Research Article |
Muhammad Z. Ahmed‡§, Matthew R. Moore‡, Eric A. Rohrig‡, Cindy L. McKenzie§, Di Liu|, Jinian Feng|, Benjamin B. Normark¶, Douglass R. Miller#‡
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Corresponding author: Muhammad Z. Ahmed ( muhammad.ahmed@usda.gov ) Academic editor: Roger Blackman
© 2021 Muhammad Z. Ahmed, Matthew R. Moore, Eric A. Rohrig, Cindy L. McKenzie, Di Liu, Jinian Feng, Benjamin B. Normark, Douglass R. Miller.
This is an open access article distributed under the terms of the CC0 Public Domain Dedication.
Citation:
Ahmed MZ, Moore MR, Rohrig EA, McKenzie CL, Liu D, Feng J, Normark BB, Miller DR (2021) Taxonomic and identification review of adventive Fiorinia Targioni Tozzetti (Hemiptera, Coccomorpha, Diaspididae) of the United States. ZooKeys 1065: 141-203. https://doi.org/10.3897/zookeys.1065.69171
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Abstract
This work provides general descriptions, illustrations, molecular diagnostic data, taxonomic keys, slide mounting recommendations, and Florida distribution records for Fiorinia Targioni Tozzetti species occurring in the USA. Species treated are F. externa Ferris, F. fioriniae (Targioni Tozzetti), F. japonica Kuwana, F. pinicola Maskell, F. phantasma Cockerell & Robinson, F. proboscidaria Green, and F. theae Green. New descriptions of second-instar males and females of all seven species in addition to first-instar nymphs and adult females of F. phantasma and F. proboscidaria are presented. Taxonomic keys to second-instar males and females are developed for the first time and previously available taxonomic keys to first-instar nymphs and adult females are improved. DNA sequences were used to further evaluate the monophyly of Fiorinia and provide additional diagnostic tools for Fiorinia species. Multigene phylogenetic analyses, COI barcoding methods, and examination of type material indicate that F. yongxingensis Liu, Cai & Feng, 2020, syn. nov. is a junior synonym of F. phantasma. A morphological survey of the genus demonstrates, for the first time, the utility of second-instar males for diagnostics. This study will help inform regulatory and pest management decisions by facilitating morphological and molecular identification of adventive Fiorinia species occurring in the USA.
Keywords
Armored scale insects, DNA barcodes, Florida, palms, phantasma scale, slide mounting
Introduction
The genus Fiorinia (Hemiptera, Diaspididae) comprises 70 species (
Fiorinia phantasma, commonly known as phantasma scale, was described from the Philippine Islands in 1915. Subsequently, a major global expansion of F. phantasma occurred over the last decade through movement of nursery stock (
The first North American continental report of F. phantasma was in Florida and included more than twenty heavily infested Canary Island date palms (Phoenix canariensis Chabaud) along both sides of a road in Miami-Dade County (
The main pest management challenge is detection of new F. phantasma infestations. Fiorinia phantasma occurs in two Florida counties, Miami-Dade and Palm Beach, and is usually found on palms (
The purpose of this study is to provide taxonomic keys for immatures of seven Fiorinia species occurring in the USA. We also provide line drawings and diagnoses of slide-mounted second-instar males and females, DNA sequence data for multiple loci for molecular diagnostics, and extensive records of the species’ distributions in Florida. We newly describe and illustrate first-instar nymphs and adult females of two species, F. phantasma and F. proboscidaria. In addition, we provide updated taxonomic keys for first-instar nymphs (adapted from
Materials and methods
Taxon sampling
Four species of Fiorinia (F. fioriniae, F. phantasma, F. proboscidaria, F. theae) were collected from Florida (Suppl. material
In addition to the freshly collected Fiorinia specimens described above, additional specimens and sequences were included in analyses of DNA sequences (Suppl. material
Slide mounting of immature stages
Slide mounting is considered mandatory for morphological identification of armored scale insects because it is nearly impossible to identify taxonomic features without doing so. Moreover, for museum curation purposes, slide mounting is the best way to archive scale insects in a reference collection. There are studies available on methods for mounting hemipterans (
(i) Standard slide mounting method (6 steps)
