Corresponding author: Scott A. Schneider (
Academic editor: Roger Blackman
Four new species of armored scale insect,
Wei J, Schneider SA, Normark RD, Normark BB (2021) Four new species of Aspidiotini (Hemiptera, Diaspididae, Aspidiotinae) from Panama, with a key to Panamanian species. ZooKeys 1047: 1–25.
Armored scales are the most species-rich family of scale insects, comprising over 2600 species in 418 genera (
Some armored scale insect species are extremely polyphagous, with host ranges among the widest known for any herbivorous insect, comprising in some cases over 100 families of plants (
Armored scales are currently classified into four subfamilies:
The sampling locality for new species described in this paper was the canopy crane in San Lorenzo National Park, Colón (
In this paper, morphological terminology conforms to descriptions and illustrations provided by
Depositories are abbreviated as follows:
This new species is most similar in appearance to
The epithet
Panama (Colón).
This species is placed in the genus
The specific epithet is the Latin genitive of the host plant genus,
Panama (Colón).
DNA sequences from 3 loci of the holotype of
The holotype of
The only other known species in this genus is
The specific epithet is the Latin genitive of the host plant genus,
Panama (Colón).
Several DNA sequences of
Specimens of
This is an unusual species both in its life history, showing intraspecific variation in the pupillarial habit, and in its biogeography, having affinities to African species. Some slide-mounted specimens are unequivocally pupillarial, having well-developed 1st instars inside of adult females that are themselves inside of 2nd-instar cuticles. More often than not, these adult females are flipped inside their puparia, with their head at the posterior end of the puparium. Other specimens are apparently non-pupillarial, and some of these have a sclerotized cephalothorax, a feature not seen, to our knowledge, in adult females of any pupillarial species. We had originally intended to describe the pupillarial and non-pupillarial forms as two different species, but the three sequenced gene regions show no differences between them and there are no consistent morphological differences either; therefore, we consider them to comprise a single species that includes both pupillarial and non-pupillarial developmental phenotypes. The second instar has a more completely developed secretory system than the adult, with more ducts, plates, and lobes – a pattern typical of pupillarial species and opposite to what is typical of non-pupillarial species. This may imply that this species is derived from a pupillarial ancestor and that the non-pupillarial form represents a secondary loss of the pupillarial habit.
Molecular phylogenetic studies have shown that
We tentatively place this species in the genus
Morphologically,
Our study of Neotropical and African species that resemble
The specific epithet is a Latin adjective; here it alludes to this species’ unusual biogeography as a Neotropical member of a mostly African clade.
Panama (Colón).
Panama • 1 adult female; Parque Metropolitano Canopy Crane;
Panama (Parque Metropolitano).
The key incorporates some modified excerpts drawn from
1 | With deep thoracic constriction between prothorax and mesothorax or mesothorax and metathorax |
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– | Without deep thoracic constriction on thorax |
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2 | With deep thoracic constriction between mesothorax and metathorax; |
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– | With deep thoracic constriction between prothorax and mesothorax; |
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3 | Paraphyses absent on pygidium |
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– | Paraphyses present on pygidium |
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4 | Perivulvar pores absent |
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– | Perivulvar pores present |
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5 | With |
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6 | L4 absent, plates beyond |
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– | L4 present, plates beyond |
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7 | Dorsal ducts present in submedian areas of pygidium; entire body strongly sclerotized at maturity |
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– | Dorsal ducts absent from submedian areas of pygidium; cephalothorax slightly sclerotized at maturity or body membranous |
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8 |
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9 | Pre-pygidial marginal macroducts absent; with total of 15–29 dorsal macroducts on each side of body |
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– | Pre-pygidial marginal macroducts present; with total of 22–38 dorsal macroducts on each side of body |
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10 | Plates present anterior to |
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– | Plates absent anterior to |
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11 | Prosoma of mature female reniform; with 3 long fleshy plates laterad of |
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– | Prosoma of mature female elongate, round, oval or turbinate, not reniform; combination of plates laterad of |
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12 | Body elongate and more or less parallel-sided, 3× or 4× as long as wide ( |
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– | Body round, turbinate, or oval, less than 3× as long as wide |
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13 | Body long and quite slender, cephalic margin almost straight; apical angle of pygidium more than 90 degrees; perivulvar pores in 5 groups |
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– | Body elongate but broad, cephalic margin broadly rounded; apical angle of pygidium less than 90 degrees; perivulvar pores in 4 groups |
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14 | Most paraphyses shorter than or similar in length to |
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– | Paraphyses typically longer than |
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15 | Anal opening relatively large, distance between posterior edge of opening and base of |
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– | Anal opening small, distance between posterior edge of opening and base of |
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16 | Perivulvar pores absent |
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– | Perivulvar pores present |
