insect

Watson, Gillian W. & Ouvrard, David, 2021, Towards the identification of the scale insects (Hemiptera: Coccomorpha) of continental Africa: 1. Identification of the families, Zootaxa 5052 (2), pp. 151-190 : 155-157

Key to scale insect families currently known from continental Africa, based on adult females (23 families). Figures 1‒26 are given after the key.

Caution. It is recognised that some entomologists may use this key to identify specimens from outside Africa. In some cases, this could result in misidentifications because the key was not developed to identify scales from other continents, particularly Australia, which has many unique scale insects.

1 (0) Adult female with anterior legs much enlarged and sclerotized (adapted for digging); mouthparts absent; found in soil, feeding on roots. Legless immature cyst stages in soil, forming ‘ground pearls’........................ MARGARODIDAE ( Fig. 1 )

– Adult female with anterior legs not enlarged, similar in size to middle legs, or absent; mouthparts present or absent; found above ground or on roots. Immature stages, if forming legless cysts, found above ground .................................... 2

2 (1) Abdomen with small spiracles present on at least posterior segments, numbering 2–8 pairs..............................3

– Abdominal spiracles absent................................................................................6

3 (2) Ovisac band on venter of abdomen composed of spines. Antennal terminal segment with a stout spine-like apical seta. Anal opening situated near or at apex of abdomen, formed of a sclerotised ring bearing 6 setae. Eyes often situated on sclerotized stalks...................................................................................... ORTHEZIIDAE ( Fig. 2 )

– Ovisac band on venter of abdomen, if present, composed of pores. Antennal terminal segment without a stout spine-like apical seta. Anal opening, if visible, situated near apex of abdomen or on dorsum at some distance from abdominal apex. Sclerotised anal ring, if present, situated at inner end of an anal tube, not on body surface; anal ring lacking setae but sometimes with pores. Eyes never on sclerotized stalks................................................................................ 4

4 (3) Antennae with bases separated by more than width of antennal segment I; each antenna with 8‒11 segments and with basal segment length up to about 2x longer than other segments. Mouthparts usually well-developed but occasionally degenerate. Ovisac or marsupial band, if present, composed of pores. Immature stages not forming legless cysts.... MONOPHLEBIDAE ( Fig. 3 )

– Antennae set close together on head, separated by less than width of antennal segment I; each antenna with 8 or 9 segments, and with basal 1 or 2 segments each more than 2x longer than other segments. Mouthparts often reduced or absent. Ovisac or marsupial band absent. Some immature stages forming legless cysts above ground....................................... .. 5

5 (4) Tarsus 2 segmented, proximal segment short. Coxa apparently divided vertically. Ventral apex of each tibia with stout acute setae. Claw without a denticle. Abdominal spiracles each without pores in atrium. On Pinaceae ...... MATSUCOCCIDAE ( Fig. 4 )

‒ Tarsus 1 segmented. Coxa not divided longitudinally. Ventral apex of each tibia with numerous capitate setae. Claw with a denticle. Abdominal spiracles each with pores in atrium. On Fagaceae , especially Quercus ...... KUWANIIDAE in part ( Fig. 5 )

6 (2) Antenna with 8 segments. Antennal bases situated close together on front of head, separated by less than basal width of antennal segment I. Antennal segments II–VIII much shorter and narrower than antennal segment I, each no more than 1.8x as long as wide, and antennal segment I large,>2x as long as wide and at least 3x as long as segment II. Labium weakly developed; stylets absent. Living in the bark..................................... Kuwania ologostigma De Lotto (Kuwaniidae) ( Fig. 6 )

‒ Antenna with 1–9 segments. Antennal bases normally separated by more than 2x width of antennal segment I, situated on front or venter of head. If present, antennal segments II–VIII slimmer than antennal segment I, each up to 3x as long as wide, and antennal segment I usually <1.5x as long as wide and <1.5x as long as segment II. Mouthparts usually developed and functional. Habitats various including under bark.............................................................................. .. 7