Initially, a 67 mm beveled-edge watch glass (Prolab Scientific) and micro spatula were used. Fisher 10% potassium hydroxide (KOH) was used in step 1 for heating and maceration. Following this step, specimens were placed into a Humboldt mesh (Replacement Mesh Disk 5 cm dia. No. 325H-3807.325) container that was then placed inside the watch glass, eliminating the need for the micro spatula in the following steps until the final mount. Forceps (Bioquip Swiss style #4) were used to remove the mesh from the 4.8 cm watch glass (Item#742300, Carolina) while switching between steps. Glacial acetic acid (Fisher) was used in step 2 for removal of the remaining 10% KOH from step 1. Acid fuchsin stain (Bioquip) was used with a 3:7 dye to acetic acid ratio in step 3 to stain the specimens. For dehydration of the cuticle in the 4th step, 75% and 95% EtOH were used. Clove oil (Spectrum Chemical) was used in step 5 to remove any remaining wax from the specimens. A disposable transfer pipette (13-711-9D, Fisher) which holds 3.2 ml, was used in steps 2–5. In the final step 6 filtered Canada balsam (CAS 8007-47-4, Fisher) was used as the medium and placed on a glass slide (22-038-103, Fisher). A glass coverslip (12-545-80P, Fisher) was placed on the specimen in balsam to complete the mount. The 6 steps required for the standard slide mounting method are as follows:
- Heating: specimens were set in a watch glass filled with 10% KOH and heated at 85 °C for 5–10 mins. After heating, gut contents were teased out using a micro-spatula to gently tap the dorsum.
- KOH removal: specimens were moved to a watch glass of 95% glacial acetic acid for 10 mins to remove any remaining KOH.
- Staining: Acid fuchsin stain was added and let sit for 5 mins.
- Stain correcting: specimens were moved to a watch glass of 75% EtOH for 10 mins. Specimens were then placed in 95% EtOH for another 10 mins to dehydrate.
- Wax removal: specimens were soaked in clove oil for 5 to 10 mins. This helps to remove any remaining wax or lipids and makes specimen bodies flexible to be easily spread on a slide.
- Mounting: on a labeled slide, a drop of balsam was placed in a center and spread to avoid specimen drift. The specimen was then placed in the balsam dorsoventrally (i.e. ventral side up) and legs and antennae were positioned properly. A coverslip was placed on the balsam, and the slide was placed on a hot plate at 30 °C for 10 mins to remove any bubbles.
Due to the multiple steps in this method, which require each specimen to be moved from 5 different watch glasses before mounting, many first-instar nymphs were lost or damaged. Additionally, this method was time consuming. In an attempt to reduce the loss of first-instar nymphs, minimize damage, reduce the amount of chemical usage, and save time, we subsequently developed alternative methods – see below.
(ii) Modified slide mounting method A (1 step)
For fresh specimens (not preserved in ethanol).
- Mounting: on a labeled slide, a drop of Hoyer’s medium was placed in the center and spread to avoid specimen drift. Fresh specimens picked from plant material were placed in a Hoyer’s medium dorsoventrally and legs and antennae were positioned properly. In this protocol, we omitted steps 1–5 of the standard method and mounted specimens directly into Hoyer’s medium. This was effective in preventing loss of specimens and reducing the amount of chemical usage.
(iii) Modified slide mounting method B (4 steps)
For ethanol-preserved specimens.
- Heating: specimens were placed in a watch glass filled with 10% KOH and heated for 5 mins at 85 °C.
- Rehydrating: specimens were placed in water and left to soak for 5–10 mins. We found that heating the specimens prior to submerging them in water aided in the rehydration process.
-
Cleaning: specimens were moved to a watch glass filled with Hoyer’s medium. Because Hoyer’s medium is a self-cleaning fluid (
Anderson 1954 ), specimens were placed in the dish to accelerate the cleaning. - Mounting: on a labeled slide, a drop of Hoyer’s medium was placed in the center and spread to avoid specimen drift. The specimen was placed in the Hoyer’s medium dorsoventrally and legs and antennae were positioned properly.
(iv) Modified slide mounting method C – balsam method with mesh container (7 steps)
For fresh specimens and ethanol-preserved specimens.
- Heating and cleaning: specimens were placed in a watch glass filled with 10% KOH and heated at 85 °C for 5–10 mins. After heating, cavity contents were teased out using a micro-spatula. Once this step was completed, specimens were moved to a container modified using mesh placed in a watch glass (Fig.
