Ovaticoccus Kloet 1944

Miller, Douglass R. & Stocks, Ian C., 2022, New genera and species of felt scales (Hemiptera: Coccomorpha: Eriococcidae), with descriptions of new species and immature instars of described species, Zootaxa 5221 (1), pp. 1-213 : 85-90

Ovaticoccus Kloet 1944

Gymnococcus Cockerell 1894b: 1053 . Type species: Coccus agavium Douglas 1888 by monotypy (junior homonym discovered by Kloet 1944: 86). Notes: junior homonym of a genus of Mycetozoa ( Gymnococcus Zopf 1887 )

Ovaticoccus Kloet 1944: 86 (replacement name for the junior homonym Gymnococcus Cockerell 1894b ). Type species: Coccus agavium Douglas 1888 by monotypy.

Cornoculus Ferris 1955: 81 syn. n. Type species: Cornoculus cornutus Ferris 1955 , by monotypy.

Onceropyga Ferris 1955: 208 syn. n. Type species: Eriococcus neglectus Cockerell 1895 , by original designation (junior homonym discovered by Hoy 1963: 179). Notes: junior homonym of a genus of Lepidoptera : Zygaenidae , Onceropyga Turner 1906 .

Oregmopyga Hoy 1963: 179 syn. n. Type species: Eriococcus neglectus Cockerell 1895 , by original designation (replacement name for the junior homonym Onceropyga Ferris 1955 )

Generic diagnosis of adult female: Body oval or rotund. Anal lobes either not protruding, or protruding slightly. Flagellate setae on dorsum and venter, longest setae usually medially on ventral abdomen. Often with enlarged setae of species-specific shape. Usually with microtubular ducts and macrotubular ducts (only in adult females and second-instar males). Multilocular pores present with from 3–12 or 14 loculi. Cruciform pores present or absent. Anal ring variable, configuration usually diagnostic, normally with 2 lateral plates, sometimes with plates joined by sclerotization anteriorly and/or posteriorly forming complete or partial ring; with or without pores, usually with 3 setae on each side of ring, anal tube and/or anal opening sometimes sclerotized, sometimes with anal flap covering anal opening. Legs well-developed; hind coxae each with translucent pores in adult and second-instar females; each femur with 3–5 setae, without translucent pores; each tibia with 3–5 setae, rarely with 1 in middle of tibia; each tarsus with campaniform sensillum near junction of tibia and with 2 capitate digitules; claws each with denticle and 2 capitate digitules. Antennae each 6- or 7-segmented, apical 3 segments each with 1 or more fleshy setae. Without frontal lobes. With preantennal pore. Microtrichia usually present dorsally at least on posterior abdominal segments, present ventrally from head or thorax to posterior abdominal segments, often present on coxae.

Notes: Including Oregmopyga and Cornoculus as junior subjective synonyms of Ovaticoccus gives a broader definition of the genus. No single character is diagnostic for all Ovaticocccus species, but the genus can be recognized by the following combination of characters: anal lobes not protruding or slightly protruding; anal-lobe areas each without enlarged setae or rarely with one or two; anal ring often reduced and with zero to three setae on each side of ring; enlarged setae often characteristically shaped; claw digitules same length as, or longer than claw, and capitate. Ovaticoccus is similar to Acanthococcus in having: enlarged setae; microtubular ducts each with two internal scleroses; claw digitules longer than claw and capitate. Species of Acanthococcus differ by (character states of Ovaticoccus species are given in brackets): anal lobes protruding, each lobe usually with three or four enlarged setae (anal lobes not protruding or slightly protruding, each lobe with two or fewer enlarged setae); anal ring with four setae on each side of ring and with at least one ring of cells (anal ring with zero to three setae on each side of ring and often with reduced number of cells); enlarged setae often conical, not pear- or nipple-shaped (enlarged setae sometimes conical, often pear- or nipple-shaped). Species of Ovaticoccus are similar also to species of the Old-World genus Hispaniococcus Kozár in having: three setae on each side of anal ring; enlarged setae nipple-shaped in one species of Hispanococcus; and microtubular ducts with two internal sclerotized areas. Species of Hispaniococcus differ from species of Ovaticoccus by (character states in brackets are of Ovaticoccus ): claw digitules shorter than length of claw and apically acute (claw digitules same length as, or longer than, length of claw and apically capitate); anal lobes protruding; and each lobe with three or more enlarged setae (anal lobes not protruding or slightly protruding, and each lobe with two or fewer enlarged setae).

