Puto, Signoret, 1876

Powell, Erin C. & Miller, Douglass R., 2024, Three new species of Puto giant mealybugs (Hemiptera: Coccomorpha: Putoidae) from Mexico and Guatemala, with keys to New World adult females and World adult males, Zootaxa 5443 (3), pp. 301-336 : 303-304

Key to adult females of Puto from the New World (revised from McKenzie 1967)

Note: Puto orthezioides ( Cockerell, 1903) is known only from a third-instar female and is not included in the key.

1(0) Circulus complete, without constriction or division, or circulus absent............................................ 2

- Circulus partially or completely divided into two parts....................................................... 27

2(1) Circulus absent....................................................................................... 3

- Circulus present...................................................................................... 4

3(2) Ostioles without conical setae on lips; anal-lobe cerarius usually with fewer than 30 cerarian setae.................................................................................... Puto kosztarabi Miller & Miller, 1993 (in part)

- Ostioles with conical setae on lips; anal-lobe cerarius usually with more than 30 cerarian setae.................................................................................................... Puto acirculus McKenzie, 1960

4(2) Cerarii without oral-rim type tubular ducts in sclerotized area....................................................5

- Cerarii with oral-rim type tubular ducts in sclerotized area (beware of dorsal oral-rim type tubular ducts adjacent to, but not in, sclerotized area)..................................................................................... 16

5(4) Circulus oval or circular, approximately as wide as long, less than three times as wide as long, often with smooth margin... 6 Circulus laterally expanded, often three times as wide as long, usually with crenulate margin.......................7

6(5) Medial or mediolateral areas with oral-rim type tubular ducts.................. Puto pacificus McKenzie, 1967 (in part)

- Medial or mediolateral areas without oral-rim type tubular ducts............................ Puto bryanthi Ferris, 1950

7(5) Twenty-five or more conical setae in each anal-lobe cerarius.....................................................8

- Less than 25 conical setae in each anal-lobe cerarius............................................................9

8(7) Abdomen without ventral submarginal multilocular pores.......................................................40

- Abdomen with ventral submarginal multilocular pores......................... Puto barberi ( Cockerell, 1895) (in part)

9(7) More than five multilocular pores in area between meso- and metacoxa............................................. 10

- Fewer than five multilocular pores in area between meso- and metacoxa...........................................13

10(9) Abdominal segment VI with multilocular pores near anterior margin............................................ 11

- Abdominal segment VI without multilocular pores near anterior margin............... Puto atriplicis McKenzie, 1961

11(10) Ostioles with setae on lips.................................................................................12

- Ostioles without setae on lips........................................ Puto kosztarabi Miller & Miller, 1993 (in part)

12(11) Anal ring with six setae, setae with acute apices; abdominal segment VII without multilocular pores near anterior margin.................................................................... Puto calcitectus ( Cockerell, 1901)

- Anal ring with more than six setae, setae with conspicuously capitate apices; abdominal segment VII with multilocular pores near anterior margin................................................... Puto decorosus McKenzie, 1967 (in part)

13(9) Ventral multilocular pores on most abdominal segments...................................................... 14

- Ventral multilocular pores restricted to abdominal segments VII and VIII............... Puto profusus McKenzie, 1967

14(13) Anal-lobe cerarii each with fewer than 16 conical setae; apices of anal-ring setae acute..........................15

- Anal-lobe cerarii each with 16 or more conical setae; apices of anal-ring setae sometimes slightly enlarged............................................................................. Puto californicus McKenzie, 1967 (in part)

15(14) Multilocular pores scattered on thorax and head; length of antennae longer than 1.1 mm ...................................................................................................... Puto pricei McKenzie, 1960

- Multilocular pores on thorax restricted to around spiracles; length of antennae shorter than 1.1mm .................................................................................................. Puto sandini Washburn, 1965

16(4) Medial or mediolateral areas of dorsum with oral rim-type tubular ducts...........................................17

- Medial or mediolateral areas of dorsum without oral-rim type tubular ducts.........................................22

17(16) Venter of thorax with multilocular pores abundant, with more than five in area between meso- and metacoxa.......... 18

- Venter of thorax with multilocular pores sparse or absent, with five or less in area between meso- and metacoxa.........37

18(17) Cerarii on head without associated oral-rim type tubular ducts................................................. 19

- Cerarii on head with associated oral-rim type tubular ducts................................................... 20

19(18) With multilocular pores scattered around each spiracle; without ventral multilocular pores on margins of anterior abdominal segments.................................................................... Puto cupressi ( Coleman, 1908)

- With conspicuous cluster of multilocular pores around each spiracle; with numerous multilocular pores on margins of anterior abdominal segments................................................. Puto laticribellum McKenzie, 1961 (in part)

20(18) Without clusters of oral-rim type tubular ducts surrounding cerarii................................................21

- With clusters of oral-rim type tubular ducts surrounding cerarii....................... Puto salinasi Foldi & Kozár, 2006

21(20) With fewer than 25 oral-rim type tubular ducts on dorsum excluding those in cerarii........................................ Puto echinatus McKenzie, 1961 (in part) (San Miguel (in part), Santa Cruz, and San Nicholas Islands, winged male) With more than 25 oral-rim type tubular ducts on dorsum excluding those in cerarii............................................................................ Puto echinatus McKenzie 1961 (in part) (inland mainland, winged male)