1 ). The modified mesh container was made using a plastic 5 ml screw-top tube and fine wire mesh (Humboldt, Elgin, IL United States). The top was cut out of the screw-top tube and the mesh was put in its place, allowing liquid to move through the mesh while keeping the specimens inside. - Rehydration: specimens were placed in water and left to soak for 5 to 10 mins.
- KOH removal: specimens were moved to a watch glass of 95% acetic EtOH (a few drops of glacial acetic acid with 95% ethanol) for 10 mins.
- Staining: after removing 95% acetic EtOH, acid fuchsin stain was added and let sit for 5 mins.
- Stain correction and dehydration: specimens were moved to a watch glass of 75% EtOH for 10 mins. Specimens were then placed in 95% EtOH for another 10 mins to dehydrate the cuticles.
- Wax removal: specimens were soaked in clove oil for 5 to 10 mins.
- Mounting: on a labeled slide, a drop of balsam was placed in a center and spread to avoid specimen drift. The specimen was placed in the balsam and legs and antennae were positioned properly. A coverslip was placed on the balsam and the slide was placed on a hot plate at 30 °C to remove any bubbles.
Although the mesh is effective in keeping first- and second-instar nymphs in the container without damage, a few problems were noted. The mesh does not sit flat against the glass bottom of the watch glass, so the cleaning step cannot be done in the mesh. Cleaning must be done in a watch glass and then specimens must be moved back into the mesh for the remaining steps. Due to the smaller size of the mesh container, range of motion using microtools throughout this process can be limited. Similar to processing in a watch glass without mesh, specimens can get stuck on the upper sides of the modified dish. Visibility of first-instar nymphs can be hampered by the reflective coloration of the mesh.
There are several steps involved in traditional slide-mounting protocols (method i) that require each specimen to be moved to and from at least five different watch glasses before eventually being slide mounted. Many first-instar nymphs can be lost or damaged during these steps. We recommend using the mesh container during the slide-mounting protocol (method iv). Use of this container will decrease mounting time, reduce specimen loss, decrease the quantity of chemical reagents, and generate quality slides. All steps can easily be performed using the mesh container except for the cleaning step. Unfortunately, the cleaning step must be done in a watch glass and then the specimens should be moved back into the mesh container to finish the mounting process. Although this procedure is laborious, we recommend it when the aim is to make permanent mounts for deposit in archival collections. The other mounting procedure is to place first-instar specimens directly into Hoyer’s mounting medium on a slide (method ii, iii). This protocol has fewer steps and less chance of specimen loss, and yields specimens with superior visibility. We recommend this protocol for rapid species diagnosis. Unfortunately, the mounts are only temporary unless slides are ringed to prevent deterioration.
DNA extractions, polymerase chain reaction (PCR), and sequencing
DNA was extracted from individual Fiorinia and Thysanofiorinia specimens using the Qiagen Blood and Tissue Kit per the manufacturer’s protocol. Extractions were non-destructive, and recovery of individual scale vouchers was attempted. DNA was quantified on a Nanodrop 2000 and PCRs had a target input of at least 5 ng of genomic DNA. PCRs were performed using the Kapa HiFi HotStart PCR Kit, in a total volume of 25 uL.
The standard cytochrome oxidase I (COI) barcode region was targeted for each species using the primer pair PCOF1 (
Two other loci, the large ribosomal subunit (28S D2/D3 expansion region) and elongation factor 1α (EF1α) were also targeted, for comparison with the results of
PCRs were visualized on 1.5% agarose gels. Positive PCRs were purified and prepared for sequencing using BigDye Terminator v3.1 chemistry. Amplicons were sequenced bidirectionally on the ABI SeqStudio platform at FDACS-DPI. Sequence chromatograms were trimmed and assembled in Sequencher 5.4.6. Newly generated sequences were deposited in GenBank (Suppl. material
Data analysis
Cytochrome oxidase I barcode sequences (5’-COI) were initially aligned using an online version of MAFFT 7 (
Sequences were aligned using the default settings of MUSCLE (
Neighbor-joining and distance analyses of the 5’-COI matrix were conducted in MEGA X (
Phylogenetic analyses were conducted using 3 sequence regions reported in
Maximum Likelihood (ML) phylogenetic analyses were conducted on the XSEDE computing cluster as part of the CIPRES Science Gateway (
Node support was assessed by 10,000 ultrafast ML bootstrap replicates (
Results
Phylogenetic analyses
Maximum Likelihood analyses estimated a consensus bootstrap tree with a log-likelihood of -20,889.543 for the multigene tree (Fig.