7 (6) With some or all of: dorsal ostioles, ventral circuli, trilocular pores each with a swirled centre, and 1–18 pairs of cerarii. Anal ring usually with 1 to 3 concentric rows of pores.................................................................. 8

– Without dorsal ostioles, ventral circuli, swirled trilocular pores or cerarii. Anal ring with or without pores.................10

8 (7) Claws each with a denticle on plantar surface and often an additional denticle on claw base. Trochanters mostly each with 3 or 4 campaniform sensilla on each surface....................................................... PUTOIDAE ( Fig. 7 )

– Claws each with or without a denticle on plantar surface, but claw base always without a denticle. Trochanters each with 2 campaniform sensilla on each surface (except Lenania prisca De Lotto (Pseudococcidae) with 3 on each surface).............. 9

9 (8) Antennae each with 1–9 segments. Cerarii present. Tri- or bitubular cerores absent. Hind legs normally with translucent pores. Circuli, if present, fairly flat, membranous, and usually not textured. Found on any part of host-plant including roots....................................................................................... PSEUDOCOCCIDAE ( Fig. 8 )

– Antennae with up to 6 segments. Cerarii absent. Tri- or bitubular cerores usually present. Hind legs without translucent pores. Circuli, if present, often each on a raised cone and sometimes textured. Feeding on roots or in moss......................................................................................................... RHIZOECIDAE ( Fig. 9 )

10 (7) Anterior thoracic spiracles conspicuously larger than posterior pair. Dorsum of thorax with a pair of brachia, each bearing a brachial plate. Centre of abdomen with a large dorsal spine. Anal ring surrounded by a ring of fimbriations, lobes or setae, situated on anal tubercle near posterior end........................................................ KERRIIDAE ( Fig. 10 )

– Anterior and posterior thoracic spiracles of similar size. Dorsum without brachia, brachial plates or a large dorsal spine. Anal opening never surrounded by lobes or fimbriations and either situated on dorsal body surface, or at inside end of an eversible tube, sometimes on a posterior apical protruberance ............................................................... .. 11

11 (10) Anal complex oval, heavily sclerotized, situated near middle of dorsum. Anal plates numbering 2, each semicircular and situated on midline, with 1 anterior to anal opening and 1 posterior to it........................ STICTOCOCCIDAE ( Fig. 11 )

– Anal complex usually situated at or near posterior end of abdomen. Anal plate(s) either singular and situated on midline or paired, not semicircular, and situated on either side of anal opening............................................ .. 12

12 (11) Anal cleft present, with a pair of dorsal triangular or rounded anal plates situated at its base, covering anal opening. Inner margins of anal plates usually contiguous, together forming an operculum............................... COCCIDAE ( Fig. 12 )

– Anal cleft, if present, without paired plates at its base, although there may be a single plate in this position. If anal cleft absent, a pair of anal plates may be present, with one on each side of anal opening.......................................... 13

13 (12) Antenna 3 segmented. Sclerotised anal ring flanked by paired curved anal plates or a ∩-shaped sclerotised crescent. Largest tubular ducts each with an invaginated, cup-shaped inner end. Vulva located on abdominal segment VI. On grasses.......................................................................................... .. MICROCOCCIDAE ( Fig. 13 )

– Antenna with 1–9 segments, almost never with 3. Anal opening with or without a sclerotised anal ring; if ring present it is associated with anal plates, these situated anterior or posterior to anal ring, not flanking it. Largest tubular ducts with inner end sometimes cup-shaped. Vulva located between abdominal segments VII and VIII. Hosts various including grasses......... 14

14 (13) Abdomen with furrows and ridges along posterior margin. Short anal cleft usually present, with a single triangular or oval anal plate situated at anterior end; occasionally anal plate split longitudinally almost to its anterior margin. Anal opening leading to an eversible anal tube, inner end with an anal ring bearing 10 or more setae. Legs usually absent. Mostly found on grasses........................................................................... ACLERDIDAE ( Fig. 14 )

