Review of the Neotropical scale insects formerly assigned to Coelostomidiidae and here transferred to a new tribe within the Monophlebidae (Hemiptera: Sternorrhyncha: Coccoidea).

This study reviews the status of all Neotropical genera and species of Coelostomidiidae (Hemiptera: Coccoidea) and transfers them to the family Monophlebidae in the Cryptokermesini Foldi & Gullan tribe n. (the tribe Cryptokermini Tao & Hao is recognised here as a nomen nudum). This change of family placement for Neotropical taxa is based on the morphology of adult males, as supported by the phylogenetic study of Hodgson & Hardy (2013), and by unpublished DNA data. New diagnoses are provided for each of the four recognised genera of Cryptokermesini: Cryptokermes Hempel, Mimosicerya Cockerell, Neocoelostoma Hempel and Paracoelostoma Morrison. The genus Nautococcus Vayssière is considered here to be a junior synonym (syn. n.) of Mimosicerya and the type species of Nautococcus, N. schraderae Vayssière, thus becomes M. schraderae (Vayssière) comb. n. Cryptokermes mexicanus Morrison is transferred to Mimosicerya as M. mexicana (Morrison) comb. n. Also Cryptokermes mimosae Foldi does not fit the morphological concept of Cryptokermes and is excluded from this genus and revision, and from the new tribe; its taxonomic position is uncertain and requires further study. All type species of the Cryptokermesini, including N. schraderae (as M. schraderae), are redescribed and illustrated based on most female instars and available adult males, examined using optical and scanning electron microscopes. Adult males are described and illustrated only for M. schraderae and N. xerophila. Keys are provided to distinguish the Neotropical monophlebid tribes Cryptokermesini and Llaveiini and to recognise each cryptokermesine genus based on female instars and first-instar nymphs. The included species of Cryptokermesini and their known distributions are: Cryptokermes brasiliensis Hempel from Brazil and C. oaxaensis Foldi from Mexico; Mimosicerya hempeli (Cockerell) from Brazil, M. mexicana from Mexico, M. schraderae from Panama and M. williamsi Foldi from Venezuela; Neocoelostoma xerophila Hempel from Argentina, Bolivia, Brazil, Paraguay and Uruguay; and Paracoelostoma peruvianum Morrison from Peru. All these insects live exposed on their host plant, either inside a secreted test (as for female and immature male instars of Cryptokermes, Neocoelostoma and Paracoelostoma) or the strongly sclerotised derm of the preadult female protects the adult (as for all species of Mimosicerya). Adult females of Mimosicerya are pupillarial, remaining within the exuviae of the previous instar, whereas adult females of the other three genera either remain within their test (and some species may be pupillarial) or escape the test to oviposit. The morphology of the adult female and often the preadult female is strongly modified, with reduction of antennae and legs, and with legs lacking in some species. 


Introduction
The Coccoidea, or scale insects, are small sap-sucking insects with about 8000 species known worldwide and including many of economic importance . Extreme sexual dimorphism exists between the paedomorphic females and dipteriform males. These small insects are divided into two informal groups, the archaeococcoids and the neococcoids, based on the presence or absence of features interpreted as ancestral or derived (Foldi, 2005;Gullan & Cook, 2007). The archaeococcoids are characterised by the presence of abdominal spiracles in all instars, compound eyes in adult males and an XX-XO sex determination system (features found in most other Hemiptera), whereas the neococcoids lack abdominal spiracles in all instars, lack compound eyes in adult males and have the synapomorphic paternal genome elimination (PGE) system (Cook et al., 2002;Normark, 2003;Ross et al., 2010) and share derived DNA sequences (Cook et al., 2002;Gullan & Cook, 2007;Yokogawa & Yahara, 2009). The archaeococcoids comprise up to 15 extant families and the neococcoids up to 18 extant families (Foldi, 2005;Gullan & Cook, 2007;Ben-Dov et al., 2014). Although neococcoids have been shown to form a monophyletic group, the relationships of most archaeococcoid higher taxa are not resolved at present (Foldi, 1997;Cook et al., 2002;Gullan & Cook, 2007;Hodgson & Foldi, 2005;Hodgson & Hardy, 2013). Furthermore, the affiliations of some archaeococcoid genera and even their family-level status are uncertain. However, the continuing acquisition and analysis of DNA sequence data, and phylogenetic analyses based on adult female and male morphology are helping to reconstruct scale insect family relationships.
Originally,  erected the subfamily Coelostomidiinae with three tribes: Coelostomidiini, Marchalini and Platycoelostomini, with the New Zealand Coelostomidia Cockerell as the type genus of Coelostomidiini. Koteja (1974) elevated the subfamily to family rank as Coelostomidiidae, but later Koteja (1996) narrowed the family to include just members of Morrison's Coelostomidiini, and this classification is accepted widely now (Foldi, 2005;Hodgson & Foldi, 2006;Gullan & Cook, 2007). Currently the Coelostomidiidae includes Coelostomidia (six species) and Ultracoelostoma Cockerell (three species) from New Zealand (Morales, 1991), and Cryptokermes Hempel (four species), Nautococcus Vayssière (one species), Neocoelostoma Hempel (one species), Mimosicerya Cockerell (two species), and Paracoelostoma Morrison (one species) from the Neotropical area (Mexico, Central and South America) Williams & Gullan, 2008;Foldi, 2009). The validity of the genus Nautococcus (Vayssière, 1939) from Panama was questioned by Williams & Gullan (2008), who suggested that Mimosicerya and Nautococcus are identical, although the described species are not the same. They excluded Nautococcus from the tribe Llaveiini of the Monophlebidae. It was placed in Monophlebidae by Ben-Dov (2005, 2014. Vayssière and Hughes-Schrader (1948) commented that Nautococcus undoubtedly belonged with the coelostomidiids based on morphology but that it had cytology (2n = 6 in females; 2n = 5 in males) similar to Llaveiini. However, there is no information on karyotypes in coelostomidiids.
Most Neotropical coelostomidiids are test forming (Figs 1A,D,E & F), with the immature insects secreting a thick-walled protective test that enlarges during their development. At maturity, the adult female either stays and oviposits within the test or exits and oviposits elsewhere. In contrast, species of Mimosicerya do not secrete a test but the preadult cuticle becomes heavily sclerotised and forms a protective cover for the adult female ( Fig. 1B, C), which can, therefore, be considered as "pupillarial" because it remains and oviposits within the exuviae of the previous instar.  noted that the anal tube of the preadult females of Cryptokermes and Paracoleostoma is often retained within that of the adult, which may suggest that they are pupillarial too. In Neocoelostoma, the adult female escapes from the test to seek a sheltered position on the bark, and exudes a mass of cottony white filaments prior to ovipositing within (from notes of H.L. Parker associated with specimens in USNM). Related to this behaviour, the adult female of Neocoelostoma has well-developed legs, as typical in monophlebids, whereas the adult females of Cryptokermes, Mimosicerya and Paracoelostoma have highly reduced legs and those of Nautococcus are completely absent.
This paper revises the classification of the Neotropical archaeococcoid genera that currently are placed in the family Coelostomidiidae. Nautococcus schraderi Vayssière, the only species in Nautococcus, shares a number of morphological features with the other genera, as outlined by Williams & Gullan (2008) and documented in this paper. We formally recognise Nautococcus as a junior synonym of Mimosicerya and thus N. schraderae is transferred to Mimosicerya. We also formally transfer Cryptokermes, Mimosicerya, Neocoelostoma and Paracoelostoma to the Monophlebidae in their own tribe, and provide keys to the genera based on the morphology of adult and preadult females and first-instar nymphs. These genera are close to the Neotropical tribe Llaveiini (as defined by Williams & Gullan (2008)), but in the Llaveiini, no genus forms a test and nor do adult females have such reduced appendages.

Material, methods and terminology
Materials. Most specimens used in this study were borrowed from museums. Extensive use was made of the dry collections in the USNM, especially those made in the early 1940s by H.L. Parker, who also made notes on the insects in life. Some specimens also were collected in South America by P.J. Gullan and I. Foldi. The following abbreviations are used for museum collections: BME: The International Code of Zoological Nomenclature (ICZN, 1999) requires the designation of lectotypes after 1999 to "contain an express statement of deliberate designation" (amended Article 74.7.3). We use the statement "here designated" to fulfill this requirement. A lectotype has been designated for Nautococcus schraderae Vayssiere because this name lacks a holotype or lectotype and unambiguous syntypes have been identified. The purpose is to provide stability of nomenclature, and designation is done in a revisionary context in agreement with the amended Recommendation 74G of Article 74.7.3. We have not designated a lectotype for either Cryptokermes brasiliensis Hempel or Icerya (Crypticerya) hempeli Cockerell, as we explain in the type notes for those two names.
Methods. Some dry museum material and the freshly collected specimens were slide-mounted in Canada balsam using the method described in Williams & Granara de Willink (1992) except that xylene was used instead of clove oil. Scanning electron microscopy (SEM) was used to interpret dermal microstructures. For SEM study, the whole insects were treated in KOH (10% in water), washed in water and dehydrated in a series of alcohols -50 %, 70 % to absolute alcohol, cleaned in an ultrasonic cleaner, critical point-dried and finally the specimens were coated in gold. Slide-mounted specimens were measured under a compound microscope; body length and width were recorded in mm, whereas other measurements of various useful features were made in microns. Length is measured from the apex of the head to the posterior end of the body, whereas width is measured as the greatest width. Each figure shows an entire insect with the venter depicted on the right side of the illustration and the dorsum shown on the left. Special features of the specimen are enlarged around the main illustration, although enlargements are not in direct proportion to each other.