Ferris (1955) and Miller and McKenzie (1967) characterized Oregmopyga as having: slightly protruding anal lobes; and often possessing characteristically shaped enlarged setae, e.g., nipple-shaped. Species in the genus usually possess an anal ring with two rows of pores and three pairs of elongate setae, and 7-segmented antennae. The only character that consistently characterizes species of Oregmopyga is the presence of slightly protruding anal lobes, but even these vary from clearly protruding to barely perceptible. However, Oregmopyga species with protruding anal lobes present a mosaic of other character states that include an anal ring without pores, e.g., Or. neglecta (the type species); an anal ring with reduced setae, e.g., Or. neglecta ; and antennae that are 6-segmented, e.g., Or. eriogoni Miller. There are species of Ovaticoccus that possess similarly shaped enlarged setae to Oregmopyga species, e.g., Ov. exoticus Pellizzari and Kozár , and have 7-segmented antennae, e.g., Ov. agavium (Douglas) . Based on this variation, we place Oregmopyga as a junior subjective synonym of Ovaticoccus .

Ferris (1955) characterized the single species of Cornoculus ( C. cornutus ) as having: unusually large hornlike eyes; characteristically shaped setae, e.g., nipple-shaped; antennae 7-segmented; multilocular pores on dorsum and venter; and anal ring reduced. Miller and McKenzie (1967) added a second species to Cornoculus ( C. densus Miller ), based primarily on the large number of macrotubular ducts on the lateral areas of the venter but not having unusual eyes. With the exception of the unusual eyes, the species of Cornoculus possess all of the characteristics of species of Ovaticoccus and are here considered to be part of the latter genus. The unusual large eyes are believed to be an autapomorphy.

Enlarged setae are common, obvious and distinctive characters in Ovaticoccus species and often have been used as diagnostic of many species. However, we have found their presence or absence to be unreliable for species recognition except in the first-instar nymphs, and even in this instar there can be variation. Adult females and second-instar nymphs tend to show a great deal of variation; surprisingly, transformation of flagellate setae from enlarged setae appears to occur with ease in the process of molting. We have found specimens of the same instar, from the same locality, both with and without enlarged setae, e.g., second-instar males of Ov. agavium and adult females of Ov. betsyae Miller and Stocks sp. n. We also have observed specimens with the presence of enlarged setae on one side of the body, and not on the other. Because of this variation we have avoided using the presence or absence of enlarged setae as primary diagnostic characters of species whenever possible. However, the shape of enlarged setae is reliable in characterizing many Ovaticoccus species, e.g., Ov. davesmithi Miller and Stocks versus Ov. johnsoni Miller.

The arrangement of the dorsal setae (either enlarged or flagellate or both) is segmental, usually with a single row of setae across each abdominal segment and a less obvious segmental arrangement on the head and thorax. In most species there also is a longitudinal arrangement of these setae, often characterized as lateral, mediolateral and medial. This longitudinal pattern is most obvious in the first-instar nymph, but can be seen in the adult female and even some of the male instars in some species. This arrangement can be useful in distinguishing among different species. Although the descriptions often describe this pattern as occurring on the entire dorsum, it may not be as obvious on the head and thorax as it is on the abdomen.

The seta that is normally located near the anal ring and often is mentioned as “additional pair of closely associated setae not attached to anal ring” in the descriptions of the anal ring, is most likely the same as the suranal seta. It often is difficult to determine the anterior limits of the so-called anal-lobe area, which may cause inconsistencies in the data presented in the text. The suranal seta is usually included in these data for the ventral anal lobe.