– Abdomen without furrows or ridges along posterior margin. Anal cleft, if present, without a single, triangular or oval anal plate at anterior end, although a thin sclerotised crescent may be present anterior to anal opening and / or a shield- or butterfly-shaped plate may be present near posterior margin. Anal opening on surface or at inner end of an eversible tube; anal ring present or absent. Legs present or absent. Hosts various, including grasses.........................................................15

15 (14) 8-shaped pores present somewhere on body................................................................. 16

– 8-shaped pores absent, although dorsal bilocular pores may be present.............................................18

16 (15) Cribriform plates present at least on dorsum of abdomen. Posterior spiracular furrows often divided. Tubular ducts each with a cup-shaped inner end and a narrow terminal filament. Anal plate posterior to anal opening well developed, shield- or butterflyshaped; sometimes with an arched plate anteriorly in addition................................................... 17

– Cribriform plates absent. Posterior spiracular furrows not divided. Tubular ducts each without either a strongly cup-shaped inner end or a narrow terminal filament. Dorso-anterior margin of anal opening with arched anal plate poorly developed or absent........................................................................... ... ASTEROLECANIIDAE ( Fig. 15 )

17 (16) Antennae each 1- or 2-segmented, with a group of quinquelocular pores present near base. Single anal plate triangular to shieldshaped. Venter of abdomen with transverse rows of pores, usually of both 8-shaped and multilocular types .................................................................................................. CEROCOCCIDAE ( Fig.16 )

– Antennae each with 7–9 segments, without quinquelocular pores near base. With 2 triangular anal plates present posterior to anal opening, linked by a narrow yoke to form a single butterfly shaped plate, plus a narrow, arched plate present more anteriorly. Venter of abdomen with transverse rows of multilocular pores only................... LECANODIASPIDIDAE ( Fig. 17 )

18 (15) With dorsal clusters of 3-, 4- or 5-locular pores, each cluster usually including 1 central tubular duct. Thick truncate setae numerous, each with sides almost parallel................................................ .. DACTYLOPIIDAE ( Fig. 18 )

– Without dorsal clusters of 3-, 4- or 5-locular pores each containing 1 central duct. If thick truncate setae present, their sides are distinctly convergent.................................................................................... 19

19 (18) Abdomen with posterior segments fused into a pygidium or pygidium-like area.................................... 20

– Abdomen with posterior segments distinct, not fused into a pygidium-like structure................................. 22

20 (19) Legs usually present.Antenna with 3‒5 segments. Dorsum of head with 1 pair of large circular ocellar spots. Disc pores each with more than 7 loculi, often flower- or star-shaped. Scale cover not visibly containing exuviae of previous instars............................................................................................ CONCHASPIDIDAE ( Fig. 19 )

– Legs absent or vestigial. Antenna with only 1 segment. Dorsum of head without large ocellar spots. Disc pores, when present, each with no more than 5 loculi. Scale cover, if present, obviously containing exuviae of previous instars.....................21

21 (20) Pygidium simple, its margin without lobes, plates or gland-spines. Detachable scale cover absent. Adult female enclosed in a test, possibly formed from second-instar exuviae, posterior end of test with a hinged flap (operculum) surrounded by a sclerotized rim. On palms or Pandanus only..................................................... HALIMOCOCCIDAE ( Fig. 20 )

‒ Pygidium with dorsal macroducts, and usually lobes and plates or gland-spines on margin. Detachable scale cover usually present, incorporating exuviae of previous instars. If adult female remains inside exuviae of previous stage (pupillarial), test of those on mango may have a posterior hinged operculum. Hosts various including palms and Pandanus ....... DIASPIDIDAE ( Fig. 21 )

