Revista Facultad Nacional de Agronomía Medellín

Field characteristics

The female scale is circular, smooth, flat, light brown in the center, surrounded by a white halo (Figure 2A). The insect is reniform, membranous and yellow when it is young (Figure 2B) and it turns orange and sclerotized at maturity (Figure 2C). The female has two nymphal instars, all of them slightly darker than the adult female (Figure 2D).

Figure 2: Live specimens of Aonidiella comperei. A. Two stages of development, N1: first-instar nymph, N2: second-instar nymph B. Adult female cover. C. Young adult female separated from its scale; D. Mature adult female after removal of scale.

Slide-mounted characters

The adult female is reniform, highly sclerotized at maturity (Figure 3A). The following features are given by McKenzie (1937): pygidium not heavily sclerotized; three pairs of lobes present, the median pair only slightly larger than the second pair (Figure 3B); paraphyses small, short, and slender; tubular ducts broad and conspicuous (Figure 3C); anal opening large; venter with a few small ducts, situated close to the margin of the pygidium; perivulvar pores present, in two groups only, of apparently no more than two pores in each group (Figure 3D). The differences between A. comperei and A. orientalis are listed in Table 1.

Figure 3: Slide-mounted features of Aonidiella comperei: A. Reniform and highly sclerotized mature adult female. B. Three pairs of lobes present on the pygidium, the median pair (a) only slightly larger than the second pair (b). C. Tubular ducts. D. Perivulvar pores in two groups, each with two pores. E. Prosomatic lobes well developed. F. Fringed plates lateral to third lobe.

Table 1: Morphological differences between Aonidiella orientalis and A. comperei, showed as attributes. Character states taken from Beardsley (1966), Mckenzie (1937) and Williams and Watson (1988).

Material studied

Aonidiella comperei McKenzie. Colombia, Tolima, Chicoral, vereda Las Mercedes, finca El Diamante, 318 m.a.s.l., 04°13′41.34″ N, 75°00′11.1636″ W, 18th november 2015, coll. A. Ramos ex. leaves, branches and fruits of key lime, Citrus x latifolia (Rutaceae), 32♀♀, Catalogue No. UNAB 1837; Tolima, Espinal, vereda La Morena, Citriexpinal, 331 m.a.s.l., 04°13ʹ59.62ʺ N, 74°54ʹ23.96ʺ W, 2nd march 2016, coll. A. Ramos, ex. stored fruits of key lime, Citrus x latifolia (Rutaceae). 10♀♀, Catalogue No. UNAB 1837.

NOTE: The scale insects were found in high populations (Figure 1).

Parlatoria ziziphi (Lucas)

Coccus ziziphi Lucas, 1853: xxix. Type data: FRANCE: on Ziziphus pinnachristi. Syntypes, female. Described: female.

Chermes aurantii Boisduval, 1867: 338-339. [Synonymized by McKenzie, 1945: 54].

Parlatoria lucassi Targioni Tozzetti, 1868: 735. Nomen nudum.

Parlatoria (Websteriella) ziziphus (Lucas); Ramakrishna Ayyar, 1919: 26. Change of combination.

Apteronidia ziziphi (Lucas); Lindinger, 1934: 62. Change of combination.

Diaspis ziziphus (Lucas); Lindinger, 1934: 62. Change of combination.

Parlatoreopsis ziziphi (Lucas); Kawai, 1972: 23. Change of combination.

Field characters

According to Miller and Davidson (2005) the adult female cover is flat, broadly elongate oval, black with narrow white fringe, with two or three longitudinal ridges, shed skins marginal, black, primary component of cover second shed skin; white; body of newly matured adult female yellow brown, margin of body lateral of mouthparts with a small lobe; normally on leaves, also on branches and fruit.

Slide-mounted characters

The description and character states that differentiates P. ziziphi from others species of Parlatoria agrees well with those proposed by Miller and Davidson (2005): body oval, length usually less than two times the greatest width; presence of perivulvar pores; marginal macroducts barrel shaped, length of ducts usually less than three times width of inner end of duct; a barrel shaped macroduct present between median lobes; plates or gland spines in space between median and second lobes with at least two apical fimbriations, usually more; with conspicuous ear-like lobes on body margin laterad of mouthparts.

The differences between P. ziziphi and others species of Parlatoria present on Citrus spp. in Colombia are given in the key.

Material studied

Parlatoria ziziphi (Lucas). Colombia, Atlántico, Sabanagrande, Vda Los Caracoles, 9 m of altitude, 10°47’16.3” N, 74°46’07.2” W, 8th december 2015, coll. O. Dix; E. Palacino ex. Citrus x limonia (Rutaceae), 23♀♀, Catal. UNAB 1885.