For second-instar nymphs, we were unable to determine whether the specimens examined were male or female, nor whether there is any sexual dimorphism in the second instar. Larger collections of this instar are required.
Terminology. The terms used to describe the adult female and immature insects follow those of  and Foldi (2001Foldi ( , 2009, and for the adult male those of Hodgson & Foldi (2006). The term cicatrix refers to an approximately circular structure with a membranous surface surrounded by a sclerotised rim; the number, size and distribution of cicatrices are useful taxonomic characters. In the genera dealt with here, there are large variations in cicatrix size and mostly the surface appears smooth. Disc-like tubercles are small circular structures that in side view have a slight to pronounced dome.  described disc-like and spine-like tubercles on the derm of first-instar nymphs and preadult females of Cryptokermes species, and on the first-instar nymph of Mimosicerya hempeli. In the preadult female of M. hempeli, there are two kinds of disc-like tubercles: one smaller type, more densely distributed and with no central opening (Fig. 7G), and another of larger size, less common but with a visible central opening (Fig. 7J). Derm pores are named according to the presence or absence of loculi: simple pores have no loculi, whereas loculate pores can be uni-, bi-, tri-, quadri-or quinquelocular (i.e., from one to five loculi), or multilocular if there are six or more outer loculi. Raised pores are each located on a raised, circular sclerotised area of derm that has a sunken centre in which is located the multilocular pore (see Figs 3M, 22H).

Phylogenetic relationships and a revised taxonomic classification
The adult females and immature instars of the South American and New Zealand Coelostomidiidae share a number of morphological features as described by . The adult females exhibit much variation among the genera especially in antennal and leg development and degree of sclerotisation, but the two New Zealand genera differ from the South American species in having a well-developed anal tube bearing hairs, pores and sometimes lobes, and always have a sclerotised inner ring lacking polygonal pores (Morales, 1991), whereas the anal tube in the Neotropical genera is probably always membranous but often retains the tube of the previous instar, which has a band of polygonal pores at the inner end. The second-and third-instar females of the New Zealand and South American coelostomidiid genera share the following morphological features: posterior end of the abdomen surrounding anal opening with derm sclerotised to varying degrees; antennae and legs reduced in size but segments usually distinct; anal tube long with a ring or band of polygonal wax pores at inner end; cicatrices present and usually present on both dorsum and venter; however, disc-like tubercles are absent in the New Zealand species. The first-instar nymphs of New Zealand coelostomidiids have simple pores (Morales, 1991) but lack the distinctive disc-like tubercles present in some of the South American genera, and there is only one pair of caudal setae in New Zealand species whereas the South American species have two to many pairs, as in the first-instar nymphs of most Monophlebidae.
A relationship between the Coelostomidiidae (sensu Koteja, 1996;Hodgson & Hardy, 2013) and the Monophlebidae was suggested by Gullan & Sjaarda (2001), based on a cladistic analysis of morphological data from 27 archaeococcoid genera. A recent work (Hodgson & Hardy, 2013) on the phylogeny of Coccoidea, based on the morphology of macropterous males, found the Coelostomidiidae to be paraphyletic with respect to Monophlebidae. The included South American coelostomidiid representatives, Nautococcus (synonymised with Mimosicerya later in this paper) and Neocoelostoma, formed a clade sister to Monophlebidae rather than being grouped with the New Zealand members of Coelostomidiidae; the authors suggested that these South American taxa could be treated either as a separate family-level group or transferred to the Monophlebidae. The available adult males of the Neotropical coelostomidiids show a combination of features found in the Monophlebidae and the New Zealand coelostomidiids, with the adult male of Neocoelostoma resembling those of monophlebids such as Laurencella colombiana Foldi & Watson and Drosicha dalbergiae (Stebbing) more than does the adult male of Nautococcus. Neocoelostoma and Nautococcus share the following adult male apomorphies with the Monophlebidae but not with the New Zealand Coelostomidiidae (character states typical of NZ taxa in parentheses): ocelli located dorsal to compound eyes (ocelli posterior to compound eyes); claws with one pair of non-capitate, usually setose, digitules (claws with capitate digitules and often more than one pair); paired caudal extensions on at least one posterior abdominal segment (lacking caudal extensions); anus at apex of a sclerotised tube (anal tube apparently membranous) (Hodgson & Foldi, 2006: Hodgson & Hardy, 2013. Preliminary analysis of nucleotide sequence data (L.G. Cook, T. Kondo and P.J. Gullan, unpublished) from the small subunit ribosomal RNA gene (SSU rRNA or 18S) and including taxa of New Zealand and Neotropical Coelostomidiidae, Australian and Neotropical Monophlebidae and representatives from other archaeococcoid families, show the Neotropical coelostomidiids (Nautococcus, Neocoelostoma and Cryptokermes) to be related to the monophlebid tribes Llaveiini (Laurencella and Protortonia) and Monophlebulini (Monophlebulus and Melaleucoccus) and to be unrelated to the New Zealand coelostomidiids.
Although the insects of most genera of coelostomidiids (both New Zealand and South American) develop within a test on their host plant, the nature of the test appears to differ between the New Zealand and Neotropical genera, with those of the latter typically being composed of a yellow to orange, brown or reddish-brown resin and fully exposed on the plant, whereas the New Zealand species have tests that are more waxy and often are at least partially hidden in the bark. Similarities in female morphology between New Zealand and South American taxa presumably result from convergent adaptation to living confined within a test. Examples of such adaptations are (i) the reduction of appendages in instars with limited or no mobility, and (ii) the apical anal opening and its associated pores and well-developed anal tube to allow efficient elimination of honeydew from the test.
Here we formally transfer the genera Cryptokermes, Mimosicerya (including Nautococcus), Neocoelostoma and Paracoelostoma to the Monophlebidae and place these four genera into their own tribe, which is named and described below.  Tang & Hao (1995) used the name Cryptokermini to include four genera and five species in the subfamily Coelostomidiinae (which they included in Monophlebidae), but they did not specify which genera and species were included nor did they provide a description or definition of their new tribe. Their name is thus unavailable (see Article 13 of the International Code of Zoological Nomenclature (ICZN 1999)). Williams (2013) emended the name Cryptokermini to Cryptokermesini but did not realise that the earlier name was invalid. Here we validate the name Cryptokermesini by providing a description of the tribe and a key to the included genera of the tribe.

Diagnosis of tribe Cryptokermesini
Species of Cryptokermes, Neocoelostoma and Paracoelostoma are test forming (Fig. 1A, D, E & F), whereas in Mimosicerya, the immature insects develop exposed on the host plant and the preadult cuticle becomes heavily sclerotised as a protective cover for the pharate adult female (Fig. 1B, C). Adult female. Major morphological features such as a sclerotised zone on the head, presence or absence of mouthparts and degree of development of antennae and legs vary among females of the different species and genera. Cryptokermes and Mimosicerya have a large, sclerotised zone on the head, which extends onto the thorax. In contrast, the adult females of Neocoelostoma and Paracoelostoma lack a sclerotised zone anteriorly, but the anal area may become slightly sclerotised at maturity in Paracoelostoma. The antennae range from well developed in Neocoelostoma (11 segmented) and Paracoelostoma (5 or 6 segmented) to reduced to a group of setae in Cryptokermes and Mimosicerya. Mouthparts are present in Paracoelostoma, but absent in the other genera, although the apparent presence in Paracoelostoma may be due to retention of the preadult mouthparts in the adult cuticle. Legs range from well developed in Neocoelostoma to considerably reduced in Cryptokermes, Mimosicerya hempeli and Paracoelostoma and absent in M. mexicana and M. schraderae. The thoracic spiracles lack atrial pores but each spiracle has a cluster of perispiracular multilocular pores and a well-developed apodeme; the 7 pairs of abdominal spiracles each have atrial multilocular pores. The anal opening is usually apical to slightly dorsal; the anal tube is probably always membranous and, if a sclerotised ring of polygonal wax glands is present at the inner end, this may due to the preadult anal tube being retained within the tube of the adult. Multilocular pores mostly have a circular, oval, triangular or quadrate centre and various numbers of outer loculi. Hairs, hair-like setae, flagellate setae, spiniform setae and spines are present. Cicatrices and disc-like tubercles are absent.
Third-instar (preadult) female. Antennae 8 or 9 segmented in Neocoelostoma and Mimosicerya, 6 segmented in Paracoelostoma and either 3-7 segmented or reduced to a group of setae in Cryptokermes. Legs well developed in Mimosicerya but considerably reduced in size and/or structure in all other genera. Mouthparts always present. Thoracic spiracles lacking atrial pores but with numerous perispiracular multilocular pores; all 7 pairs of abdominal spiracles with atrial pores. Sclerotised anal plate large in Neocoelostoma and Paracoelostoma, but smaller and less complex in Cryptokermes and Mimosicerya. Anal tube with a band of polygonal wax pores at inner end and with a circle of pores on middle or distal part of tube in all genera except apparently Paracoelostoma. Multilocular pores numerous and mostly similar those of adult females. Hairs, hair-like setae, flagellate setae, spiniform setae and spines present. Cicatrices and disc-like tubercles with slightly domed centre present.
Second-instar nymph. Specimens of this instar were available for only two genera, Cryptokermes and Neocoelostoma, and it is not clear whether they are female or male. Antennae 5 segmented in Neocoelostoma and 6 segmented in Cryptokermes. Legs with 5 distinct segments, of normal development in Cryptokermes but leg structure reduced in Neocoelostoma. Thoracic spiracles lack atrial pores but have numerous perispiracular multilocular pores; all 7 pairs of abdominal spiracles with atrial pores. Anal tube with a sclerotised inner end and anal opening located at centre of a sclerotised area. Multilocular pores mostly with a circular, oval, triangular or quadrate centre and various numbers of outer loculi. Hairs, hair-like setae, flagellate setae, spiniform setae present. Cicatrices and disc-like tubercles always present.