22 (19) Derm covered with papillae, especially around margins.Anal ring simple, with 0–2 setae and no pores. Tubular ducts each divided longitudinally by a septum, appearing 8-shaped in cross section; duct with inner end almost flat. On palms only...................................................................................... PHOENICOCOCCIDAE ( Fig. 22 )

– Derm without papillae. Anal ring variable. In eriococcids currently known from Africa, tubular ducts each not divided longitudinally by a septum nor 8-shaped in cross-section (but not true world-wide); duct with inner end often cup-shaped. Hosts various but not on palms....................................................................................... 23

23 (20) Anal lobes small or absent. Macrotubular ducts present in a dense ventral submarginal zone. Almost exclusively on Fagaceae ............................................................................ KERMESIDAE (in part) ( Fig. 23 )

– Anal lobes, if present, prominent. Most species without a dense submarginal zone of macrotubular ducts. Hosts various, including Fagaceae ............................................................................................. 24

24(23) Microtubular ducts absent. Sclerotised anal lobes present and prominent. Hind legs without translucent pores. On Juniperus sp. only.................................................. Eriokermes juniperinus (De Lotto) (Kermesidae) ( Fig. 24 )

‒ Microtubular ducts present. Sclerotised anal lobes present or absent. Hind legs often with translucent pores. On a wide variety of hosts but almost never on Juniperus sp. ............................................ ERIOCOCCIDAE ( Figs 25 , 26 )

Figures

FIGURE 1. Margarodidae adult female morphology, from Foldi (2005a: 99, Fig. 8), reproduced with permission from the Société entomologique de France. A. Enlarged anterior leg for digging; B. Abdominal spiracle; note the absence of mouthparts.

FIGURE 2. Ortheziidae adult female morphology, from Gill (1993: 4 Fig. 1), © California Department of Food & Agriculture, Sacramento, California, U.S.A. Note the spine on the antenna tip; presence of abdominal spiracles; ovisac band formed of spines, and sclerotised anal ring with six setae.

FIGURE 3. Monophlebidae adult female morphology, from Williams & Watson (1990: 24, Fig. 5), © CABI, Wallingford, U.K. Note well-spaced antennae with basal segments only moderately enlarged, and each antenna with 8‒11 segments; ovisac band composed of pores; anal opening often distinctly dorsal, simple, not sclerotised; and well-developed mouthparts.

FIGURE 4. Matsucoccidae adult female morphology, from Gill (1993: 56, Fig. 22), © California Department of Food & Agriculture, Sacramento, California, U.S.A. Note coxa appears to be divided vertically; 2-segmented tarsus, proximal segment very short; claw without a denticle; ventral apex of tibia with stout, acute setae (A); and abdominal spiracles without pores in atria.

FIGURE 5. Kuwaniidae adult female morphology, modified from Gill (1993: 54, Fig. 20), © California Department of Food & Agriculture. Note ventral apex of tibia with numerous capitate setae (A); abdominal spiracle with pores in the atrium (B); coxa normal, not divided vertically (C); distal end of tibia with numerous capitate setae; tarsus one segmented; claw with one or more denticles; and antennal bases close together.

FIGURE 6. Kuwania oligostigma De Lotto adult female morphology, from De Lotto (1959a: 388, Fig. 2), reproduced with permission of the Entomological Society of Southern Africa. Note ventral apex of tibia with numerous capitate setae (A) and presence of mouthparts, otherwise structure similar to Fig. 5 apart from the absence of abdominal spiracles.

FIGURE 7. Putoidae adult female morphology, from Danzig & Gavrilov-Zimin (2014: 160, Fig. 2.1.1-8). Note dorsal ostiole (A); trochanter with 3 or 4 campaniform sensilla on each surface (B), and claw with plantar denticle (C). Note also that Putoidae often have a pair of denticles on the claw base, and have 9-segmented antennae; 18 pairs of cerarii-like marginal groups of setae; and have trilocular pores, a circulus and ostioles.