Updated list of species of Diaspididae on Citrus spp. from Colombia. Author(s) after the colon (:) correspond to whom recorded each species.

Acutaspis scutiformis (Cockerell, 1893): Figueroa (1977).

Aonidiella comperei (McKenzie, 1937): Present study.

Aspidiotus nerii (Costa, 1829): Gallego and Vélez (1992), Kondo et al. (2012), Posada (1989).

Aulacaspis tubercularis (Newstead, 1906): Kondo et al. (2012), Posada (1989).

Chrysomphalus aonidum (Linnaeus, 1758): Balachowsky (1959), Figueroa (1946, 1952, 1977), Gallego and Vélez (1992), Kondo et al. (2012), Mosquera (1979), Posada (1989).

Chrysomphalus dictyospermi (Morgan, 1889): Figueroa (1946, 1952, 1977), Gallego and Vélez (1992), Kondo et al. (2012), Mosquera (1979), Posada (1989).

Hemiberlesia lataniae (Signoret, 1869): Kondo et al. (2012), Posada (1989).

Hemiberlesia palmae (Cockerell, 1892): Kondo et al. (2012), Posada (1989).

Ischnaspis longirostris (Signoret, 1882): Balachowsky (1959), Gallego and Vélez (1992) Kondo et al. (2012), Mosquera (1979), Posada (1989).

Lepidosaphes beckii (Boisduval, 1868): Balachowsky (1959), Figueroa (1952, 1977), Gallego and Vélez (1992), Mosquera (1979), Kondo (2001), Kondo et al. (2012).

Lepidosaphes gloverii (Packard, 1869): Balachowsky (1959), Figueroa (1952, 1977), Kondo (2001), Kondo et al. (2012), Mosquera (1979), Posada (1989).

Lopholeucaspis sp.: Mosquera (1979), Posada (1989).

Parlatoria cinerea (Hadden in Doane and Hadden, 1909): Kondo (2001), Kondo et al. (2012), Mosquera (1979), Posada (1989).

Parlatoria pergandii (Comstock, 1881): Kondo (2001), Kondo et al. (2012), Mosquera (1979), Posada (1989).

Parlatoria ziziphi (Lucas, 1853): Blackburn and Miller (1984)

Pinnaspis aspidistrae (Signoret, 1869): Figueroa (1952, 1977), Kondo et al. (2012), Mosquera (1979), Posada (1989).

Pinnaspis strachani (Cooley, 1898): Balachowsky (1959), Kondo et al. (2012), Mosquera (1979), Posada (1989).

Pseudaonidia trilobitiformis (Green, 1896): Kondo et al. (2012), Mosquera (1979), Posada (1989).

Selenaspidus articulatus (Morgan, 1889): Balachowsky (1959), Kondo et al. (2012), Mosquera (1979), Posada (1989).

Unaspis citri (Comstock, 1881): Balachowsky (1959), Figueroa (1952, 1977), Gallego and Vélez (1992), Kondo (2001), Kondo et al. (2012), Mosquera (1979), Posada (1989).

Key to subfamilies, genera and species of Diaspididae on Citrus spp. from Colombia (Compiled from Beardsley (1966), Claps and Wolff (2003), Kosztarab (1996), McKenzie (1946), Miller (2005), Miller and Davidson (2005) and Watson (2016)).

1. Macroducts normally of the “one-barred type”, length of each one at least six times the width; second pygidial lobes unilobulate; anterior spiracles usually without associated pores; antennae of adult female usually bearing only one seta; pygidial plates present in second instar if not in adult; gland spines absent; duct tubercles absent; body circular or pyriform (Figure. 4A) … Aspidiotinae …12

1’. Macroducts of the “two barred” type, length of each one rarely longer than three times their width; second pygidial lobes uni-or bilobulate; anterior spiracles usually with associated pores; antennae of adult female often bearing more than one seta; pygidial plates or gland spines often present in second instar if not in the adult; gland spines present or absent; duct tubercles may be present; body usually elongate spindle-shaped, rarely pyriform (Figure 4B) … Diaspidinae …2

Figure 4: Habitus with magnified structures of two main subfamilies of Diaspididae. A. Aspidiotinae (Illustration from Miller and Davidson 1998); B. Diaspidinae. Illustration by Davidson (Miller, 2005).