First-instar nymph. Antennae 6 segmented with apical segment largest. Legs well developed, with a denticle and a pair of setose digitules on each claw. Thoracic spiracles each with 1-3 perispiracular pores; all 7 pairs of abdominal spiracles lack atrial pores, apart from Paracoelostoma which has one atrial pore in each spiracle. Anal tube well developed, with a band of polygonal pores at inner end; anal opening apical to subapical, surrounded either by spines, spiniform setae and pores (Cryptokermes and Mimosicerya) or by a sclerotised plate bearing setae and pores (Neocoelostoma and Paracoelostoma). Multilocular pores mostly with a circular, oval, triangular or quadrate centre and various numbers of outer loculi. Raised pores present around anal opening in Paracoelostoma.
Derm covered with small hair-like setae, flagellate setae with a large, flat basal collar, and hairs with swollen bases; spines or spiniform setae present on posterior abdomen in Cryptokermes and Mimosicerya. Posterior abdomen with 2 pairs of long caudal setae in Neocoelostoma and Paracoelostoma, 2 pairs of long caudal and 2 pairs of long lateral setae present in Cryptokermes, and 3 pairs of long caudal and 1-6 pairs of long lateral setae found in Mimosicerya species. Cicatrices present on posterior ventral abdomen in an arc of 3, with one medially and 2 laterally in C. brasiliensis, Mimosicerya and Neocoelostoma, and a submedial longitudinal row of 6 cicatrices on each side of abdomen in Paracoelostoma. Disc-like tubercles present in Mimosicerya (toadstool shaped in side view), Cryptokermes and Neocoelostoma (discs with domed centre), but absent in Paracoelostoma.
Adult male. The adult males of two species were available: M. schraderae and N. xerophila. Antenna 10 segmented, each segment without nodes and setae not arranged in whorls. Ocelli dorsal to compound eyes. One or more posterior abdominal segments with paired caudal extensions. Collared setae either absent or with a shallow collar-like basal socket. Bifurcated setae either absent from all legs or present only on profemur. Claws with a pair of setose digitules. Apex of penial sheath divided (bifurcate) and broader anteriorly than posteriorly, and endophallus strongly setiferous.

1.
Females secreting and developing within a test (Fig. 1A Antenna 8-9 segmented; anal opening surrounded by a "secretory unit" formed by a dense ring of elongate wax pores . . . . . .

Descriptions of genera and species
Cryptokermes Hempel Cryptokermes Hempel, 1900: 398. Type species: Cryptokermes brasiliensis Hempel, by monotypy and original designation. Cryptokermes Hempel; Morrison, 1928: 100. This genus was erected by Hempel (1990) for C. brasiliensis collected from a species of Schinus (Anacardiaceae) at Poços de Caldas in the state of Minas Gerais, Brazil. Subsequently, three more species, all from Mexico, were added to the genus: C. mexicanus Morrison collected from Mimosa (Fabaceae) , C. mimosae Foldi also from Mimosa (Foldi, 1995(Foldi, , 2011 and C. oaxaensis Foldi from Prosopis (Fabaceae) (Foldi, 2011). However, after careful examination of published descriptions and specimens of C. mexicanus from the two type localities, we have determined that it is a species of Mimosicerya and this species is discussed further under that genus. Also Cryptokermes mimosae Foldi does not fit the present morphological concept of Cryptokermes or any other named genus and is excluded from this revision pending further study of its relationships, and is not transferred to Cryptokermesini. The type species, C. brasiliensis is redescribed below but, because C. oaxacaensis was described and illustrated recently by Foldi (2011), it is not dealt with again here. In the generic diagnosis below, the features of the second-and first-instar nymphs are based only on C. brasilensis as these stages are not known for C. oaxacaensis. Generic diagnosis. All stages live within a test on twigs or stems of the host plant; the adult female remains within the test to oviposit.
Adult female (based on C. brasiliensis and C. oaxacaensis). Enclosed within a spherical test; mostly in groups on twigs and stems, each group usually with 2-10 individuals. Test hard but brittle, with surface rough, and with an orifice about 1 mm in diameter, in position of anal opening. Body broadly oval, almost circular, with derm membranous apart from anterior, which is strongly sclerotised, mainly on dorsum; this sclerotised zone includes antennae and eyespots and bears spines and a few setae; adjacent unsclerotised derm densely covered by spines, some multilocular pores and few slender setae. Antennae each a flat structure or a small tubercle bearing a group of setae of various sizes and, in C. brasiliensis, also with 2 circular sensilla. Mouthparts absent. Legs of reduced size, segments not discernable. Thoracic spiracles each with numerous perispiracular multilocular pores. Abdominal spiracles numbering 7 pairs, each with atrial pores. Vulvar opening apparently membranous. Anal opening apical to subapical; anal tube membranous, with adult female often retaining anal tube of preadult stage. Dorsal and ventral multilocular pores each with a circular, bilocular or triangular centre. Long hair-like setae present in a transverse row on each segment except at anterior and posterior ends; shorter, thick, spiniform or slender hair-like setae mostly present on abdomen, scattered. Spines present on sclerotised zone of head and thorax, also in a band on adjacent derm. Cicatrices and disc-like tubercles absent.
Third-instar (preadult) female (based on both C. brasiliensis and C. oaxacaensis). Body broadly oval, almost circular, inside a test (as in adult female). Derm membranous, but sclerotised in a circular area around anal opening. Antenna short, indistinctly 3-7 segmented or reduced to a group of setae. Mouthparts present, with labium 3 segmented. Legs reduced, but segments distinct. Thoracic spiracles each with numerous perispiracular pores. Abdominal spiracles numbering 7 pairs, each with a few to numerous atrial pores. Anal opening with a broad, sclerotised margin, opening in a sclerotised area, surrounded by several rings of pores, each with a very wide rim. Anal tube with polygonal wax pores at inner end and a circle of multilocular pores more distally. Dorsal and ventral derm covered in abundant pores and setae of various types, plus spines, cicatrices and disc-like tubercles. Dorsal and ventral multilocular pores present, each with a central loculus either circular, bilocular or trilocular; raised pores present only on dorsum. Various setae present; stout, enlarged conical spines restricted to posterior end, particularly around anal opening. Cicatrices and dorsal disc-like tubercles scattered.
Second-instar nymph (based on C. brasiliensis only). Body elongate-oval, with anal area slightly dorsal and sclerotized, bearing multilocular pores, spines and long flagellate setae. Antenna 6 segmented. Mouthparts present, with labium 3 segmented. Legs fully developed but with short, stout segments. Thoracic spiracles small, each with perispiracular pores. Abdominal spiracles each with several atrial pores. Anal opening apical, at centre of a sclerotised anal plate. Anal tube with a band of polygonal wax pores at inner end and a single circle of pores at mid length. Dorsal and ventral multilocular pores mostly with a circular centre, but sometimes with a triangular or quadrate centre. Dorsal hair-like setae short; flagellate setae and hairs scattered; stout spine-like setae present near coxae, and a few present on posterior abdominal segments. Disc-like tubercles, each with a domed centre, and cicatrices scattered on head and thorax and present in a single transverse row on each abdominal segment.
First-instar nymph (based on C. brasiliensis only). Body elongate-oval. Antenna 6 segmented. Clypeolabral shield and labium well developed. Legs well developed, slender; trochanter with 2 campaniform sensilla on each side; claw long and slender with a single denticle near apex; claw digitules setose. Thoracic spiracles each with a single perispiracular pore. Abdominal spiracles numbering 7 pairs; atria without pores. Anal opening surrounded by a ring of pores, spiniform setae and hair-like setae. Anal tube conspicious, inner end with polygonal wax pores and a circle of pores at opening. Dorsal and ventral multilocular pores, each with 3-6 central loculi and 8-16 outer loculi, in a scattered row among setae on each segment. Short stiff setae on margins and longest hair-like setae in a row across each dorsal and ventral body segment. Enlarged, stout, conical spiniform setae surrounding anal opening. Posterior end of body with 2 pairs of long caudal setae and 2 pairs of long lateral setae. Cicatrices circular, 3 in number, in a medial transverse line on posterior ventral abdomen. Dorsal disc-like tubercles with conical domed centre, present across all segments of body.

Cryptokermes brasiliensis Hempel
Cryptokermes brasiliensis Hempel, 1900: 398. Cryptokermes brasiliensis; Morrison, 1928: 100. According to Ben-Dov (2005), there are supposed to be Hempel type specimens of this species in the Instituto Biologico de São Paulo. One of us (I. Foldi) tried to examine these in November 1985 during a visit to Brazil. Although accompanied by Dr Saulo J. Soria, entomologist at EMBRAPA in Brazil, it was not possible to examine or borrow material from the Hempel collection at Instituto Biologico de São Paulo during their time in São Paulo. In his original description, Hempel (1900) stated that he sent specimens of this insect to T.D.A. Cockerell for his opinion. This explains how material of C. brasiliensis from the type locality came to be deposited in the USNM (see below). We have not designated a lectotype because we have not examined Hempel's type material housed in Brazil.
Adult female (Fig. 2). Unmounted material. Adult female enclosed in a spherical test (Fig. 1A) about 6-8 mm in diameter, dark brown, with a rough external surface and an internal wall covered by a coating of white wax secretion, and with an orifice about 1 mm in diameter in position of female's anal opening.