22(16) Cerarii on head each with oral-rim type tubular ducts in sclerotization.............................................23

- Cerarii on head each without oral-rim type tubular ducts in sclerotization..........................................38

23(22) Multilocular pores present in submarginal areas of abdominal segments III–VI.....................................24

- Multilocular pores absent from submarginal areas of abdominal segments III–VI.................................. 26

24(23) Anal-lobe cerarii each smaller than hind coxae............................................................. 25

- Anal-lobe cerarii each larger or equal in size to hind coxae............................ Puto arctostaphyli Ferris, 1950

25(24) Anal-lobe cerarii each with fewer than eight oral-rim type tubular ducts; tubular ducts in anal-lobe cerarii each about same diameter as setal socket of largest conical seta.............................................................. 39

- Anal-lobe cerarii each with eight or more oral-rim type tubular ducts; tubular ducts in anal-lobe cerarii each twice as large as diameter of setal socket of largest conical seta.................................. Puto brom Powell & Miller , sp. n.

26(23) Anal-lobe cerarii each normally with more than five oral-rim type tubular ducts.......... Puto mexicanus ( Cockerell, 1893)

- Anal-lobe cerarii each normally with fewer than five oral-rim type tubular ducts........... Puto yuccae ( Coquillett, 1890)

27(1) Oral-rim type tubular ducts present within confines of sclerotization of cerarii......................................28

- Oral-rim type tubular ducts absent from within confines of sclerotization of cerarii................................. 30

28(27) Circulus with constriction but not divided into two pieces.................................................... 29

- Circulus divided into two pieces............................................... Puto antioquensis ( Murillo, 1931)

29(28) Claw and tarsal digitules acute; claw digitules not reaching tip of claw.................... Puto usingeri McKenzie, 1962

- Claw and tarsal digitules clubbed; claw digitules extending beyond tip of claw........... Puto ulter Ferris, 1950 (in part)

30(27) Circulus divided into two or more pieces....................................................................31

- Circulus with constriction but not divided into two pieces..................................................... 36

31(30) Claw denticle present................................................................................. 32

- Claw denticle absent.....................................................................................35

32(31) Circulus on abdominal segment III only; ventral multilocular pores scattered over entire surface.................. 33

- Circuli on abdominal segments III and IV; ventral multilocular pores restricted to abdomen and near spiracles..................................................................................... Puto lamottei Matile-Ferrero, 1985

33(32) Ostioles each poorly developed or absent, lips with few or no conical setae........................................34

- Ostioles each well developed, lips with numerous conical setae.................... Puto spath Powell & Miller , sp. n.

34(33) Tarsal and claw digitules apically enlarged.................................. Puto bicirculatus Gavrilov-Zimin, 2023

- Tarsal and claw digitules apically acute................................ Puto kosztarabi Miller & Miller, 1993 (in part)

35(31) Area between meso- and metacoxa with multilocular pores rare or absent............... Puto philo Powell & Miller , sp. n.

- Area between meso- and metacoxa with multilocular pores abundant............. Puto paramoensis Matile-Ferrero, 1985

36(30) Tarsal digitules simple; body margin on anterior abdominal segments with multilocular pores; usually with 17 pairs of cerarii, frontal cerarius usually comprising fusion of frontal and antefrontal cerarii......... Puto barberi ( Cockerell, 1895) (in part)

- Tarsal digitules clubbed; body margin on anterior abdominal segments and thorax with multilocular pores absent; usually with 18 pairs of cerarii, frontal cerarii and antefrontal cerarii separate........................ Puto ulter Ferris, 1950 (in part)

37(17) Ventral abdominal segment VII with posterior row of multilocular pores only........... Puto simmondsiae McKenzie, 1961 Ventral abdominal segment VII with anterior and posterior rows of multilocular pores.................... Puto echinatus McKenzie, 1961 (in part) (coastal, with associated wingless male; inland mainland, with associated winged male; San Miguel (in part) and Santa Barbara Islands, with associated winged male)

38(22) Submarginal areas of abdomen with multilocular pores abundant; anal-ring setae apically acute; occurring in mountainous areas on conifers........................................................ Puto laticribellum McKenzie, 1961 (in part)

- Submarginal areas of abdomen with multilocular pores absent or sparse; anal-ring setae apically enlarged; occurring in salt marshes on Sarcocornia ................................................ Puto ambiguus ( Fullaway, 1910) (in part)

39(25) Submarginal area of thorax and head with multilocular pores sparse or absent; dorsomedial setae on posterior abdominal segment VII with basal sclerotization............................................ Puto lasiorum ( Cockerell, 1901) Submarginal area of thorax and head with multilocular pores abundant; dorsomedial setae on abdominal segment VII without basal sclerotization............................................................ Puto albicans McKenzie, 1967

40(8) Anal ring with three pairs of setae..........................................................................41

- Anal ring with more than three pairs of setae................................ Puto decorosus McKenzie, 1967 (in part)

41(40) Area between meso- and metacoxa with 5 or more multilocular pores............ Puto ambiguus ( Fullaway, 1910) (in part)

- Area between meso- and metacoxa with less than 5 multilocular pores......... Puto californicus McKenzie, 1967 (in part)