FIGURE 8. Pseudococcidae adult female morphology, from Williams (2004: 18, Fig. 2), © The Trustees of the Natural History Museum, London. Note dorsal ostiole (A); trochanter with 2 (rarely 3) campaniform sensilla on each surface (B); claw without a basal denticle, with or without plantar denticle (C); cerarii (numbered C1 - C18), and trilocular pores.

FIGURE 9. Rhizoecidae adult female morphology, from Williams (2004: 755, Fig. 338), © The Trustees of the Natural History Museum, London. A. Antenna usually geniculate, with no more than 6 segments; B. Bitubular cerore (tritubular in some species); C. Circulus. Note that each circulus is often raised on a sclerotised cone (see sclerotization encircling each circulus in the FIGURE); also the absence of cerarii, and of translucent pores from the hind legs.

FIGURE 10. Kerriidae adult female morphology, from Gill (1993: 6, Fig. 3), © California Department of Food & Agriculture, Sacramento, California, U.S.A. BP. Brachial plate. Note anterior thoracic spiracle much larger than posterior one; anal tubercle with anal opening surrounded by fimbriations, lobes and setae (in this case, setae); dorsum of thorax with a pair of brachia, each bearing a brachial plate, and presence of a large dorsal spine medially on abdomen. Body covered in a hard waxy test.

FIGURE 11. Stictococcidae adult female morphology, from Williams et al. (2010: 25, Fig. 10), © Magnolia Press, www. mapress.com/j/zt, reproduced with permission. Note anal complex situated at centre of dorsum, with anal opening between anterior and posterior anal plates.

FIGURE 12. Coccidae adult female morphology, from Gill (1993: 7, Fig. 4), © California Department of Food & Agriculture, Sacramento, California, U.S.A. Note well-developed anal cleft, paired triangular anal plates with inner margins usually contiguous, and tubular ducts with cup-shaped inner ends.

FIGURE 13. Micrococcidae adult female morphology, from Miller & Williams (1995; 219, Fig. VII). Note 3-segmented antenna (A); vulva on abdominal segment VI (B); paired anal plates with inner margins not contiguous (C), and tubular ducts with cup-shaped inner ends (D).

FIGURE 14. Aclerdidae adult female morphology, from Gill (1993: 8, Fig. 5), © California Department of Food & Agriculture, Sacramento, California, U.S.A. Note posterior margin with furrows and ridges; legs usually absent; short anal cleft usually present, with a single triangular or oval anal plate situated at anterior end; anal opening leading to an eversible anal tube, inner end with an anal ring bearing 10 or more setae. Mostly found on grasses.

FIGURE 15. Asterolecaniidae adult female morphology, modified from Gill (1993: 10, Fig. 7), © California Department of Food & Agriculture, Sacramento, California, U.S.A. Note tubular duct with inner end only slightly cupped and, inner filament not narrow; posterior spiracular furrow not divided; arched anal plate poorly developed or absent; 8-shaped pores present somewhere on body; cribriform plates absent.

FIGURE 16. Cerococcidae adult female morphology, from Gill (1993: 11 Fig. 8), © California Department of Food &Agriculture, Sacramento, California, U.S.A. Note 1- or 2-segmented antenna; derm around antennal base with a group of quinquelocular pores; venter of abdomen with transverse rows of 8-shaped and multilocular pores; and anal plate approximately triangular or shield-like.

FIGURE 17. Lecanodiaspididae adult female morphology, from Gill (1993: 12, Fig. 9), © California Department of Food & Agriculture, Sacramento, California, U.S.A. Note antenna with 7‒9 segments; derm around antennal base without associated quinquelocular pores; venter of abdomen with transverse rows of multilocular pores only (no 8-shaped pores, although they are present elsewhere); and anal plates forming a butterfly shape, plus a small arched plate usually present more anteriorly. Additional diagnostic features are labelled on the illustration.