Venter. Multilocular pores mostly similar to those on dorsum but perhaps slightly smaller, each about 9 µm wide, distributed in transverse bands 4-5 pores wide on anterior abdominal segments but up 10-14 pores wide more posteriorly where pores mixed with (i) hair-like setae, each seta 30-90 µm long ( Third-instar (preadult) female (Fig. 3). Unmounted material. Living within a test, as described above for adult female. Derm of live preadult female light yellow according to Hempel (1900), who mistakenly described the third-instar female as the adult.
Venter. Multilocular pores (Fig. 5P), each 10-11 µm in diameter with a triangular centre and 8 outer loculi, on head and thorax; also pores with either 4 or 5 central loculi and 6 outer loculi or with a quadrate centre and 12 outer loculi ( Cockerell (1902a) erected Mimosicerya as a section of Palaeococcus, not as a section of Icerya as stated by  and Ben-Dov (2014). Icerya (Crypticerya) hempeli was described by Cockerell (1899) based on insects collected by A. Hempel on Mimosa(?), on May 12, 1898, at Campinas, São Paulo state, Brazil. One year later, Hempel (1900) published the name as Crypticerya hempeli and soon after Palaeococcus (Mimosicerya) was established by Cockerell (1902a) as a section of the genus Palaeococcus. Cockerell's designation of its rank corresponds to the nomenclatural requirement of a subgenus (Article 10.4 of ICZN) and is in the form of a genusgroup name with a description and a designated type species, i.e. Icerya (Crypticerya) hempeli Cockerell. Later, Newstead (1920) received from Brazil (São Paulo, on spiny branches of tree or shrub (mimosa?), collected in 1906) scale insects that he compared with a specimen of I. (Crypticerya) hempeli from the collection of Cockerell via E.E. Green and, based on these, erected a new genus Clypeococcus for its reception. The genus name was based on the possession of a supposed clypeus but, in reality, this corresponds to a fold in the derm anterior to the attachment of the labium in the preadult female. Shortly after his publication, Newstead (in correspondence to G.F. Ferris, March 16th, 1920, housed in BME) noted that he had overlooked Cockerell's (1902a) paper and regretted that he had "added another useless synonym".  revised and diagnosed the taxon, changed the status to Mimosicerya, and formally recognised Clypeococcus as its synonym.
Mimosicerya remained a monotypic genus for over 100 years, until Foldi (2009) added a new species from Venezuela, M. williamsi Foldi, with descriptions of the adult and preadult females. Here we add two more species: Nautococcus schraderae Vayssière, which becomes a species of Mimosicerya by synonymy of Nautococcus with Mimosicerya (see under M. schraderae below), and Cryptokermes mexicanus Morrison, which we transfer to Mimosicerya (see under M. mexicana below). We redescribe and illustrate all species except M. williamsi, which was recently treated in detail and compared with M. hempeli by Foldi (2009).
Generic diagnosis. All stages live either exposed, or hidden in cracks or under bark on twigs or stems of the host plant; the adult female is pupillarial, remaining inside the sclerotised preadult exuviae for reproduction.
Adult female. Body broadly oval, female remaining within preadult exuviae after last moult. Derm membranous except for an approximately circular sclerotised area anteriorly bearing antennae, eyespots, multilocular pores, setae and numerous spines of varying stoutness and length. Ventral dermal pocket-like depressions (apodemes) present, 2 per segment in a submedian line on abdomen (M. hempeli) or absent (M. mexicana, M. schraderae and M. williamsi). Antenna greatly reduced, platelike or bulbous, segments not discernable on slide-mounted specimens, but bearing a group of sensory setae. Mouthparts absent. Legs either absent (M. schraderae and M. mexicana), present but strongly reduced (M. hempeli) or represented by vestigial protuberances (M. williamsi). Thoracic spiracles wide, each with a group of perispiracular multilocular pores. Abdominal spiracles numbering 7 pairs, with posteriormost pair close to anal opening; peritremes each situated on a truncate sclerotized protrusion in M. hempeli but not in M. mexicana, M. schraderae or M. williamsi; each spiracle with or without atrial pores. Anal tube of preadult retained. Multilocular pores circular or broadly oval with a thickened rim; each with a circular or oval, rarely triangular or quadrate, centre and with 10-16 outer loculi, distributed throughout both dorsum and venter, most abundant toward posterior end; raised pores, if present, only on sclerotised head area. Spines present on sclerotised anterior area and sometimes spines or spiniform setae present elsewhere, particularly on posterior end of body. Flagellate and hair-like setae sparsely scattered over body. Hairs, if present, sometimes with a swollen apex (M. hempeli) and most common on posterior end of abdomen. Cicatrices and disc-like tubercles absent.
Third-instar (preadult) female. Body globular to nearly spherical in life. Derm thick and sclerotised dorsally and marginally in mature specimens, often membranous only on mid-ventral area. Dorsal posterior end of abdomen of mature specimens sclerotised and bearing a wide circular area of derm, surrounded by a wide rim in M. hempeli; in M. schraderae, this circular area of derm is known to form an operculum that lifts, allowing abdomen of adult female to protrude (Fig. 14). Antenna 7 (M. mexicana) or 9 segmented (M. hempeli, M. schraderae and M. williamsi); each segment with a few fleshy setae as well as slender hair-like setae. Eyespot situated lateral to scape. Mouthparts present; labium small. Legs well developed, trochanter with a long ventral seta and with 2 campaniform sensilla on each side; claw stout, without a denticle but with a pair of setose digitules. Thoracic spiracles large, each with perispiracular multilocular pores. Abdominal spiracles small, numbering 7 pairs; all atria with pores. Anal opening apical and anal tube with polygonal wax pores at inner end, also with 2 or 3 rows of multilocular pores nearer to anal opening. Multilocular pores with either a triangular, quadrate or circular centre, distributed throughout on dorsum and margin; mid-ventral area with a few multilocular pores distributed in transverse rows. Flexible cylindrical long hairs, each with an expanded base and mostly with a swollen apex, either densely covering sclerotised margins of mid-ventral area and thorax (M. hempeli), or near mouthparts only (M. schraderae), or densely covering medial to submedial thorax (M. williamsi), or rare (M. mexicana). Short stout setae, slender setae and spinose setae sparsely scattered on dorsum. Spines absent (M. hempeli) or present (M. mexicana, M. schraderae and M. williamsi). Ventral cicatrices, varying in size, numerous, usually scattered on thorax and in transverse segmental rows on abdomen; sometimes also on dorsum. Disc-like tubercles present in all species, with various distributions.
First-instar nymph. Body elongate, elliptical. Antenna 6 segmented, third segment very short in M. hempeli and M. schraderae, pedicel and apical segment longest; apical segment swollen and bearing fleshy setae at apex. Clypeolabral shield well developed; labium 3 segmented, apical segment with sensory setae. Legs well developed; claw elongate with a pair of setose digitules. Thoracic spiracles each with 1 or 2 perispiracular pores. Abdominal spiracles numbering 7 pairs, small, without atrial or perispiracular pores. Anal opening apical to subapical; anal tube short, inner end with a double collar of polygonal wax pores. Dorsal and ventral pores small, mostly circular, each with 3-6 inner loculi and a variable number of outer loculi. Spines stout, cylindrical, pointed at apex, distributed on posterior dorsum, densest around anal area in all species. Slender hair-like setae with a stout conical base, sparse across abdominal segments; hairs and flagellate setae sparsely distributed. Long caudal setae in 3 pairs on posterior abdominal segment, plus 3-6 pairs of long setae on margin. Cicatrices flat and circular, 3 in number, in a transverse row ventromedially on posterior abdomen. Disc-like tubercles, each toadstool shaped in side view with a slightly domed centre, numerous in transverse rows on dorsum and either numerous towards posterior end (M. hempeli) or absent at posterior end (M. schraderae). This species was described by Cockerell (1899) based on what he believed to be the adult female. However, he described 9-segmented antennae and small stout legs, which are characteristic of the third-instar (preadult) female. The collection data for the type material was given as "Campinas, Brazil, May 12, 1898, on Mimosa (?). (A. Hempel, 215d)". Later, Newstead (1920) also described and illustrated the preadult female, believing it to be the adult and described and illustrated the first-instar nymph. Later still,  recognised that the preadult exuviae concealed the pharate adult female and described the adult and preadult females and the first-instar nymph. Males are not known.
Note: It is clear that the above four boxes in the USNM contain Cockerell's type material, from which specimens probably later were slide-mounted for  monograph. Morrison provided detailed illustrated descriptions of the adult and preadult females and the first-instar nymph that would have required slidemounted specimens. It seems that the labelling of the three USNM slides of type specimens with the collection date of 1900 was just an error on the labels, and we believe that these females are part of the type material. There is one slide of Cockerell's type material in the Natural History Museum in London (Williams, 1985), but it consists of pieces of cuticle only, and although the collection data show that the specimen is from the type material, the slide is not labelled as being a type. We have not designated a lectotype because there is no doubt as to the identity of this species; also the one box that has a label ("TYPE Ckll. Coll.") indicating that it is primary type (rather than "CO-TYPE") contains a single dry mature preadult female that would be difficult to slide-mount. Thus all original specimens are listed as syntypes.
Clypeococcus hempeli Newstead: BRAZIL: Syntypes: 1 slide with part of 1 preadult female, 1 slide with 3 first-instar nymphs and one box with 1 dry preadult female, all labelled as "Clypeococcus hempeli", São Paulo, Brazil, on spiny branches of unknown tree, 1906, purchased from O. E. Janson (BME). Note: These specimens were sent to G.F. Ferris by Newstead (as noted in Newstead's correspondence to G.F. Ferris, March 16th, 1920) and were part of Newstead's type series. The Natural History Museum, London, has dry specimens of this Newstead species but no slide preparations could be located.