Maximum Likelihood bootstrap consensus tree of the subtribe Fioriniina based on 28S, EF1-α, 5’-COI, 3’-COI, and COII. The clade highlighted in red indicates a monophyletic Fiorinia. The close-up of Fiorinia clade is presented on right side. High-resolution figure of the main tree is in Suppl. material
The slide-mounted cuticle of D4815B and other specimens in the same lot have been re-examined by BBN and they clearly belong to a pupillarial species whose morphology is completely consistent with the genus Fiorinia. These results might imply that the lineage leading to D4815B and D4815C represents a second origin of the pupillarial habit in Fioriniina. These two isolates were placed within a section of a Fiorinia + Rolaspis + Pseudaulacaspis (in part) clade in the ML phylogenetic tree using only 28S data (Suppl. material
COI barcoding
This study produced 43 new sequences of the COI barcode region, 37 of which represent nine Fiorinia species (Fig.
Neighbor-joining tree of Diaspididae 5’-COI barcodes. Terminal taxa are labeled to their narrowest identification-level. Numbers in parentheses after terminal taxa indicate how many sequences are represented in each cluster. The cluster of Fiorinia species is highlighted in red. Bootstrap support values greater than 75 are indicated on the tree. Nodes with 100 percent bootstrap support are indicated by a “*”. The close-up of the Fiorinia clade is presented on the right side. High-resolution figure of the main tree is in Suppl. material
The alignment for calculating K2P distances among Fiorinia species included 37 terminal taxa and was 645 bp long. Based on the 95% site cutoff, calculations involved 560 total positions. Intraspecific K2P distances were low, except for specimens identified as F. theae (Table
| Species | Intra. K2P Dist. | Inter. K2P Dist. |
|---|---|---|
| Fiorinia externa (n = 6) | 0.00% | 9.1%–12.4% |
| Fiorinia fioriniae (n = 7) | 0.00%–0.02% | 11.8%–14.8% |
| Fiorinia phantasma (n = 7) | 0.00%–0.09% | 9.1%–13.7% |
| Fiorinia pinicola (n = 3) | 0.00% | 9.1%–15.2% |
| Fiorinia proboscidaria (n = 6) | 0.00%–0.02% | 9.9%–14.2% |
| Fiorinia theae (n = 5) | 0.00%–8.00% | 9.5%–14.8% |
| Fiorinia sp. isolate D4778A (n = 1) | N/A | 9.5%–13.9% |
| Fiorinia sp. isolate D4674F (n = 1) | N/A | 9.5%–12.7% |
| Fiorinia sp. isolate D4682A (n = 1) | N/A | 9.1%–15.2% |
General descriptions of second-instar nymphs of Fiorinia species occurring in the USA
Second-instar females
With two definite pairs of lobes; third lobes and sometimes fourth lobes represented by series of points. Median lobes yoked, medial margins divergent or nearly parallel, longer than lateral margin, with series of notches. Second lobes bilobed, usually smaller than median lobes, sometimes wider, medial lobule largest, sometimes with small notches, lateral lobule sometimes with one or two small notches. Third lobes usually represented by raised sclerotized area with small series of notches, often divided into two lobules by seta marking segment VI. Fourth lobes sometimes represented by series of sclerotized points. Gland spine arrangement of two types: F. proboscidaria and F. theae with single gland spine on each side of each of segments II–VIII, gland spines on each side of segments II–IV larger than those on segments V–VIII, without gland spines on segment I; remaining species with single gland spine on each side of segments II–V, absent from segment VI, present on each side of segments VII and VIII, gland spines on each side of segments II–V larger than those on segments VI–VIII, with two or three smaller gland spines on each side of segment I; gland spines with barely perceptible sclerotization posterolaterad of each spiracle. Macroducts barrel shaped, marginal, with four or five on each side of pygidium from segments III or IV–VII. Microducts restricted to venter, three different patterns on abdomen; in F. proboscidaria and F. theae longitudinal lines on each side of abdomen from II–VI, each line composed of one or more ducts on each segment, mediolateral line on segments III or IV, V or VI, submarginal line on segments II–VI; in F. externa, F. fioriniae, F. japonica, and F. pinicola longitudinal lines on each side of abdomen from II–VI, each line composed of one or rarely two ducts on each side of each segment, mediolateral line on segments II–V or VI, submarginal line on segments II–VI; in F. phantasma longitudinal lines restricted to mediolateral areas of segments II–IV or V, other lines absent; microducts on head and thorax usually anterior of clypeus, laterad of labium, and posterior of each spiracle. Perispiracular pores associated with anterior spiracles only, with three loculi, one or two pores associated with each spiracle. Anal opening normally located in center of pygidium mesad of fourth marginal macroduct counting forward from posterior macroduct. Dorsal setae present near body margin on head and thorax, with one seta submarginally on each side of each abdominal segment; also present in mediolateral area on each side of body on any or all of abdominal segments I–VI; usually with one mediolateral seta on each side of head. Ventral setae in small numbers in marginal areas of head and thorax, with one seta usually present laterad of each spiracle; abdominal segments with one marginal and one submarginal seta on each side of each segment and with one mediolateral seta on each side of segments IV–VI. Antennae each normally with one long seta and two small sensillae. Cicatrices present or absent on each side of abdominal segment I. Two inconspicuous lobes present submarginally on head of F. proboscidaria and F. theae.