Other structures useful in species recognition in Ovaticoccus that merit special mention are multilocular pores, the anal ring, and the setae on the femur. The predominant number of loculi in multilocular pores is a reliable diagnostic character. Many species have the predominant kind of multilocular pores with five loculi, e.g., Ov. sanguineus Miller , but several species have pores predominantly with seven loculi, e.g., Ov. cornutus , or even nine loculi, e.g., Ov. tippinsi . The same loculus-number predominance often occurs throughout the life history of the species in first-instar nymphs, second-instar males and females, and adult females.

The structure of the anal ring also is surprisingly consistent throughout the life history of a species. For example, the anal ring of Ov. neglectus is horseshoe shaped, lacks pores, the setae are minute or absent, the anal tube and anal opening are weakly sclerotized, and there is no anal flap. This is true for all instars except the male prepupa, pupa, and adult. In contrast to Ov. neglectus , the consistent structure of the anal ring in the first-instar nymph, second-instar male and female, and adult female of Ov. agavium is composed of two lateral plates, each with three setae on each side of the ring that are each about as long as the greatest diameter of the ring, which lacks pores; furthermore, these instars of Ov. agavium have the anal tube unsclerotized, the anal opening is sclerotized, and lacks an anal flap.

The number and distribution of the setae on the femur follow the same pattern, i.e., they remain the same throughout the life history of each species. There are several diagnostic patterns including: a total of four setae (one proximal seta, three distal setae), e.g., Ov. haigi Miller and Stocks sp. n.; five setae (two proximal setae, three distal setae), e.g., Ov. sanguineus ; and three setae (zero proximal setae, three distal setae), e.g., Ov. neglectus .

Unlike most species of Acanthococcus , species of Ovaticoccus rarely have a middle seta on the tibia. Miller and McKenzie (1967), Gill (1993), and Kozár et al. (2013) used this as a diagnostic character state for species recognition within the Eriococcidae , but surprisingly the setae on the femur are much more diagnostic for species in Ovaticoccus .

Intraspecific variation occurs in certain species of Ovaticoccus causing the use of “in part” designations in dichotomous keys. Examples include: the number of segments in the antennae of the adult female of Ov. maryfoleybensonae and of Ov. davesmithi (6- or 7-segmented); the sclerotization of the anal ring on the adult female of Ov. mackenziei (complete or divided anteriorly); and cruciform pores on the second-instar females of Ov. salviae (present or absent).

Etymology: The genus epithet “ Ovaticoccus ” is formed from the Latin word “ ovatus” meaning “egg-shaped” and the Greek “ kokkos” meaning “round structure” or “scale insect” and apparently refers to the oval body shape of this scale insect. The generic name is a masculine noun.

Field features: Found in a diversity of microhabitats including within the leaf-blade sheaths of grasses, the tightly adhering bases of leaves of yucca and agave, under the bark of shrubs such as Salvia and Hypericum , and in cyst-like encasements in the soil. Generally, they are found in tight, protected areas of the host.

Key to Ovaticoccus and Hypericicoccus species based on adult females

1(0) Dorsal cruciform pores present.......................................................................... 2

– Dorsal cruciform pores absent.......................................................................... 21

2(1) Pores present in anal ring.............................................................................. 3

– Pores absent from anal ring............................................................................ 11

3(2) Dorsum without dorsal dermal projections; with macrotubular ducts; legs well formed.............................. 5

– Dorsum with numerous dorsal dermal projections; without macrotubular ducts; legs poorly formed.................... 4

4(3) Hind coxae without pores; without enlarged setae or with fewer than 15................ gordoni Miller and Stocks sp. n.