Adult female (Fig. 6). Unmounted material. Protected inside sclerotised cuticle of preadult female and reproducing therein. For slide preparation, adult females were extracted from preadult exuviae.
Third-instar (preadult) female (Figs 1B, 7). Unmounted material. Living exposed on bark; body subglobose, dark slate-gray with a thin but dense coating of cream-coloured mealy secretion; subdorsal areas marked by a longitudinal series of small round spots free from secretion; legs blackish brown to dark brown (Cockerell, 1899). Young preadults reddish-brown but becoming darker with age, with derm strongly sclerotised in old specimens.
Venter. Medial to submedial area of abdomen membranous and clear, delimited by a dense band of hairs (Fig.  7N) and sclerotised submarginal derm. This clear area with cicatrices (Fig. 7I) of various sizes, each 15-30 µm wide in an irregular transverse row 1-2 cicatrices wide on each abdominal segment, even extending a short distance onto sclerotised venter; also with a few multilocular pores, each about 10 µm wide and mainly with 3 or 4 (rarely 6) loculi at centre and 10-12 outer loculi (Figs 7K, L). Small, circular pores (Figs 7R, S), each 6-7 µm in diameter with 3 or 4 loculi, sparsely scattered on venter. Long, flexible, cylindrical hairs (Fig. 7N) each mainly 70-110 µm long but with a few only 40-50 µm long, and with an expanded base and a swollen apex, densely distributed on weakly sclerotised ventral margins of abdomen, also sparse on membranous ventral area and on head and thorax, where mixed with sparsely scattered short flagellate setae (Fig. 7O), each 20-30 µm long. Sclerotised margins and submargins with disc-like tubercles with a central opening (Fig. 7J) plus a few smaller, domed disclike tubercles.
Venter. Multilocular pores (Figs 8H, N), each 7-8 µm in diameter with a triangular centre and with 3 paired (Fig. 8H) or 3 single (Fig. 8N) tiny outer loculi, but pores with 3 paired outer loculi sparse; present in longitudinal lines on abdominal segments. Hair-like setae, each 15-20 µm long, scattered on head and thorax, and in transverse rows 1 seta wide, across all abdominal segments; longest hair-like setae (Fig. 8O), each 50 µm long, on head, thorax and in medial area of each abdominal segment. Hairs (Fig. 8M) scattered on abdomen. Caudal setae in 3 pairs on posterior abdominal segment, each seta 360-430 µm long (Fig. 8R), plus 3 pairs of lateral setae on margin of abdomen, 360 µm to 140 µm long, becoming progressively shorter anteriorly. Cicatrices circular, 3 in a transverse line medially on posterior abdomen, middle one small, about 11 µm in diameter, lateral ones each 21-23 µm in diameter. Disc-like tubercles absent.  (Cockerell). Adult female. A. plate-like antenna with sensory setae; B. dorsal and ventral stout, straight and curved spines within sclerotised area; C. multilocular pore at centre of raised, circular, sclerotised derm; D. slender hair-like setae; E. common multilocular pore with circular centre and 15 outer loculi; F. abdominal spiracle with protruding peritreme; G. short hair-like seta; H. tube anal with polygonal wax glands at inner end; I. short or long flexible hair with swollen apex; J. ventral tubular structure near posterior end; K. metathoracic leg without discernable segments; L. straight and curved spines around spiracles; M. thoracic spiracle with perispiracular multilocular pores.  (Cockerell). Third-instar female (preadult). A. apical segments of antenna; B. multilocular pore with quinquelocular centre; C. short hair-like seta; D. multilocular pore with 6-locular centre; E. view from above of abdominal spiracle located within thick derm, a=atrium, p= peritreme; F. anal tube; G. disc-like tubercle; H. quinquelocular pore at center of raised sclerotised derm; I. cicatrices of variable sizes; J. disc-like tubercle; K. multilocular pore with triangular centre; L. multilocular pore with quadrate centre; M. tibia, tarsus and claw of metathoracic leg; N. long, flexible, cylindrical hair with swollen apex; O. flagellate seta; P. thoracic spiracle with perispiracular multilocular pores; R. simple quadrilocular pore; S. simple trilocular pore.  (Cockerell). First-instar nymph. A. disc-like tubercle; B. apical segment of antenna; C. short hair-like setae; D. quinquelocular pore; E. pore with rectangular quadrilocular centre and 4 outer loculi; F. straight or curved spines; G. tube anal with polygonal wax pores at inner end; H. pore with large triangular centre and 3 tiny paired outer loculi; I. abdominal spiracle; J. pore with 6 loculi; K. claw with a denticle and digitules; L. metathoracic tarsus and claw, proximal part of tarsus with a campaniform sensillum (cs); M. flagellate seta; N. pore with triangular centre and 3 tiny outer loculi; O. ventromedial long hair-like seta; P. thoracic spiracle with one perispiracular multilocular pore; R. 3 pairs of long caudal setae. There have been several descriptions of this species. Cockerell (1902b) and Ferris (1918) both erroneously referred to the species as Cryptokermes brasiliensis. Later,  named the species as Cryptokermes mexicanus and provided the details of the type specimens (see below).  pointed out that Ferris' (1918) and Cockerell's (1902) descriptions were actually of C. mexicanus. There is no mention of a test in any of the above four papers, and Ferris (1918, page 221) says of the adult female: "remaining enclosed within the derm of the penultimate stage", which is a characteristic of the genus Mimosicerya. Also Morrison (1927, page 103) describes the first-instar nymph as having curious derm discs that are flattened or with the centre slightly convex. We examined these structures under the compound microscope and they are toadstool shaped in side view and exactly match the disc-like tubercles of the first-instar nymph of M. hempeli, rather than the disc-like tubercles of the firstinstar nymph of C. brasiliensis. Based on the morphological features of the first-instar nymph and the preadult and adult females, we have confirmed that C. mexicanus belongs to Mimosicerya and here transfer it to that genus as M. mexicana. As the genus name Mimosicerya (based on Icerya Signoret) is feminine and the species name is an adjective (Pellizzari & Williams, 2013), the latter should be amended to "mexicana". We describe the adult female, preadult female and first-instar nymph, but illustrate only the latter because that instar is the best for species recognition.
Morrison (1927, page 103) listed two type collections, as follows: "Zapotlan, Jalisco, Mexico, on Mimosa sp., 1903, collected by C.H.T. Townsend (T. & B. Cy. #22) (holotype and paratypes); and from Cuautla, Morelos, Mexico, on Mimosa sp., July, 1897, collected by A. Koebele (#1609 -Div. Ent. #7894 -and #1672 -Div. Ent. #7918) (paratypes)". The types are in the USNM, as stated by , but the BME also holds slidemounted specimens with the above data and these were apparently mounted by G.F. Ferris; the BME also has one collection of dry material from the Zapotlan locality. Cockerell sent some of the Zapotlan specimens to Ferris, as stated by Ferris (1918), but it is not known how the Cuautla specimens were obtained by Ferris. However, given the collection data, both BME collections must have come from the original collections made by Townsend and Koebele, but were not the specimens seen by  when he described C. mexicanus, and thus they are not part of the type series. Each of Ferris' slides of specimens from Zapotlan is labelled as "heautotype; this name was applied to a specimen used by the original describer as an illustration of the species and compared with the type or cotype. The USNM has 15 slides from the Zapotlan series from Townsend and 23 slides from the Cuautla series from Koebele. The slide in the USNM marked as "holotype" has label data: "Cryptokermes / brasiliensis Hemp / mexicanus / On Mimosa sp. (wild) / Zapotlan, Jalisco, /Mexico / T. & B. Cy. #22 / Rec'd. July 1903" with the word "holotype" scratched on the glass of the slide. We have not examined Morrison's type specimens in the USNM.
Adult female. Mounted specimen (n=1). Body broadly oval, 5.2 mm long, 5.0 mm wide. Derm membranous throughout except for a wide circular sclerotised area on anterior end of body and a smaller slightly sclerotised area surrounding anal opening on posterior end of body. Antennae not seen. Eyespots appear as 2 light spots within sclerotised area. Mouthparts absent. Legs absent. Thoracic spiracles located on border of sclerotised area; each peritreme about 80 µm wide with a group of perispiracular pores and a large apodeme about 285 µm long. Abdominal spiracles numbering 7 pairs; each spiracle with a peritreme 35-45 µm wide, an internal sclerotised atrium and an external membranous atrium with 4-8 atrial pores; posteriormost pair of spiracles much reduced in size, about 30 µm long and, 22 µm wide, without atrial pores. Anal tube retained from preadult. Multilocular pores scattered, each 10 µm wide with a wide rim; most widespread type with a wide single circular loculus at centre and about 16 loculi in outer rim; these pores more numerous toward posterior end of body; some multilocular pores apparently with an oval centre containing 2 small loculi, sparsely distributed. Short spines sparsely scattered throughout; stouter and longer spines, each 50-55 µm long, densely distributed within sclerotised area on anterior end of body. Hair-like setae and flagellate setae both short and slender, each 25-35 µm long, sparsely scattered throughout. Longest flagellate setae, each 120-130 µm long with a wide, enlarged and sclerotised collar, densely distributed around vulva. Cicatrices and disc-like tubercles absent.