Notes
Characters most useful in distinguishing among species are: a) number of marginal macroducts; b) arrangement of gland spines; c) arrangement of microducts; d) presence or absence of cicatrices; e) presence or absence of lobes on head; f) relative size of median lobes compared to medial lobule of second lobe; g) shape of median lobes.
Second-instar females of Fiorinia species can be distinguished from most similar genera by having the following: median lobes yoked, usually divergent, medial margin longer than lateral, with one pair of setae between; dorsal macroducts confined to body margin, with four or five on each side of pygidium; with two pairs of definite lobes, second pair bilobular. However, we have been unable to distinguish between second-instar females of the Fiorinia species treated here and Pseudaulacaspis cockerelli (Comstock) and P. pentagona (Targioni Tozzetti). There are consistent differences in the distribution of the gland spines in most species of Fiorinia, but F. proboscidaria and F. theae are identical to P. cockerelli and P. pentagona. It is remarkable that the second-instars are so similar, but the adult females are quite different.
Second-instar males
With two definite pairs of lobes; remaining body margin often with numerous projections, not organized into clear lobes. Median lobes spaced apart, without zygosis, usually with small medial lobule and large, conspicuous lateral lobule, medial lobule with one or two projections, lateral margin with several notches and projections. Second lobes usually associated with a dense cluster of marginal ducts, with series of projections, rarely bilobed, smaller than median lobes. Gland spines of three sizes: largest in clusters posterolaterad of each anterior spiracle, posterolaterad of posterior spiracle, and submarginal on abdominal segment I and sometimes II; medium-sized gland spines on body margin of anterior abdominal segments; small gland spines laterad of anterior spiracle on F. externa and F. theae. Macroducts barrel shaped, of two sizes: larger ducts grouped into communal ducts (= glanduliferous craters;
Notes
Characters most useful in distinguishing among species are: a) arrangement and number of communal ducts b) organization of duct clusters c) arrangement of microducts; d) arrangement of gland spines. Second-instar males of Fiorinia are remarkably similar to the same instar of Pseudaulacaspis species by each having unusual lobes, duct clusters, and communal ducts (
First-instar nymphs
First-instar nymphs of Fiorinia species can be recognized by having the following combination of characters: antennae five segmented; apical segment annulate; large duct on each side of dorsum of head; submedial longitudinal line of microducts on each side of thorax; second lobes bilobulate. First-instar nymphs of Fiornia species are similar to some species of Pseudaulacaspis (P. cockerelli and P. pentagona) but differ by normally having a submedial longitudinal line of microducts on each side of thorax, whereas these ducts are absent from P. cockerelli and P. pentagona (
Keys to Fiorinia species occurring in the USA using immature instars
First-instar nymphs (adapted from Howell 1977 )
| 1 | Lobules of pygidial lobe 2 rounded | 2 |
| – | Lobules of pygidial lobe 2 truncate | F. externa |
| 2 | Gland spines on segment VI more than ½ length of gland spine on segment VII | 3 |
| – | Gland spines on segment VI less than ½ length of gland spine on segment VII | 5 |
| 3 | Sclerotized pattern outside of oval surrounding large duct on dorsum of head mostly thin and serpentine like; large duct short, broad, with inner apex nearly flat | 4 |
| – | Sclerotized pattern outside of oval surrounding large duct on dorsum of head mostly in clumps; large duct elongate, narrow, with inner apex mushroom shaped |
F. phantasma (Fig. |
