which appear to be characteristic of the Matsucoccidae are: (i) what are considered here to be fleshy setae on all legs; (ii) a single tarsal segment; (iii) presence of clavate claw digitules; (iv) trochanter campaniform sensilla more or less in a line and in a shallow groove; (v) each wing with a narrow alar lobe along wing margin; (vi) a single group of tubular ducts present dorsally on abdominal segment VII; (vii) abdominal spiracles small and poorly developed on segments II–VIII; (viii) thoracic spiracles poorly developed; (ix) hamulohalteres with hamuli along anterior margin; (x) aedeagus long and thin; (xi) absence of eversible endophallus; (xii) wing with a group of circular sensoria near base of costal vein; (xiii) wings with a series of parallel ridges; (xiv) derm of legs and antennae with transverse ridges (as on Steingelia ), and (xv) absence of tarsal campaniform sensilla.

At the time, Beardsley (1968) discussed at considerable length the possible relationships of Matsucoccus to aphids and other Coccoidea , based on adult male characters, and he concluded (p.1458): 1. that the male of Matsucoccus was primitive and not particularly closely related to Margarodes , Xylococcus or to Monophlebidae ; 2. Matsucoccus was most closely related to Orthezia among the Coccoidea he considered in his paper; and 3. that the genus Matsucoccus should be treated as a distinct subfamily (Matsucoccinae) within the Margarodidae rather than as a tribe within the Xylococcinae. On the other hand, Koteja (1974) considered that the mouthparts of Matsucoccus were amongst the most specialised within the Orthezioidea (= Margarodidae ) and could not be considered primitive!

Figures

FIGURE 1. Male aphid (Eucallipterus tiliae (L.) with a few additions) labelled using the terminology used for Coccoidea in this paper (for abbreviations, see Table 2). Where similar structures are absent from Coccoidea, the name of the structure is given in full. Left side dorsal surface, right side ventral surface.

FIGURE 7. Male Matsucoccus josephi Bodenheimer & Harpaz. Letters as for Fig. 6 except P = side view of thorax, N = side view of end of abdomen and U = apical antennal segment.