2. Adult female with second lobes bilobulate (Figure 5A); gland spines usually present, occasionally replaced by plates; pores often associated with anterior and posterior spiracles; marginal macroducts usually with the long axis of each orifice perpendicular to margin; antennae each usually with two or more setae … 6

2’.Adult female with second lobes unilobulate (Figure 5B); gland spines absent, plates usually present; spiracular pores associated with anterior spiracles only; marginal macroducts often with the long axis of each orifice oriented parallel to margin; antenna with one or more setae … 3

Figure 5: A. Pigydium with second lobe bilobulate; B. Pigydium with second lobe unilobulate. Illustrations taken from Miller and Davidson (2005).

3. Antenna with two to six setae; adult female pupillarial; body more or less elongate; plates, if present, confined to pygidium; marginal pygidial ducts not enlarged; abdominal disc pores sometimes present on some prepygidial segments … Lopholeucaspis sp .

3’ Antenna with one seta; adult female usually not pupillarial; body oval to circular (elongate if pupillarial); plates often present on pre pygidium and pygidium; marginal pygidial ducts often enlarged; abdominal disc pores never present on some prepygidial segments … Parlatoria (Targioni Tozzetti) … 4

4. With a large, conspicuous, lateral and membranous lobe on each side of head, marginal to each anterior spiracle; female scale composed principally of a large black second exuvium … Parlatoria ziziphi (Lucas) (Figure 6).

4’ Without such a membranous lobe on each side of head; female scale cover not largely black … 5

Figure 6: Habitus of Parlatoria ziziphi (Lucas) with magnified structures. Illustration taken from Miller and Davidson (2005).

5. Dorsum of pygidium with two longitudinal rows of macroducts on each side of pygidium on abdominal segments six and seven, extending cephalad to well anterior of anal opening; three plates between third and fourth lobs … Parlatoria cinerea (Hadden in Doane and Hadden) (Figure 7A)

5’. Dorsum of pygidium without such rows of macroducts; macroducts not found anterior to anal opening on these segments; four plates between third and fourth lobes … Parlatoria pergandii (Comstock) (Figure 7B)

Figure 7: Habitus with magnified structures of: A. Parlatoria cinerea (Hadden in Doane & Hadden) (Illustration taken from Gerson 1977); B. Parlatoria pergandii (Comstock). Illustration taken from Miller and Davidson (2005).

6. Median lobes zygotic (Figure 8A) … 7.

6’. Median lobes not zygotic (although crowding or secondary sclerotization may make this difficult to see) (Figure 8B) … 9

Figure 8: A. Median lobes zygotic; B. Median lobes not zygotic. Illustrations taken from Watson (2016).

7. Gland spines and macroducts absent from area anterior to abdominal segment 1; body of characteristic shape, with head, prothorax, and mesothorax swollen and rectangular; lateral margin side of prothorax on older females with 1 swollen tubercle on each side of body; labium set in groove, with sclerotized areas on each side … Aulacaspis tubercularis (Newstead) (Figure 9).

7’.Gland spines and macroducts present anterior to abdominal segment 1; head, prothorax, and mesothorax not swollen; without lateral tubercles on lateral margin side of prothorax on older females; labium set not in groove, without sclerotized areas on each side … Pinnaspis (Cockerell) … 8

Figure 9: Habitus with magnified structures of Aulacaspis tubercularis (Newstead). Illustration taken from Miller and Davidson (2005).

8. Preanal sclerosis lacking or represented by light sclerotized patch; median lobes protrude less than or about same distance as second lobes; posterior spiracles each with one to 12 pores, four pores on average … Pinnaspis aspidistrae (Signoret) (Figure 10A)

8’. Preanal sclerosis represented by sclerotized bar; median lobes protrude beyond or about same distance as second lobes; posterior spiracles each with 0 to four pores, two pores on average … Pinnaspis strachani (Cooley) (Figure 10B)

Figure 10: Habitus with magnified structures of: A. Pinnaspis aspidistrae (Signoret) B. Pinnaspis strachani (Cooley). Illustrations taken from Miller and Davidson (2005).

9. Median lobes with only a pair of marginal setae between their bases … 11

9’. Median lobes with only a pair of gland spines between their bases … Lepidosaphes (Schlectendal) … 10

10. Mature female without sclerotized pattern on thorax; without sclerotized dermal pockets on thorax; macroducts on dorsal margin of prothorax present … Lepidosaphes beckii (Boisduval) (Figure 11A)

10’. Mature female with distinctive pattern of punctures on dorsum of thorax extending to ventral margin; with sclerotized dermal pockets on pro- and mesothorax; macroducts on dorsal margin of prothorax absent … Lepidosaphes gloverii (Packard) (Figure 11B)

Figure 11: Habitus with magnified structures of: A. Lepidosaphes beck ii (Boisduval); B. Lepidosaphes gloverii (Packard). Illustrations taken from Miller and Davidson (2005).