– Hind coxae with pores; with more than 20 enlarged setae........................................ hyperici (Ferris)

5(3) Antennae 7-segmented................................................................................ 6

– Antennae 6-segmented............................................................................... 17

6(5) Multilocular pores predominantly with 5 loculi............................................................ 10

– Multilocular pores predominantly with more than 5 loculi, or with equal numbers of 5-locular pores and 7-locular pores... 7

7(6) Eyes not enlarged; without large clusters of macrotubular ducts in ventrolateral areas of abdominal segments............ 8

– Eyes enlarged; with large clusters of macrotubular ducts in ventrolateral areas of abdominal segments..... cornutus (Ferris)

8(7) Enlarged setae present near body margin; anal ring complete.................................................. 9

– Enlarged setae absent; anal ring divided posteriorly................ maryfoleybensonae Miller and Stocks sp. n. (in part)

9(8) Without pores on hind coxae; with conical setae on margin of abdominal segment VIII................................................................................................ telotrichus Miller and Stocks sp. n. (in part)

– With pores on hind coxae; with dome-shaped setae on margin of abdominal segment VIII................................................................................................. davesmithi (Miller & Stocks) (in part)

10(6) Dorsal enlarged setae uncommon or absent, if present, then most abundant dorsally on margin; cruciform pores absent or uncommon in ventromedial areas between legs (<10), often restricted to marginal areas............................ 39

– Dorsal enlarged setae common, present over entire dorsal surface; cruciform pores abundant over entire ventral surface including between legs (>10).................................................................. parvispinus (Chaffin)

11(2) Without cluster of dorsal cruciform pores on segment VIII................................................... 13

– With cluster of dorsal cruciform pores on segment VIII...................................................... 12

12(11) Without distinct pores on hind coxae; on grass.................................................. adoxus (Ferris)

– With distinct pores on hind coxae; on cactus................................ villanuevorum Miller and Stocks sp. n.

13(11) Macrotubular ducts present on dorsum; on several hosts..................................................... 14

– Macrotubular ducts absent from dorsum; on Hymenoclea ................................. viscosa (Kondo) (in part)

14(13) Multilocular pores predominantly with 5 loculi............................................................ 15

– Multilocular pores predominantly with more than 5 loculi................................................................................................................................. tuttlei Miller and Stocks sp. n.

15(14) Anal ring with setae; enlarged setae conical or pear-shaped, with base narrower than length of seta................... 16

– Anal ring without setae; enlarged setae dome-shaped, basal width same as or wider than length of seta..... densus (Miller)

16(15) Antennae 7-segmented; without small pores on derm anterior to hind coxae; on Artemisia and Eriogonum . variabilis Miller

– Antennae 6-segmented; with small pores on derm anterior to hind coxae; on grass............... haigi Miller and Stocks

17(5) Anal ring incomplete, divided either dorsally or ventrally or both.............................................. 18

– Anal ring complete, not divided........................................................................ 20

18(17) Translucent pores present on hind coxae; enlarged setae conical or absent....................................... 19

– Translucent pores absent from hind coxae; enlarged setae dome-shaped...................... tippinsi (Miller & Miller)

19(18) Enlarged setae absent........................................ maryfoleybensonae Miller and Stocks sp. n. (in part)

– Enlarged setae present............................................................ strongyla (Miller & Miller)

20(17) Without pores on hind coxae; with conical setae on margin of abdominal segment VIII................................................................................................ telotrichus Miller and Stocks sp. n. (in part)

– With pores on hind coxae; with dome-shaped setae on margin of abdominal segment VIII................................................................................................. davesmithi (Miller & Stocks) (in part)

21(1) Anal ring complete, not divided........................................................................ 22

– Anal ring divided anteriorly or posteriorly, or both......................................................... 29

22(21) With dorsal multilocular pores......................................................................... 23

– Without dorsal multilocular pores....................................................................... 27

23(22) Anal ring pores absent................................................................................ 24

– Anal ring pores present............................................................................... 26

24(23) Antennae each 6-segmented........................................................................... 25

– Antennae each 7-segmented........................................................ viscosa (Kondo) (in part)

25(24) Small pores present on derm anterior to hind coxa................................................ salviae Miller

– Small pores absent from derm anterior to hind coxa..................................... mackenziei Miller (in part)

26(23) Enlarged setae longer than wide; antennae each 7-segmented........................... peninsularis (Ferris) (in part)

