A review of neococcid scale insects (Hemiptera: Sternorrhyncha: Coccomorpha) based on the morphology of the adult males
Author
Hodgson, Chris
text
Zootaxa
2020
2020-04-16
4765
1
1
264
journal article
22494
10.11646/zootaxa.4765.1.1
8af5a092-f062-48c7-8d7c-f96ae6282bfa
1175-5334
3774174
urn:lsid:zoobank.org:pub:C442D94C-0EB4-4509-B762-913707214819
RHIZOECIDAE
Williams, 1969
Rhizoecini
Williams 1969
, 335
.
Rhizoecinae
Williams 1969
, 335
;
Koteja 1974b
, 45
;
Tang 1992
, 42
.
Rhizoecidae Williams
;
Hodgson, 2012
, 4
.
Type
genus:
Rhizoecus
Künkel d’Herculais.
Type
species:
Rhizoecus falcifer
Künkel d’Herculais 1878
, 164 by monotypy.
FIGURE 2
.
Rhizoecus coffeae
Laing. Macropterous
male. (
Rhizoecidae
, Rhizoecina). Where A=hair-like seta, B=fleshy seta, C=loculate pores, K=proximal end of metathoracic leg, and Z=mouth tubercle. For abbreviations, see p. 8.
FIGURE 3.
Ripersiella hibisci
(Kawai & Takagi)
. Macropterous male. (
Rhizoecidae
, Ripersiellina). Where A=hair-like seta, B=fleshy seta, C =loculate pores, F=alar sensoria, and K=proximal end of metathoracic leg. For abbreviations, see p. 8.
Introduction.
The taxa with the hypogaeic and myrmecophilous mealybugs have, until recently, been included in the
Pseudococcidae
as the subfamily
Rhizoecinae
. However, a phylogenetic study based on DNA sequences and including representatives from all five pseudococcid subfamilies, found that the
Rhizoecinae
(represented by 4 species in 3 genera) formed a monophyletic group sister to the remaining pseudococcids (
Downie & Gullan 2004
). Later, based on adult male characters only,
Hodgson and Foldi (2005)
found that
Rhizoecus
formed a separate clade with
Pityococcus
, sister to the
Pseudococcidae
. Since then, also largely based on the morphology of the adult males,
Hodgson (2012)
reviewed the relationships of the
Pseudococcidae
and the hypogaeic and myrmecophilous mealybugs, and concluded that the rhizoecine mealybugs formed a separate family from the
Pseudococcidae
(
Rhizoecidae Williams
) and that this family included two subfamilies,
Rhizoecinae Williams
and Xenococcinae Tang. A more recent study (
Hodgson & Hardy 2013
), also based on adult male morphology, also found the
Rhizoecidae
to be a separate clade, sister to the
Pseudococcidae
. However,
Danzig and Gavrilov-Zimin (2014)
did not accept this and argued that: (a) the hypogaeic mealybugs belonged within the
Pseudococcidae
, possibly sister to
Mirococcopsis
Borchesenius
, and that (b) the myrmecophilous Xenococcinae were unrelated to the
Pseudococcidae
and might be closest to the
Margarodidae
s.l
. Danzig and Gavrilov-Zimin therefore raised the Xenococcinae to family status, Xenococcidae Tang, although this was without phylogenetic support. Whilst it is still the view of the present author that the
Rhizoecidae
are sister to the
Pseudococcidae
, he agrees that the relationships of the Xenococcinae are problematic and that it should be raised to family status. As understood here, the
Rhizoecidae
contains 16 genera and 214 species (
García Morales
et al
. 2019
).
Family diagnosis based on adult male morphology
. Many species
apterous
but some with
brachypterous
or fully
macropterous
adult males. Unless otherwise stated, diagnosis refers to all morphs.
Body
generally
small to minute
, cylindrical, somewhat pointed at both ends.
Head
without ocelli
, although two pairs of simple eyes generally present; preocular ridges generally recognisable ventrally although often not strongly developed;
postocular ridge absent although a faint line occasionally present marking posterior margin of ocular sclerite
;
postoccipital ridge absent
;
dorsal midcranial ridge absent
; ventral midcranial ridge poorly developed or absent; base of antennae placed close together near anterior margin of head; number of antennal segments variable, each 3–10 segmented, with fs and hs setae on most segments;
capitate setae on antennae generally absent or not differentiated
;
neck entirely absent
.
Loculate pores frequently present on head, thorax and/or abdomen
.
Thorax
.
Pronotal ridge and pronotal sclerite absent
; propleural ridge short, at most only extending anteriorly about half-way to head; apterous species without any mesothoracic sclerotisation. Metaprecoxal ridge absent; leg setae hs or spinose, not fs;
each trochanter with 3 round sensoria arranged in a curve or triangle on each side
; trochanter often divided by a Y-shaped sclerotisation;
tarsi with 2 spurs
; tarsi generally clearly 2 segmented, occasionally 1 segmented; tarsal digitules often undifferentiated;
claws long and narrow with setose digitule
s.
Abdomen.
Ostioles absent
;
abdominal segment VIII without glandular pouches and without a group of disc-pores
, although 1 or more long setae may be present in this position; abdominal segment IX not apparently differentiated, but position indicated by anal opening found dorsally at anterior end of dorsal ridge on penial sheath; ventral opening of penial sheath often indistinct and never with lateral finger-like processes;
penial sheath about as broad as long
; penial sheath without a large, U-shaped, sclerotised structure extending anteriorly within abdomen from base of aedeagus (if an obvious internal sclerotisation present, this never as long as penial sheath
). Macropterous and brachypterous species
also with: scutum without a membranous area; basisternum without a median ridge;
alar lobes
present on each wing but
hamulohalteres absent
;
alar setae present
; alar sensoria present or absent.
Discussion
. In his 2012 paper, Hodgson found that the morphology of the adult males of the
Rhizoecidae
was very variable, particularly in the number of antennal segments, and that male morphology did not entirely support the classification based on the morphology of the adult female of
Kozár & Konczné Benedicty (2007)
. The males of the macropterous rhizoecine mealybugs,
Rhizoecus coffeae
Laing
(
Fig. 2
),
Ripersiella hibisci
(
Figs 2
&
3
) and
Rhizoecus dianthae
Green
(
Fig. 4
), plus an apterous rhizoecine mealybug,
Kissrhizoecus hungaricus
Kozár & Konczné Benedicty
(
Fig. 5
), are illustrated here to illustrate the diversity of the family. A key to the described rhizoecine adult males plus descriptions and other illustrations can be found in
Hodgson (2012)
.