11. Dorsum of pygidium with a coarse ‘lattice-work’ sclerotized pattern … Ischnaspis longirostris (Signoret) (Figure 12A)

11’. Dorsum of pygidium without such a sclerotized pattern; with four or fewer perivulvar pores on each side of body … Unaspis citri (Comstock) (Figure 12B)

Figure 12: Habitus with magnified structures of: A. Ischnaspis longirostris (Signoret); B. Unaspis citri (Comstock). Illustrations taken from Miller and Davidson (2005).

12. Dorsal surface of pygidium with a conspicuous mosaic or areolate appearance … Pseudaonidia trilobitiformis (Green) (Figure 13A)

12’. Dorsal surface of pygidium without mosaic areolate pattern … 13

Figure 13: A. Habitus with magnified structures of Pseudaonidia trilobitiformis (Green). Illustration from Miller and Davidson (2005); B. Pygidial apex acute, forming an angle of 90° or less of Acutaspis scutiformis (Cockerell). Illustration taken from Watson (2016).

13. Pygidial margin with paraphyses present, these sometimes small (arising from bases of lobes or from the margin itself); paraphyses sometimes present lateral to the outer lobes … 14

13’. Pygidial margin without paraphyses … 19

14. Pygidial apex acute, forming an angle of 90° or less, although basal half of pygidium may be broad (Figure 13B); pygidial margins tending to be concave and sclerotized; plates confined to interlobular spaces … Acutaspis scutiformis (Cockerell)

14’. Pygidial apex more or less broad, forming an angle greater than 90°; pygidial margins convex or straight, with or without sclerotization; plates often present lateral to third lobes … 15

15. Mature specimens with prosoma very enlarged and sclerotized, usually reniform so its lateral lobes more or less enclose the pygidium. If prosoma not reniform at maturity (A. orientalis), then paraphyses between second and third lobes never longer than the lobes … Aonidiella comperei McKenzie (Figure 14)

15’. Mature specimens with prosoma membranous or sclerotized, not expanded sufficiently to become reniform; paraphyses between second and third lobes shorter or longer than the lobes … 16

Figure 14: Habitus with magnified structures of Aonidiella compereiMcKenzie. Illustration taken from McKenzie (1937).

16. Paraphyses obviously longer than median lobes (Figure 15A) … Chrysomphalus (Ashmead) … 17

16’ Paraphyses same length or shorter than median lobes (Figure 15B) … Hemiberlesia (Berlese and Leonardi) … 18

Figure 15: A. Paraphyses obviously longer than median lobes; B. Paraphyses same length or shorter than median lobes. Illustrations taken from Miller and Davidson (2005).

17. With one cluster of macroducts on submarginal areas of prepygidial segments; plates just lateral to third lobes fringed; nine to 13 prevulvar pores on each side of pygidium … Chrysomphalus aonidum (Linnaeus) (Figure 16A)

17’. Without clusters of macroducts on submarginal areas of prepygidial segments; plates just lateral to third lobes clavate; five or six prevulvar pores on each side of pygidium … Chrysomphalus dictyospermi (Morgan) (Figure 16B).

Figure 16: Habitus with magnified structures of: A. Chrysomphalus aonidum (Linnaeus); B. Chrysomphalus dictyospermi (Morgan). Illustrations taken from Miller and Davidson (2005).

18 Medial notch, first and second space of pygidium with band of fringe plates less strongly developed; plates in third space, if present, reduced in size, not forming such a dense uniform fringe; median lobes very close together, almost in touch; … Hemiberlesia lataniae (Signoret) (Figure 17A)

18’. Medial notch, first, second and third space of pygidium with a band of broad apically fringed plates of nearly uniform; plates in third space always present in a dense, nearly uniform fringe; median lobes separated, at least in the half of the width of each median lobe … Hemiberlesia palmae (Cockerell) (Figure 17B)

Figure 17: Habitus with magnified structures of A. Hemiberlesia lataniae (Signoret). Illustrations taken from Miller and Davidson (2005); B. Hemiberlesia palmae (Cockerell). Illustrations taken from Ferris (1938).

19. Prosoma strongly constricted between meso- and metathorax, and sclerotized; third lobe represented by an elongate, acute, sclerotized spine … Selenaspidus articulatus (Morgan) (Figure 18a)

19’. Prosoma without strong constriction between meso- and metathorax, and usually membranous; third lobe short, either rounded or pointed, or absent … Aspidiotus neri i (Costa) (Figure 18b)

Figure 18: Habitus with magnified structures of: A. Selenaspidus articulatus (Morgan); B. Aspidiotus nerii (Costa). Illustrations taken from Miller and Davidson (2005).