– Enlarged setae about as wide as long; antennae each 6-segmented................................. eriogoni (Miller)

27(22) Without small pores anterior to hind coxae................................................................ 28

– With small pores anterior to hind coxae.................................................... senarius McKenzie

28(27) Anal ring without setae or with 1 inconspicuous seta on each side of ring........................... parkerorum Miller

– Anal ring with 3 setae on each side of ring............................................. variabilis Miller (in part)

29(21) Protruding anal lobes absent........................................................................... 32

– Protruding anal lobes present, sometimes small............................................................ 30

30(29) Pores absent from derm anterior to hind coxa.............................................................. 31

– Pores present on derm anterior to hind coxa............................................... neglectus (Cockerell)

31(30) Anal ring with pores; with enlarged setae; alcohol with preserved specimens clear................................ 38

– Anal ring without pores; without enlarged setae; alcohol-preserved specimens turn liquid crimson...... sanguineus (Miller)

32(29) Cruciform pores absent from ventromedial areas of abdomen................................................. 36

– Cruciform pores present in ventromedial areas of abdomen................................................... 33

33(32) Macrotubular ducts present at least on posterior abdominal segments........................................... 34

– Macrotubular ducts absent....................................................... exoticus Pellizzari and Kozár

34(33) Macrotubular ducts present on head and/or thorax.......................................................... 35

– Macrotubular ducts restricted to abdomen.................................................. agavium (Douglas)

35(34) Dorsal multilocular pores scattered over entire surface, with more than 10............. agavacearum Pellizzari and Kozár

– Dorsal multilocular pores restricted to head and thorax, with fewer than 10................... variabilis Miller (in part)

36(32) Microtubular ducts absent............................................................................. 37

– Microtubular ducts present........................................................ mackenziei Miller (in part)

37(36) Multilocular pores with about equal numbers of 5-locular and 7-locular pores; cruciform pores common lateroventrally, forming clusters of 9–18 pores on each of abdominal segments II or III to VII; occurring on Agave sp. ...... californicus McKenzie

– Multilocular pores primarily 5-locular, with 3-locular pores uncommon; cruciform pores in small numbers lateroventrally, forming clusters of 1 or 2 pores on each of abdominal segments II, III, or IV to VIII; occurring on Yucca sp. ........................................................................................ betsyae Miller and Stocks sp. n.

38(31) Hind coxae with translucent pores restricted to ventral surface.......................... peninsularis (Ferris) (in part)

– Hind coxae with translucent pores present on both surfaces....................................... johnsoni (Miller)

39(10) Hind coxae greatly enlarged, with numerous translucent pores on both surfaces.......... amplicoxae Williams and Martin

– Hind coxae not enlarged, with few translucent pores on ventral surface.......... peruvianus (Granara de Willink & Díaz)

Key to Ovaticoccus and Hypericicoccus species based on second-instar females

1(0) Dorsal cruciform pores present.......................................................................... 2

– Dorsal cruciform pores absent.......................................................................... 11

2(1) With pores in anal ring................................................................................ 3

– Without pores in anal ring.............................................................................. 8

3(2) Most abundant multilocular pores with more than 5 loculi..................................................... 4

– Most abundant multilocular pores with 5 loculi............................................................. 6

4(3) Anal ring setae thin, flagellate, apices acute................................................................ 5

– Anal ring setae thick, apices blunt........................................................... hyperici (Ferris)

5(4) Dorsal multilocular pores present.................................................... tippinsi (Miller & Miller)

– Dorsal multilocular pores absent except on head............................................... cornutus (Ferris)

6(3) Enlarged setae approximately as long as wide, dome-shaped................................................... 7

– Enlarged setae longer than wide, conical or pear-shaped............................. peruvianus (Granara de Willink)

7(6) Hind tibiae each with seta medially (total of 5 setae)...................................... densus (Miller) (in part)

– Hind tibiae each without seta medially (total of 4 setae)............................... davesmithi (Miller & Stocks)

8(2) Enlarged setae conical or pear-shaped, length greater than width................................................ 9