Dorsum. Multilocular pores, each about 10 µm wide with either a circular or a triangular centre and 12-16 outer loculi, distributed in an irregular transverse row or band on each body segment. Smaller quadrilocular pores sparsely scattered. Stout, enlarged conical spines each 20-25 µm long, scattered on abdomen, most abundant posteriorly. Hair-like setae with conical basal collars, of various sizes: short and thin or thick, each 15-25 µm long, some setae with enlarged apex; longest and thickest setae 45-55 µm long, scattered on all segments. Cicatrices numerous, each 12-30 µm in diameter, in a transverse row on each body segment, 1 row sometimes 2 cicatrices wide. Disc-like tubercles, each 7-8 µm in diameter, with a thick rim and domed slightly in centre, densely distributed on body, scattered on head and thorax, and present in transverse segmental rows, 3-5 tubercles wide, on abdomen.
Venter. Multilocular pores each about 10 µm wide, with circular, oval or triangular centre, scattered. Stout conical spines as on dorsum, scattered across abdomen, most abundant on posterior end. Hair-like setae, as on dorsum, each 45-55 µm long, most numerous on medial and submedial area. Hairs with expanded bases rare; short flagellate setae present throughout. Cicatrices, each 12-30 µm in diameter, present in a band 1 or 2 cicatrices wide across each abdominal segment. Disc-like tubercles, each about 8 µm in diameter and domed in centre, present throughout, densest across abdominal segments.
Venter. Multilocular pores (Fig. 9E), each about 10 µm wide, similar to those on dorsum, mostly with a quinquelocular centre and about 14 outer loculi, densest on posterior abdominal segments; multilocular pores ( Fig.  9J) with a triangular centre and about 12 outer loculi, sparse on thorax. Hair-like setae short, each 12-30 µm long, sparsely scattered on head and thorax, and with about 8-10 setae in a transverse row across each abdominal segment. One long seta of 75-80 µm, present near lateral-most cicatrix on each side of body. Caudal setae (Fig. 9L) in 3 pairs, each seta about 450-500 µm long, on posterior end of abdomen, and another long seta laterally on each side of either sixth or seventh abdominal segment. Cicatrices circular, 3 in number, each 8-10 µm in diameter, situated medially on posterior abdomen. Disc-like tubercles present on margin only.

Mimosicerya schraderae (Vayssière) comb. n.
Nautococcus schraderae Vayssière, 1939: 124;Vayssière & Hughes-Schrader, 1948: 57. The original description by Vayssière (1939) was based on the preadult female that he had assumed was the adult female. Later Vayssière & Hughes-Schrader (1948) described the adult female and all other instars, and provided a plate of photographs and information on biology. The species was described from specimens collected in February 1937 by S. Hughes-Schrader on Barro Colorado Island in Panama, Central America, mostly from Annona spraguei (Annonaceae).
There are 56 slides (museum numbers: 6604/ 1-56) in the MHMN and each is labelled as type of N. schraderi [sic]; most are in very poor condition and cuticular structures are difficult to see in many cases. Here we redescribe and illustrate all available instars. We synonymise the genus Nautococcus with Mimosicerya based on morphology of male and female instars, and consequently N. schraderae becomes M. schraderae (Vayssière). This species is distinct morphologically from all others in Mimosicerya.
Adult female (Fig. 10). Unmounted material. Adult female enclosed within thick, strongly sclerotised derm of preadult (Fig. 1C), where it oviposits. Individuals or aggregations of the scale insects are frequently covered by a protective shelter constructed by ants.
Third-instar (preadult) female (Figs 1C, 11). Unmounted material. Body broadly ovoid, 9.0-11.0 mm long, 7-9 mm wide and 5-6 mm high; reddish-brown with dorsal and marginal derm strongly sclerotised, mid-venter of abdomen more or less membranous. Distribution of dorsal secretions characteristic of species: dorsum and dorsal margin with white secretions distributed in regular transverse ridges on body segments, also forming 5 longitudinal lines, one median, 2 submedial and 2 submarginal to marginal; each longitudinal line separated by a space covered by a light coating of secretion (Fig. 1C). Posterior end of abdomen dorsally with a circular area, 1.7-2.3 mm in diameter, with derm covered by pores and setae, forming a plate that later is pushed out allowing adult female to protrude her abdomen (Fig. 14). Venter with powdery secretions; antennae and legs brownish-black. An external waxy tube, about 4-5 mm long, produced from anus.
Dorsum. Multilocular pores with a triangular centre and about 16 outer loculi (Fig. 11B), scattered on dorsum and around each peritreme. Similar multilocular pores but with a circular centre, mainly present on posterior end of abdomen (Fig. 11G). Clusters of spines each forming a "secreting area" (Fig. 11D), with (i) hair-like setae, (ii) spines and (iii) flagellate seta, in discrete zones on each segment of body; each larger cluster containing about 100-150 mainly curved and pointed spines, mostly 30-35 µm long, but smaller clusters with about 20-40 spines; periphery of each cluster with hair-like and flagellate setae. These spines also scattered throughout dorsum but mostly broken, with only base visible and appearing as pore-like structures. Long spines most numerous, each straight with a pointed apex, about 70-90 µm long, densely distributed throughout on dorsum, mixed with less numerous and shorter (about 30-35 µm long) stout spines (Fig. 11L). Details of cicatrices difficult to discern due to dermal sclerotisation but probably similar to those on venter (Fig. 11J); larger cicatrix-like structures, each 40-70 µm across, with a rough surface, appear as clear windows in sclerotised derm [but these may be artifacts]. Disc-like tubercles not apparent.
Venter. Multilocular pores (Fig. 11G), each with a circular centre and with 10-11 outer loculi as on dorsum, scattered throughout. Hairs (Fig. 11M), each 90-100 µm long, with a swollen apex and an expanded base, present in medial area of thorax. Spines, both straight and curved (Fig. 11L), of two sizes, either 30-35 µm or about 70-90 µm long, as on dorsum, dense across body in transverse bands but not forming wide patches. Cicatrices difficult to discern; circular structures with a smooth surface (Fig. 11J), each 20-30 µm across, and larger cicatrix-like structures with a rough surface (Fig. 11I), scattered. Disc-like tubercles also difficult to discern but probably sparsely present (Fig. 11H).
Dorsum. Multilocular pores (Figs 12A, J), each with 3-6 central loculi and 10-12 outer loculi, in transverse rows, 1 pore wide, on all body segments. Short flagellate and hair-like setae (Figs 12D, E), each 25-40 µm long, scattered on head, and in transverse rows on thorax and abdomen; a few longer setae, each 50-55 µm long, on head and thorax. Stout straight spines (Fig. 12H) present in a sparse transverse row on each segment of abdomen, dense on last abdominal segments where mixed with stout curved spines (Fig. 12H). Dense groupings of robust setae (Fig. 12F), each 80-95 µm long, present around anal opening. Disc-like tubercles (Fig. 12C) dense on marginal to submarginal areas around body, and forming submedial clusters on each side of head; also present in a dense transverse row, 1-3 tubercles wide, on each body segment, apart from last 4 segments where spines replace tubercles.
Adult male (Fig. 13). Mounted specimens (n=2). Body large, 4.0-5.0 mm long, 1.30-1.33 mm wide across prealare. Body covered in many setae of 3 main types: (i) collared setae (cs) (Fig. 13E), each with a shallow collarlike basal socket but on a rounded mound, up to perhaps 100 μm long; (ii) hair-like setae (hs) (Fig. 13F), each with a shallow basal socket around setal membrane and up to 50 μm long, and (iii) hairs (hrs) (Fig. 13G), each with a cone-like base fused to seta and up to about 40 μm long. Loculate pores (lp) (Fig. 13C) abundant, each about 10 μm wide, with a star-like pattern with mainly 4 loculi; found almost wherever setae are present. Other pores of 3 types also present: (i) convex pores (cp) (Fig. 13A), each very convex, each 5 μm tall and 5 μm wide, on a shallow base, with a granulate surface, restricted to head; (ii) small heavily sclerotised pores (msp) (Fig. 13B), each about 3.5 μm wide, present on head but possibly absent elsewhere, and (iii) barely sclerotised pores (mp) with a dark centre, each about 3 μm wide, on thorax and abdomen. Antenna long, with long setae randomly distributed, each usually with satellite setae. Nodulations not noted on any sclerites. Scutum with a median membranous area with setae and pores. Legs well developed and setose, with many spur-like setae on tibia and tarsus, none bifurcate; tarsus 2 segmented (Fig. 13L); claw without a denticle; claw digitules setose. Abdominal segment I not visible ventrally; abdomen with a short, broad lateral caudal extension (Fig. 13I) on segment VIII, other abdominal segments somewhat bulbous marginally; abdomen without tubular ducts.