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17 | Having the following combination of characters, plates in the first and second interlobular spaces all simple, |
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– | Without this combination, at least some fringed plates present in first and second interlobular spaces, |
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18 | Plates anterior to position of |
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– | Plates anterior to position of |
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19 |
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20 |
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21 |
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22 |
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23 | Plates deeply fringed, definitely exceeding |
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– | Plates shallowly fringed, only slightly exceeding |
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24 | Paraphyses arising from lateral angle of |
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– | Paraphyses arising from lateral angle of |
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25 | Perivulvar pores present, at least 1 pore per side |
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– | Perivulvar pores entirely absent |
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26 | With 2 plates between |
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– | With 1 plate between |
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27 | With |
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– | With 4 pairs of well-developed lobes; plates simple and much shorter than |
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28 | Paraphyses arising only from basal angles of lobes or position of obsolete lobes, never from within interlobular spaces; paraphyses in first interlobular space typically about 2× longer than those in second interlobular space (although nearly identical in length for 1 species); perivulvar pores absent ( |
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– | With at least 1 paraphysis arising from an interlobular space; paraphyses in first and second interlobular spaces not following this pattern; perivulvar pores present or absent |
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29 | Plates reduced to short membranous lobes, rounded apically |
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– | Plates present or absent, if present, elongate |
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30 | Plates entirely lacking; with 1 notch on each side of each lobule |
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– | Plates well-developed; with 1 notch on outer margin of each lobule of |
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31 | Having combination of 3 pairs of pygidial lobes, 1 paraphysis arising from first interlobular space, and lacking paraphyses beyond |
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– | Without above combination; paired paraphyses in first space arise from outer angles of lobes, with 3–4 well-developed lobes and with or without paraphyses present beyond |
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32 | Pygidial margin anterior to L4 not heavily sclerotized, without series of short paraphyses; plates anterior to |
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– | Pygidial margin anterior to L4 heavily sclerotized, often with series of short paraphyses; plates anterior to |
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33 | First 2 plates anterior to |
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– | First 2 plates anterior to |
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34 | Pygidium long and narrow, sharply tapering to acute apical point, lateral margins slightly concave; with 3 pairs of pygidial lobes, L4 reduced to point or absent; margin anterior to L4 heavily sclerotized; most paraphyses between |
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– | Pygidium short and broad, not tapering apically to acute point, lateral margins convex; usually with 4 or 5 pairs of pygidial lobes, L4 usually well developed; margin anterior to L4 lightly to heavily sclerotized; paraphyses between |
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35 | Lateral thoracic margin produced into distinct point or rounded umbo near posterior spiracles |
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– | Lateral thoracic margin without such point or umbo, at most with small, sclerotized spot in this position |
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36 | Lateral thoracic margins produced into very pronounced umbos; sclerotization of derm developed in sharply defined marginal zone extending from umbos, along sides, and across pygidium |
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– | Umbos quite small, sclerotization forming similar pattern as above, but rather weakly developed | |
37 | 1 very long paraphysis arising from outer angle of |
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– | 1 long paraphysis arising from outer angle of |
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38 | Pygidium with longest paraphyses arising from lateral angles of lobes; large V-shaped reticulate sclerotized area on abdominal segment VI always present ( |
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– | Pygidium with longest paraphyses arising from interlobular spaces; without large V-shaped reticulate sclerotized area on abdominal segment VI |
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39 | With 5 pairs of lobes; with 2 plates anterior to L4; ventral microducts present between |
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– | With 4 pairs of lobes, without plates anterior to L4; ventral microducts absent between |
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40 | Anterior head margin of mature female forming distinctly sclerotized prominence, differentiated from lateral margin, resembling a “cap” ( |
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– | Anterior head margin of mature female not distinctly sclerotized or differentiated from lateral margin, not resembling a “cap” ( |
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41 | Perivulvar pores present in 5 small groups; cephalic area very heavily sclerotized and bearing series of conspicuous setae |
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– | Perivulvar pores absent; cephalic area without series of conspicuous setae |
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42 |
