A NEW SPECIES OF PUTa AND A PRELIMINARY ANALYSIS OF ...
A NEW SPECIES OF PUTa AND A PRELIMINARY ANALYSIS OF ... A NEW SPECIES OF PUTa AND A PRELIMINARY ANALYSIS OF ...
32 MIllER & MIllER: PUfO KOSZTARABI (convergent in Pityococcus ferrisl)j ostioles [12.1); abdominal spiracles absent [17.1); ventral plate absent [18.1.j (convergent in the clade Kermesidae + Eriococcidae)j adult female with: translucent pores on hind legs [20.lI (convergent in Ovaticoccus agavium [Douglas], Acanlhococcus araucariae [Maskell], Acanlhococcus droserae Miller, Iiu, & Howell, and Stibococcus cerinus Miller & Gonzalez); abdominal spiracles absent [24.1] (convergent in the clade Kermesidae + Eriococcidae); medial circulus [28.1j (convergent in Pityococcus ferris,'; first instar with: ostioles [35.).); simple disk pores [36.1.]. Stem 11. Adult male with: incomplete preocular ridge [9.a] (convergent in Eriococcidae); capitate tarsal digitules [10.1] (convergent in the clade Kermesidae + Eriococcidae); 2 pairs of eyes [11.2] (convergent in Eriococcidae); adult female with: 2 trochanter sensoria on each surface [32.1j (convergent,Jn the clade Kermesidae + Eriococcidae). Stem 12 (Pulo group). Adult male with: triangular scutum [4.b] (convergent in Sleingelia gorodetskia); 3-4 halter setae [14.0, a reversal]j leg setae associated with small basal seta [15.11; first instar with: 7· segmented antennae [38.1] (convergent in Pityococcus ferrisi and Phenacoleachia zealandica). CONCLUSIONS The Pulo group could be treated either as a separate family or as part of the Pseudococcidae. Our preference is for the inclusion of this group as part of the mealybugs since the only non-homoplasious character that defines the group is the presence of associated basal setae with the larger leg setae on the adult males. Although this is unique to Pulo, Phenacoleachia males have similar setae on the antennae. Our analysis confirms the problematic placement of S/eingelia and Pityococcus. The presence of spiraled trilocular pores, circuli, and ostioles suggested placement of Pityococcus in the Pseudococcidae (Miller, 1983). However, this affinity is not borne out in the current study. In addition, Sleingelia, which has pseudococcid-like genitalia and lateral tail·forming pore clusters in the adult male, is also placed separately from the mealybugs. Because we did not include a detailed analysis of the Margarodidae and because these groups are purported to belong to this family, we will not make any classification changes until a more detailed analysis can be completed. The relationships of the pseudococcids, eriococcids, and kermesids is
JEFFERSONIANA 33 puzzling. The present analysis includes kermesids and eriococcids as sister groups, but in a previous study (Miller & Miller, in press) the pseudococcids and eriococcids were hypothesized as sister groups. In the present paper, we included additional taxa and characters. Eight characters support the monophyli of the eriococcids and kermesids. Unfortunately, only two of these are non-homoplasious, and the majority of the homoplasy is in the form of convergences shared with the pseudococcid clade or the pseudococcid + the Puto group clade. We favor the results presented here since a complete penial sheath and invaginated tubular duets are complicated structures unlikely to have developed independendy, but this aspect of the phylogeny requires additional srudy. This work treats the adult males of more congeneric species than in any previous Coccoidea research. In the process of our examination, we studied adult males of 17 species of Puto including more than 50 specimens. Because many species are represented by only one or two specimens, the results must be considered preliminary, but several conclusions merit special attention. There are two distinct groups of species within the genus: the first group (Puto antennatus, P. cupressi, P. sandini, and P. superbus) occurs on conifers and has a short, non· articulated aedeagus with a blunt apex similar to other pseudococcids; the other group (Puto albicans, P. ambiguus, P. antioquensis, P. arctostaphyli, P. atriplicis, P. calcitectus, P. decorosus, P. echinatus, P. kosztarabi, P. mexicanus, P. simmondsiae, P. uTter, and P. yuccae) occurs on other hosts and has a long, articulated aedeagus with a bifurcate tip. The opinion of Williams & de Willink (1992) that Macrocerococcus and Puto are synonyms is supported in this paper. However, if the conifer·infesting group were considered a separate genus, its generic name would be Macrocerococcus. We predict that other conifer-infesting species such as P. laticribellum, P. pricei, and P. profusus also will have males with a non-articulated aedeagus and a blunt apex. The species P. kosztarabi and P. calcitectus have males with two teeth that are removed from the apex of the aedeagus. The remaining species of Puto occurring on non-
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32 MIllER & MIllER: PUfO KOSZTARABI<br />
(convergent in Pityococcus ferrisl)j ostioles [12.1); abdominal spiracles<br />
absent [17.1); ventral plate absent [18.1.j (convergent in the clade<br />
Kermesidae + Eriococcidae)j adult female with: translucent pores on hind<br />
legs [20.lI (convergent in Ovaticoccus agavium [Douglas], Acanlhococcus<br />
araucariae [Maskell], Acanlhococcus droserae Miller, Iiu, &<br />
Howell, and Stibococcus cerinus Miller & Gonzalez); abdominal<br />
spiracles absent [24.1] (convergent in the clade Kermesidae + Eriococcidae);<br />
medial circulus [28.1j (convergent in Pityococcus ferris,'; first<br />
instar with: ostioles [35.).); simple disk pores [36.1.].<br />
Stem 11. Adult male with: incomplete preocular ridge [9.a] (convergent<br />
in Eriococcidae); capitate tarsal digitules [10.1] (convergent in the clade<br />
Kermesidae + Eriococcidae); 2 pairs of eyes [11.2] (convergent in<br />
Eriococcidae); adult female with: 2 trochanter sensoria on each surface<br />
[32.1j (convergent,Jn the clade Kermesidae + Eriococcidae).<br />
Stem 12 (Pulo group). Adult male with: triangular scutum [4.b]<br />
(convergent in Sleingelia gorodetskia); 3-4 halter setae [14.0, a reversal]j<br />
leg setae associated with small basal seta [15.11; first instar with: 7·<br />
segmented antennae [38.1] (convergent in Pityococcus ferrisi and<br />
Phenacoleachia zealandica).<br />
CONCLUSIONS<br />
The Pulo group could be treated either as a separate family or as part<br />
of the Pseudococcidae. Our preference is for the inclusion of this group<br />
as part of the mealybugs since the only non-homoplasious character that<br />
defines the group is the presence of associated basal setae with the larger<br />
leg setae on the adult males. Although this is unique to Pulo, Phenacoleachia<br />
males have similar setae on the antennae.<br />
Our analysis confirms the problematic placement of S/eingelia and<br />
Pityococcus. The presence of spiraled trilocular pores, circuli, and<br />
ostioles suggested placement of Pityococcus in the Pseudococcidae<br />
(Miller, 1983). However, this affinity is not borne out in the current<br />
study. In addition, Sleingelia, which has pseudococcid-like genitalia and<br />
lateral tail·forming pore clusters in the adult male, is also placed separately<br />
from the mealybugs. Because we did not include a detailed analysis of the<br />
Margarodidae and because these groups are purported to belong to this<br />
family, we will not make any classification changes until a more detailed<br />
analysis can be completed.<br />
The relationships of the pseudococcids, eriococcids, and kermesids is
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