Head. Triangular in dorsal view, length about 625 μm, width across compound eyes 750-800 μm. Dorsally with a well-developed postoccipital suture extending across posterior part of epicranium; with a broad postocciput posteriorly, without setae. Dorsomedial part of epicranium apparently entirely membranous, without a dorsal part to midcranial ridge. Dorsal surface with many setae, mainly cs but with some hs and hrs (cs up to about 80-90 μm long; hs up to 50 μm long); and with many pores, both lp, cp (fewer than lp) and a few msp. Laterally with a pair of large compound eyes (cde), each about 270-365 μm long, with about 120 ommatidia. Each compound eye with a narrow, lightly sclerotised, ocular sclerite along dorsal and posterior margins, and a single ocellus situated dorsal to compound eye; width of each ocellus about 60-65 μm; each ocular sclerite with a narrow post-ocular ridge along dorsal margin; ventral projection not located. Ventrally with a strongly sclerotised series of ridges forming a fivearmed cross (Fig. 13O), composed of: (i) ventral midcranial ridge anteriorly; (ii) a pair of lateral preocular ridges, and (iii) a pair of preoral ridges (pror) posteriorly. Ventral part of epicranium membranous apart from some light sclerotisation in angle between vmcr and procr and between procr and pror; setae, lp and cp abundant anteriorly between ventral midcranial and preocular ridges but absent from between preocular and preoral ridges; area posterior to preoral ridge with hs and hrs setae and lp around mouth. Cranial apophysis shallow and broad. No structures representing tentorial arms, tentorial bridge or tendon-like apodeme detected. Ventral sclerites present just posterior to each compound eye. Antenna: 10 segmented; length about 3.8 mm (ratio of total-body length to antennal length 1:0.84). Scape about 190 μm long, 195 μm wide, sclerotised, basal part with a sclerotised marginal ridge and with a strong basal articular process extending posteriorly from lateral margin of each scape; with about 25 setae, each about 65-80 μm long. Pedicel 215-250 μm long, 125-130 μm wide, articulating with scape; with about 45 setae, of which about half very long, each about 270-330 μm long; others short as on scape, some with satellite setae, plus a large campaniform sensillum on dorsal surface. Segments III-IX each more or less parallelsided and about 70-105 μm wide, broadest at segment III, narrowing towards apex; lengths (μm): III  segments each with about 50-65 long setae, rather randomly distributed, plus a few hs; each long seta 185-350 μm long, with 0-3 satellite setae (Fig. 13N); antennal bristles barely differentiated from long setae, with 3+ on apical segment; possibly present on some other segments but not differentiable from other setae; no basiconic sensilla or capitate setae noted on segment X.
Thorax. Prothorax: neck broad, with no indication of a cervical groove. Dorsally: pronotum absent. With a pair of diagonal post-tergites, each about 415 μm long, broadest anteriorly. Laterally with a pair of strong cervical sclerites that articulate anteriorly with ventral sclerite and preoral ridge; cervical sclerites perhaps without a proepimeron. Pleural ridge short, extending dorsally from articulation with coxa; pleural apophysis distinct. Ventrally: prosternum with a well-sclerotised median ridge, about 500 μm long, which broadens posteriorly but with no obvious sternal apophysis. Most membranous areas covered with abundant hs (about 60 μm long), a few cs, hrs and lp, as follows: with a broad group of median pronotal setae and pores extending between post-tergites over anterior end of prescutum; a large group of lateral pronotal setae + lp anteriorly; large groups of propleural setae and pores anteriorly and posteriorly on each side, possibly without any gap between groups; anteprosternal setae either absent or in a group fused with prosternal setae; with an elongate group of prosternal setae and pores on each side of prosternum; presence of antemesospiracular setae and lp uncertain. Mesothorax: dorsally: prescutum (prsc) large and oval (length 445-500 μm, width 685-780 μm); mesoprephragma short and narrow; prescutum not nodulated; prescutal ridges short; prescutal sutures well developed; prescutum without prescutal setae and lp. Scutum (sct) without nodulations; with a large oval membranous area medially, 160-260 μm long, 455-495 μm wide, with many scutal setae (cs, each about 40-60 μm long; hs and hrs each about 25-35 μm long) and lp; and with many setae and lp on either side of scutellum and on either side of membranous area. Scutellum (scl) triangular; scutoscutellar sutures extending posterolaterally from scutal membranous area to postalare; length 380 μm, width 700-715 μm; each outer angle with a distinct oval membranous area; with about 15 scutellar setae but no lp. Membranous area immediately posterior to scutellum without setae or pores. Laterally: prealare (pra) elongate. Tegula (teg) with many tegular setae and a few lp. Mesopleural ridge well developed, with a deep pleural apophysis. Episternum not nodulated. Ventrally: basisternum large, length 700-760 μm, width 775-825 μm; with a poorly developed median ridge; without lp but with a line of setae medially on either side of median ridge; anteriorly without a marginal ridge but marginal ridge well-developed anterolaterally along border of lateropleurite; precoxal ridges well-developed, each with a short subepisternal ridge; furca (f) moderately narrow posteriorly, waisted, with quite long, stout arms, which diverge strongly; lateropleurite represented by a small narrow sclerotised area. Postmesospiracular setae and lp abundant laterally and posteriorly to mesothoracic spiracle, but with only setae medially. Mesothoracic spiracles large, width of each peritreme perhaps 180-220 μm. Wing sclerites showing nothing distinctive. Metathorax: dorsally: metapostnotum present as a pair of large, roughly triangular, sclerites, which nearly fuse with tergite of abdominal segment I; metatergal setae (mts) represented by a band of longish setae + a few lp extending across median part of segment. Laterally: dorsospiracular setae and lp in a large group. Suspensorial sclerites present. Pleural ridge well developed, without a vestigial metapleural wing process extending anterodorsally from dorsal end; precoxal ridge well developed and extending about 255 μm medioventrally; metepisternum without setae; metepimeron represented by a strong sclerotisation extending posterodorsally around metacoxae; without setae or lp. Ventrally: with a large group of antemetaspiracular setae + lp; metasternum transverse, only lightly sclerotised but with large lateral apophyses. With a large group of postmesoprecoxal ridge setae and lp on each side; anterior and posterior metasternal setae and lp abundant. Postmetaspiracular setae abundant, with many lp. Posterior spiracles large, located below mesocoxae; width of each peritreme about 215 μm.
Abdomen. With a rather bulbous, rounded caudal extension (bce; Fig. 13I) on abdominal segment VIII; all other abdominal segments also slightly rounded marginally. Narrow tergites (dat) present mediolaterally on anterior borders of all segments; sternites absent. Setal distribution not easy to see; dorsal abdominal setae abundant, in bands across segments, probably mostly hs, shorter than ventral setae, each 45-60 μm long; ventral abdominal setae also abundant but in broader bands than on dorsum; hs on ventral surface somewhat longer than on dorsum (mostly about 60-85 μm long); hs ventrally about twice as frequent as hrs. Loculate pores frequent to abundant on both dorsum and venter. Minute unsclerotised pores occasional, at least pleurally; convex pores and small sclerotised pores not detected. Pleural setae and lp not apparently divided into dorsal and ventral pleural groups but indistinctly separated from dorsal (das) and ventral (vas) abdominal setae. Pleural setae mainly fairly short but some up to 125 μm long. Abdominal spiracles (Fig. 13H) very difficult to locate but thought to be small and simple, with a sclerotised opening about 12 μm wide; believed to be present laterodorsally on anterior margins of at least segments V-VII.    Genital segment. Anus present on dorsal surface above penial sheath, margin sclerotised and about 50-60 μm wide, with a slightly sclerotised triangular area extending anteriorly. Penial sheath sclerotised, about 750-790 μm long and 240-270 μm wide posteriorly, broadest anteriorly, where 305-310 μm wide; penial sheath blunt and bifurcated posteriorly, with sparse short setae across dorsal surface, along margins and ventrally on either side of ventral opening, and with a group of small sensoria near apex. Aedeagus (Fig. 13K) possibly mainly membranous, length unknown, in a groove along ventral margin of penial sheath; with a very long, strongly setiferous, eversible endophallus (Fig. 13J).
Biological observations (Fig. 14). The following is a summary and translation from French of pages 66-73 from Vayssière & Hughes-Schrader (1948).
Research on the life cycle was conducted under laboratory and field conditions, during the years 1937 to 1941. Nautococcus schraderae was collected on the island of Barro Colorado in the Panama Canal Zone during the dry periods of February 1937, December-January 1937-38, 1939-40, 1940; the preferred host plant was Annona spraguei (Annonaceae). It was collected also at Turrialba in Costa Rica, in February to March 1944, on Inga tonduzii (Fabaceae). In Panama, the preadult females were found on the trunk and the largest branches, mostly fixed in cracks or under bark. Frequently the second-instar male nymphs formed little groups of 4 to 20 individuals under bark or under the protective shelter constructed by ants. Third-instar males produced a white filamentous cocoon in which they developed to the adult stage.
Each adult female is enclosed within the strongly sclerotised preadult exuviae (Figs 1C, 14A), which frequently is covered by a protective shelter constructed by ants; an association with ants is frequently observed with this species. The posterior end of the dorsal abdomen of the preadult has a large area of circular derm surrounded by a sclerotised rim; at maturity, this derm is strongly sclerotised and transformed into a hard plate (operculum) that is pushed out by the adult female, through which she very slowly pushes out her posterior abdomen; this process can take some hours. Sometimes only the posterior end of the abdomen, but frequently about a third of the abdomen, extends from the preadult exuviae, and is directed upwards and anteriorly (Fig. 14B) and, in the laboratory, the attraction of males started immediately. The duration of mating varied considerably, between 8 to 126 minutes, with an average of 34 minutes, and males mated once, rarely twice, whereas the female could be mated successively by two different males. However, after mating, the female retracts its abdomen and stays within the preadult exuviae. Oviposition started from the seventh day after mating, was well advanced around days 12-13 and apparently finished by about day 14. The eggs are laid within a network of filamentous wax secretions between the body of the female and the exuvial wall of the preadult. The period from oviposition to eclosion varied widely (48 and 66 days); the first-instar nymphs stay immobile inside the preadult exuviae for about two months after which they move out and settle on the trees. The most important cause of mortality of the females, nymphs and eggs was due to predators, particularly the larvae of coccinellids. Observations in the field and the laboratory suggest that there are two distinct, not overlapping, generations per year. This life cycle appears similar to those of Llaveia Signoret and Llaveiella Morrison, which also have two distinct generations per year.