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43 | With sclerotized spur on head; longest paraphysis in third interlobular space arising from center of interlobular space, posterior to L4 |
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– | Without sclerotized spur on head; longest paraphysis in third interlobular space arising near mesal angle of L4 |
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44 | Perivulvar pores present |
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– | Perivulvar pores absent |
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45 |
Perivulvar pores present in 5 small groups; pygidium moderately acute at apex, lateral margins almost straight; first interlobular space with moderately long, apically swollen paraphysis followed by small process from mesal angle of |
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Perivulvar pores present in 4 small groups; pygidium with lateral margins convergent; first interlobular space with quite long paraphysis followed by very small process from mesal angle of |
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46 |
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– | With long paraphysis between |
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47 | Pygidial lobes each with dorsal seta sunk to at least 0.25× length in distinct socket |
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– | Pygidial lobes with dorsal setae not in distinct sunken sockets |
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48 | With median band of dermal reticulations, squamations or transverse striations (median squamations may be indistinct) |
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– | Without median band of dermal reticulations, squamations or transverse striations (median striations, if present, longitudinal) |
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49 | Third interlobular space with longest paraphysis about equal in length to longest paraphysis in first and second interlobular spaces; venter without long microducts |
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– | Third interlobular space with longest paraphysis much shorter than longest paraphysis in first space or second interlobular spaces; venter with long microducts along pygidial margin anteriorly and in 2 irregular submarginal rows posterior to ventral pygidial scar |
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50 | With only 1 paraphysis in third interlobular space; with long microducts in 3 distinct longitudinal bands on each side of vulva on venter |
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– | With various numbers of paraphyses in third interlobular space; with long microducts not arranged in distinct longitudinal bands on each side of vulva, or with 2 or fewer such bands |
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51 | Usually with reduced paraphyses in third interlobular space; orifices of macroducts large and conspicuous in dorsal sclerotized areas |
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– | Usually with 1 or more well-developed paraphyses in third interlobular space; orifices of macroducts smaller and less conspicuous in dorsal sclerotized areas |
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52 | With 3 paraphyses in second interlobular space; ventral seta in middle of or anterior to base of lobe on each |
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– | With 2 paraphyses in second interlobular space; ventral seta laterad of base of lobe on each |
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53 | Orifices of macroducts absent on lateral margin of dorsal sclerotized area 2; paraphyses in third interlobular space nearly equal in length |
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– | Orifices of macroducts present on lateral margin of dorsal sclerotized area 2; paraphysis arising from outer angle of |
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54 | Anal opening between or only slightly anterior to apices of paraphyses; without definite paraphyses beyond L4 |
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– | Anal opening decidedly anterior to apices of all paraphyses; with paraphyses beyond L4 |
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55 | Pygidial margin with 2 or 3 conspicuous, spur-like processes anterior to L4 |
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– | Pygidial margin without spur-like processes anterior to L4 |
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56 | Anal opening located within posterior third of pygidium from base of median lobe; with 5 paraphyses in third interlobular space |
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– | Anal opening located near center of pygidium; with 3–4 paraphyses in third interlobular space |
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57 | Without macroduct orifices on membranous area in third interlobular space; all macroduct orifices with equal diameters |
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– | With macroduct orifices on membranous area in third interlobular space; macroduct orifices in third interlobular space usually smaller in diameter than any macroduct orifices located more mesally (on abdominal segments VI–VIII) |
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Additional online resources aiding in the identification of
For logistical support for collecting in Panama, we thank D. Windsor, M. Samaniego, D. Quintero Arias, E. Andrades, J. Herrera and the Smithsonian Tropical Research Institute, and for doing much of the actual collecting, we thank G.E. Morse, D.A. Peterson, H. Shapiro, and S. Trujillo. The specimens were mounted by M. Fizdale, T. Jackson, M. Knoll, M. Logan, R. McCarthy, K. Mullen, K. Romeu, and M. Wilken. This work was supported through a National Science Foundation grant (DEB-1258001) awarded to BBN. Additional support was provided by the U. S. Department of Agriculture’s National Institute of Food and Agriculture, and by the University of Massachusetts Amherst Biology Department and Center for Agriculture, Food, and the Environment, under project number MAS00535. Research and manuscript preparation were also supported in part by the U.S. Department of Agriculture, Agricultural Research Service. JW’s participation was supported by the China Scholarship Council. Mention of trade names or commercial products in this publication is solely for the purpose of providing specific information and does not imply recommendation or endorsement by the U.S. Department of Agriculture; USDA is an equal opportunity provider and employer. The authors report no conflicts of interest.