– Enlarged setae dome-shaped, length shorter or same as width............................... densus (Miller) (in part)

9(8) Dorsal cruciform pores not restricted to area anterior to anal ring.............................................. 10

– Dorsal cruciform pores restricted to medial area anterior to anal ring on segment VIII sometimes with 1 or 2 medial pores on segment VII............................................................................... adoxus Ferris

10(9) Cruciform pores present ventrally; not occurring on Salvia ................................................... 19

– Cruciform pores absent ventrally; occurring under bark of Salvia ............................. salviae Miller (in part)

11(1) Ventral cruciform pores absent......................................................................... 12

– Ventral cruciform pores present......................................................................... 16

12(11) Dorsal multilocular pores present....................................................................... 13

– Dorsal multilocular pores absent........................................................................ 21

13(12) Anal ring setae absent or shorter than half of diameter of ring................................................. 14

– Longest anal ring seta longer than half of diameter of ring................................................... 15

14(13) Each femur with 3 setae, without proximal seta; enlarged setae present ventrolaterally...... neglectus (Cockerell) (in part)

– Each femur with 4 setae, with proximal seta; enlarged setae absent ventrolaterally........ betsyae Miller and Stocks sp. n.

15(13) Cruciform pores usually absent ventromedially........................................... salviae Miller (in part)

– Cruciform pores usually present ventromedially...................................... agavium (Douglas) (in part)

16(11) Dorsal multilocular pores present....................................................................... 18

– Dorsal multilocular pores absent........................................................................ 17

17(16) Enlarged setae present on venter; usually without proximal seta on each femur....................... eriogoni (Miller)

– Enlarged setae absent from venter; usually with proximal seta on each femur........................ viscosa (Kondo)

18(16) Each femur with 3 or 4 setae, 1 proximal seta present or absent............................................... 20

– Each femur with 5 setae, 2 proximal setae present..................................... sanguineus (Miller) (in part)

19(10) Each femur with 4 setae, 3 distally, 1 proximally..................................... haigi Miller and Stocks sp. n.

– Each femur with 5 setae, 3 distally, 2 proximally...................................... sanguineus (Miller) (in part)

20(18) Each femur with 3 setae, without proximal seta....................................... agavium (Douglas) (in part)

– Each femur with 4 setae, with 1 proximal seta....................................... exoticus Pellizzari and Kozár

21(12) Abdominal segment IV with combined total of 8 enlarged setae on dorsum and venter; without microtubular ducts in mediolateral areas of dorsum of abdomen............................................................... variabilis Miller

– Abdominal segment IV with combined total of 9–11 enlarged setae on dorsum and venter; with microtubular ducts in mediolateral areas of dorsum of abdomen....................................................... neglectus (Miller) (in part)

Key to Ovaticoccus and Hypericicoccus species based on second-instar males

1(0) Anal ring without pores................................................................................ 2

– Anal ring with pores.................................................................................. 5

2(1) Ventral cruciform pores present.......................................................................... 3

– Ventral cruciform pores absent.......................................................................... 4

3(2) Anal ring incomplete with 2 lateral plates, divided anteriorly and posteriorly...................................... 8

– Anal ring complete or divided anteriorly..................................................... viscosa (Kondo)

4(2) Each femur with 3 setae including 3 distal setae, without proximal seta.......................................... 9

– Each femur with 4 setae including 1 proximal seta and 3 distal setae................... betsyae Miller and Stocks sp. n.