FIGURE 18. Dactylopiidae adult female morphology, from Gill (1993: 13, Fig. 10), © California Department of Food & Agriculture, Sacramento, California, U.S.A. Note dorsal clusters of 3-, 4- or 5-locular pores, each cluster usually with a single central duct, and numerous thick truncate setae, each with sides almost parallel.

FIGURE 19. Conchaspididae adult female morphology, from Mamet (1954: 212 Fig. 8), reproduced with permission from The Royal Entomological Society, St Albans, U.K. Note legs usually present; antenna with 3–5 segments; a pair of dorsal ocellar spots on head; sclerotised pseudopygidium without dorsal ducts; and disc pores each with more than 7 loculi, loculi often forming a flower- or star-like pattern. Scale cover not containing exuviae of previous instars.

FIGURE 20. Halimococcidae adult and second-instar female morphology, from the illustration by G.F. Ferris in Zimmerman (1948: 431, Fig. 236). In the pupillarial adult, abdomen with posterior abdominal segments fused into a pygidium or pygidiumlike structure without dorsal ducts, and its margin without lobes, plates or gland-spines; and second-instar nymph with a hinged flap (operculum) surrounded by a sclerotised rim, this also present in test covering adult female. Legs absent; antenna only 1 segmented; dorsum of head without large ocellar spots. Detachable scale cover absent. On palms or Pandanus only.

FIGURE 21. Diaspididae adult female morphology, from Williams & Watson (1988: 12, Fig. 3), © CABI, Wallingford, U.K. Note abdomen with posterior abdominal segments fused into a sclerotised pygidium with dorsal ducts; pygidial margin with lobes, plates and / or gland spines; 1-segmented antenna; legs absent or vestigial; and disc pores each with no more than 5 loculi. The insects usually form a detachable scale cover incorporating exuviae of previous instars unless the adult female is pupillarial.

FIGURE 22. Phoenicococcidae adult female morphology, from Gill (1993: 16, Fig. 13), © California Department of Food & Agriculture, Sacramento, California, U.S.A. Note derm covered in dermal papillae especially around margins; tubular ducts each divided lengthwise by a septum, so appearing 8-shaped in cross section; anal ring simple, with 0‒2 setae and no pores; and posterior abdominal segments distinct, not fused into a pygidium-like structure. On palms only.

FIGURE 23. Kermesidae in part, adult female morphology, from Gill (1993: 9, Fig. 6), © California Department of Food & Agriculture, Sacramento, California, U.S.A. Note dense submarginal zone of macrotubular ducts with inner ends strongly cupped; anal lobes absent or poorly developed; legs absent or vestigial; antenna with only 1 segment; dorsum of head without large ocellar spots; disc pores, if present, with no more than 5 loculi.Almost always on Fagaceae.

FIGURE 24. Eriokermes juniperinus (De Lotto) (Kermesidae), adult female morphology, from De Lotto (1954a: 217, Fig. 3), reproduced with permission of the Entomological Society of Southern Africa. Note prominent, sclerotised anal lobes, and the absence of microtubular ducts and translucent pores on hind legs. On Juniperus sp. only.

FIGURE 25. Eriococcidae, eriococcine form; adult female morphology, from Gill (1993: 20, Fig. 17), © California Department of Food & Agriculture, Sacramento, California, U.S.A. Note submarginal zone without dense band of macrotubular ducts; microtubular ducts present; hind legs often with translucent pores; anal lobes prominent and sclerotised; derm without papillae. Hosts various but not on palms.

FIGURE 26. Eriococcidae, ovaticoccine form; adult female morphology, from Gill (1993: 19, Fig. 16), © California Department of Food & Agriculture, Sacramento, California, U.S.A. Note submarginal zone without dense band of macrotubular ducts; microtubular ducts present; hind legs often with translucent pores; and anal lobes not well developed or sclerotised.