Vayssière & Hughes-Schrader concluded that the placement of Nautococcus based on morphology (e.g., absence of legs and mouthparts in the adult female) was in the Coelostomidiinae (now family Coelostomidiidae), but, based on cytological results (2n female=6, male=5), it belonged to the Monophlebinae (now Monophlebidae). Neocoelostoma Hempel, 1932: 310. Type species: Neocoelostoma xerophila Hempel, by monotypy and original designation. This genus was introduced by Hempel (1932) for Neocoelostoma xerophila Hempel, which he assigned to the Coelostomidiinae (now Coelostomidiidae) in the family Margarodidae sensu . The species was described from a collection on Piptadenia falcata [now Anadenanthera peregrina var. falcata] (Fabaceae) obtained in 1931 in Pirapitinguy, São Paulo state, Brazil. Hempel's short description was based on the preadult female and the first-instar nymph and lacked any illustrations. The species was reported subsequently from Argentina, Bolivia and Paraguay on various Acacia species and on Parkinsonia praecox (Lizer y Trelles, 1936Trelles, , 1939, and it has been recorded as the host of several chalcidoid wasp parasitoids (Noyes, 2012). Neocoelostoma remains monotypic, perhaps due to the difficulties of comparing newly collected material with the type material.

Neocoelostoma Hempel
The life cycle of females of Neocoelostoma differs from that of Cryptokermes, Mimosicerya and Paracoelostoma in that the adult female is not retained within the test or exuviae of the preadult, as occurs in the other genera Vayssière & Hughes-Schrader, 1948). In Neocoelostoma, the adult female escapes from the exuviae of the third-instar female via a circular opening at the anal end and then seeks a sheltered position on the bark where it exudes a mass of cottony white filaments prior to oviposition.
Generic diagnosis. This monotypic test-forming genus is restricted to Fabaceae. Most stages live exposed on the trunk or branches. It occurs in Argentina, Bolivia, Brazil, Paraguay and Uruguay. The fully-constructed test, enclosing the third-instar female, is up to 11 mm long and 8.5 mm wide, with an apical orifice about 2 mm in diameter. The test varies from light yellow to red-brown and has a rough texture on the outer surface, whereas the internal surface is smooth and coated with a thin layer of powdery white wax.
Adult female. Body broadly oval, posterior end rounded; derm membranous. Antenna 11 segmented, each segment with numerous setae. Eyespot situated lateral to antennal scape. Mouthparts absent. Legs well developed, each with a large apodeme and with setae on all segments; trochanter with 3 or 4 campaniform sensilla on each side; inner edges of tibia and tarsus with stout setae; claw stout and curved without a denticle. Thoracic spiracle without atrial pores but each peritreme with perispiracular multilocular pores. Abdominal spiracles numbering 7 pairs; atria of anterior spiracles with pores. Vulvar opening indistinct, near end of abdomen. Anal tube poorly developed and anal opening lightly sclerotised. Dorsal multilocular pores circular or broadly oval, with a thickened rim, each mostly with an oval bilocular centre and 10-12 outer loculi, scattered on head, thorax and abdomen; some smaller pores, each up to 8 µm in diameter, along intersegmental lines. Ventral multilocular pores as on dorsum except pores on posterior segments of abdomen, particularly around vulva, with a circular or irregularly oval centre and mostly with 9-13 outer loculi. Dorsal and ventral setae fairly evenly distributed over surface, covered with abundant hairs, hair-like and flagellate setae. Cicatrices and disc-like tubercles absent.
Third-instar (preadult) female. Enclosed in a resinous test with a thin wall. Body almost round to ovoid; derm membranous except posterior end that has a large circular, strongly sclerotised, anal disc bearing scattered multilocular pores and setae. Antenna short, conical, 8 or 9 segmented. Eyespot located posterolateral to antennal scape. Mouthparts well developed; labium 3 segmented; basal segment narrow and difficult to discern. Legs considerably reduced, but each segment present; trochanter with 3 campaniform sensilla on each surface. Thoracic spiracles with each peritreme surrounded by numerous perispiracular multilocular pores. Abdominal spiracles numbering 7 pairs, with 2 posteriormost pairs smallest; each atrium with many multilocular pores. Dorsum and venter covered in multilocular pores, mostly with a trilocular centre and 8-12 outer loculi. Derm covered by mainly hair-like and flagellate setae. Cicatrices, mostly 15-35 µm in diameter, present throughout dorsum and venter apart from anal area and medially on anterior thorax. Disc-like tubercles, each 7-8 µm in diameter, with a thick sclerotised rim and a slightly domed centre, distribution similar to cicatrices except present throughout ventral thorax.
Second-instar nymph. Similar to third-instar female but setae and pores less abundant. Body ovoid; derm membranous except posterior end of body strongly sclerotised, forming an anal disc bearing multilocular pores and setae and with a central anal opening. Antenna 5-7 (mostly 6) segmented. Eyespot situated posterolateral to antennal scape. Mouthparts well developed. Legs present, each segment reduced, total length about 260 µm. Thoracic spiracles with each peritreme surrounded by perispiracular multilocular pores. Abdominal spiracles numbering 7 pairs, with posteriormost 3 pairs smallest; each atrium with a few multilocular pores. Dorsal and ventral multilocular pores mostly with an oval or triangular centre. Derm covered by short setae, mainly hair-like and flagellate, longest at extremity of abdomen. Cicatrices, each 15-25 µm in diameter, present on all segments of dorsum and venter except anal area. Disc-like tubercles, each 7-8 µm in diameter, with distribution similar to cicatrices.
First-instar nymph. Body elliptical; derm membranous apart from a large sclerotised anal disc at posterior end with anal opening and multilocular pores and setae. Antenna 6 segmented, all segments with hair-like setae; apical segment also with fleshy setae. Labium 3 segmented, elongate. Eyespots sclerotised, each almost as large as first antennal segment. Labium 3-segmented. Legs long; claws slender, each with a pair of setose digitules and a denticle near apex. Thoracic spiracles each with 2 or 3 perispiracular multilocular pores. Abdominal spiracles numbering 7 pairs; atria with characteristic corrugated surface, but pores absent. Anal tube with polygonal wax glands at inner end. Derm with multilocular pores mostly each with a trilocular or quadrilocular centre and with scattered short setae. Posterior end of abdomen with 2 pairs of long caudal setae, each about 350-450 µm. Venter with 3 large circular cicatrices on posteromedial abdomen. Dorsum with scattered small disc-like tubercles, each tubercle domed in centre.
Adult male. Derm of membranous areas of dorsum and venter mostly covered with numerous setae, mostly hair-like (hs) and flagellate setae (fs) and with convex loculate pores (lc), convex pores (cp) and small sclerotised pores (ssp). Antenna 10 segmented without nodes; segments with randomly distributed setae; many setae with satellite setae; minute setae present on intersegmental membranes between segments IV-VI; pedicel with a campaniform sensillum distally. Ventral mid-cranial ridge complete. Scutum with a membranous area medially with hair-like scutal setae, but without loculate pores. Legs well developed and very setose, many setae spur-like and some bifurcate, particularly distally on all tibia and posterior surface of profemur; each trochanter probably with 4 round campaniform sensilla on each side; tarsus 2 segmented; claw elongate, without a denticle; claw digitules setose. Abdominal spiracles probably numbering 7 pairs. Posterior margin of basisternum from which furca arises unusually narrow; basisternal setae numerous. Hamulohalteres each with 6-9 hamuli. Abdomen with pairs of long lateral extensions on segments VI, VII and VIII; endophallus eversible and spinose; aedeagus lightly sclerotised, lying in a ventral groove along penial sheath.

Neocoelostoma xerophila Hempel
Neocoelostoma xerophila Hempel, 1932: 311;Lizer y Trelles, 1936: 116;Lizer y Trelles, 1939: 174. Adult female (Fig. 15). Note. There are no adult female type specimens of this species, but adult females of a species that appeared to be N. xerophila were available, collected in Argentina and Uruguay. In order to confirm the identity of the latter adult females, we compared the third-instar females from Brazil described by Hempel with third-instar females collected with the adult females from Argentina and Uruguay. The third-instar females from all three countries appear to be identical and thus below we describe the adult female of N. xerophila based on these specimens from Argentina and Uruguay.
Third-instar (preadult) female (Figs 1D, E, 16). Note. The description, measurements, and drawing are based on specimens from Argentina, Brazil and Uruguay. We also examined several photographs of morphological details made from 2 slides of syntype preadult females, labelled "Instituto Biológico / Neocoelostoma / xerophila H. [or 'Hemp.'] / sp. n. Type. / Coll. A. Hempel / R. 6. III: 31 [or just '6']", from IBSP, São Paulo, Brazil, made available by Dr A.L.B.G. Peronti of Universidade Federal de São Carlos, Brazil. A specimen on one of these original Hempel slides should be designated the lectotype, but we do not take this action here because we have not personally seen the IBSP specimens.
Dorsum. Multilocular pores (Figs 16A, E), each 10-11 µm in diameter, mostly with a trilocular centre (rarely an oval bilocular or quadrilocular centre) and 8-12 outer loculi, scattered on head and thorax, and across abdominal segments. Flagellate setae (Fig. 16C), each mostly 10-15 µm long, sparsely distributed throughout on head and thorax and in a transverse line across each abdominal segment; hair-like setae longer, each up to 40 µm long, rare on margin and sparse on abdomen. Cicatrices (Fig. 16M), each mostly 15-25 µm (a few up to 30 µm) in diameter, present on head and thorax, and in an irregular line across each abdominal segment. Disc-like tubercles (Fig. 16D), each 7-8 µm wide, with a sclerotised thick rim and a slightly domed centre; scattered on head and thorax, and present across each abdominal segment.
Second-instar nymph (Fig. 17). Note. Description is based only on specimens from Argentina and thus may not represent the full range of morphological variation. We also do not know whether the specimens are male or female.