5(1) 5-locular pores most abundant kind of multilocular pore...................................................... 6

– 7-locular pores most abundant kind of multilocular pore.......................................... hyperici (Ferris)

6(5) Dorsal enlarged setae present; dorsal microtubular ducts present................................................ 7

– Dorsal enlarged setae absent; dorsal microtubular ducts absent........................ peruvianus (Granara de Willink)

7(6) Enlarged setae dome-shaped, about as wide as long; anal ring with fewer than 10 pores....... davesmithi (Miller & Stocks)

– Enlarged setae conical or dome-shaped, longer than wide; anal ring with more than 10 pores........ parvispinus (Chaffin)

8(3) Each femur with 4 setae, 1 proximally and 3 distally.................................. exoticus Pellizzari and Kozár

– Each femur with 3 setae, all 3 distally...................................................... agavium (Douglas)

9(4) Ventral abdominal segments each with more than 5 multilocular pores; 7-locular pores abundant, nearly as common as 5-locular pores; microtubular ducts common on dorsum, with more than 5.............................. neglectus (Cockerell)

– Ventral abdominal segments each with fewer than 5 multilocular pores; 7-locular pores rare or absent, most multilocular pores with 5 loculi; microtubular ducts rare on dorsum, with fewer than 5.................................. salviae Miller

Key to Ovaticoccus and Hypericicoccus species based on first-instar nymphs

1(0) Dorsomedial longitudinal line of enlarged setae present on abdomen, sometimes restricted to 1 or 2 setae on posterior segments........................................................................................... 2

– Dorsomedial longitudinal line of enlarged setae absent from abdomen........................................... 9

2(1) Pores absent from anal ring............................................................................. 3

– Pores present in anal ring.............................................................................. 6

3(2) Most multilocular pores with 5 or more loculi, or with equal numbers of 3- and 5-locular pores....................... 4

– Most multilocular pores with 3 loculi........................................................ variabilis Miller

4(3) Anal ring divided posteriorly and anteriorly............................................................... 14

– Anal ring not divided or divided anteriorly only............................................................. 5

5(4) Each femur with 3 setae; each tibia without middle seta......................................... eriogoni (Miller)

– Each femur with 5 setae; each tibia with middle seta............................................. densus (Miller)

6(2) Anal ring complete................................................................................... 7

– Anal ring incomplete.................................................................................. 8

7(6) Multilocular pore associated with anterior spiracle with 7 loculi; anal ring setae enlarged, with rounded apices................................................................................................... hyperici (Ferris)

– Multilocular pore associated with anterior spiracle with 5 loculi; anal ring setae slender, with acute apices.......................................................................................... braggi (Cockerell & Robinson)

8(6) Multilocular pores absent dorsally; without ventral enlarged setae............................................. 15

– Multilocular pores present dorsally on abdominal segment VIII; lateral longitudinal line of enlarged setae present ventrally.................................................................................... parvispinus (Chaffin)

9(1) Each femur with 3 or 4 setae........................................................................... 10

– Each femur with 5 setae.............................................................................. 12

10(9) Each femur with 4 setae.............................................................................. 11

– Each femur with 3 setae.............................................................. neglectus (Cockerell)

11(10) Multilocular pores predominantly with 5 or more loculi, 3-locular pores absent or rare; enlarged setae restricted to lateral areas..................................................................................... adoxus Ferris

– Multilocular pores predominantly with 3 loculi, pores with 5 loculi absent or rare; enlarged setae forming 2 pairs of longitudinal lines (mediolateral and lateral)................................................... haigi Miller and Stocks sp. n.

12(9) Anal lobes with conical enlarged setae; with a few cruciform pores on thorax.................................... 13

– Anal lobes without conical enlarged setae; without cruciform pores...................... davesmithi (Miller & Stocks)

13(12) Enlarged setae present in mediolateral line on dorsal abdomen..................... telotrichus Miller and Stocks sp. n.

– Enlarged setae absent mediolaterally on dorsal abdomen.................................. tippinsi (Miller & Miller)

14(4) Each femur with 4 setae, including proximal seta..... agavacearum Pellizzari and Kozár , also exoticus Pellizzari and Kozár

– Each femur with 3 setae, without proximal seta.............................................. agavium (Douglas)

15(8) Two multilocular pores associated with each spiracle; multilocular pores predominantly with more than 5 loculi........................................................................... maryfoleybensonae Miller and Stocks sp. n.

– One multilocular pore associated with each spiracle; multilocular pores predominantly with 5 loculi.............................................................................................. peruvianus (Granara de Willink)