| 4 | Gland spine on segment VI ca. ½ length of gland spine on segment VII | F. fioriniae |
| – | Gland spine on segment VI nearly equal to length of gland spine of segment VII |
F. proboscidaria (Fig. |
| 5 | Dorsal submedian thoracic ducts present; inner apex of large duct on dorsum of head flat | 6 |
| – | Dorsal submedian thoracic ducts absent (occasionally orifices present); inner apex of large duct on dorsum of head mushroom shaped | F. japonica |
| 6 | Gland spine on segment VI noticeably longer than those on segments I–V; pattern of dorsal derm on abdomen fine | F. pinicola |
| – | Gland spine on segment VI equal to those on segments I–V; pattern of dorsal derm on abdomen coarse | F. theae |
Second-instar females
| 1 | With 5 pairs of marginal macroducts | 2 |
| – | With 4 pairs of marginal macroducts |
F. theae (Fig. |
| 2 | With submarginal longitudinal line of microducts on venter; with 4 large-sized gland spines on each side of body; without deep incision anterior of 5th macroduct (segment III) on older specimens | 3 |
| – | Without submarginal longitudinal line of microducts on venter; with 3 large-sized gland spines on each side of body; with deep incision anterior of 5th macroduct (segment III) on older specimens |
F. phantasma (Fig. |
| 3 | Median lobes broad, as wide as or wider than medial lobule of second lobe | 4 |
| – | Median lobes narrow, narrower than medial lobule of second lobe |
F. externa (Fig. |
| 4 | With 3 pairs of microducts on head; space between bases of median lobes wider than medial lobule of second lobes |
F. fioriniae (Fig. |
| – | With 1 pair of microducts on head; space between bases of median lobes equal to or narrower than medial lobule of second lobes |
F. japonica (Fig. |
Second-instar males
| 1 | One or 2 duct clusters on each side of body, or definitive clusters absent | 2 |
| – | Three duct clusters on each side of body |
F. externa (Fig. |
| 2 | Communal ducts present; small macroducts on pygidial margin in at least 1 cluster | 3 |
| – | Communal ducts absent; small macroducts on pygidial margin not in tight cluster |
F. fioriniae (Fig. |
| 3 | Communal ducts incorporated in cluster of small macroducts | 5 |
| – | Communal ducts separate, not in cluster with small macroducts | 4 |
| 4 | With 1 communal duct on each side of pygidium |
F. proboscidaria (Fig. |
| – | With 2 communal ducts on each side of pygidium |
F. theae (Fig. |
| 5 | With 1 communal duct on each side of body |
F. phantasma (Fig. |
| – | With 2 communal ducts on each side of body | 6 |
| 6 | With 5 or more gland spines on each side of body between anterior and posterior spiracles |
F. pinicola (Fig. |
| – | With fewer than 5 gland spines on each side of body between anterior and posterior spiracles |
F. japonica (Fig. |
Adult Females (adapted from Watson et al. (2015)
| 1 | Interantennal process absent | 2 |
| – | Interantennal process present | 5 |
| 2 | Antennae each with a long spur making them longer than wide | 3 |
| – | Antennae each with a short spur making them more or less as long as wide | F. externa |
| 3 | Fewer than 7 marginal macroducts on each side of pygidium | 4 |
| – | Seven or 8 marginal macroducts on each side of pygidium | F. pinicola |
| 4 | Four to 6 (normally 5) marginal macroducts on each side of pygidium; clusters of ventral microducts near body margins of abdominal segments III and IV | F. japonica |
| – | Three or 4 (normally 3) marginal macroducts on each side of pygidium; clusters of ventral microducts absent near body margins of abdominal segments III and IV | F. fioriniae |
| 5 | Interantennal process without spicules; body narrow, with almost parallel sides | 6 |
| – | Interantennal process with spicules; body wide, narrowing abruptly to triangular pygidium |
F. phantasma (Fig. |
| 6 | Seven or 8 marginal macroducts on each side of body; head rounded | F. theae |
| – | Three to 5 marginal macroducts on each side of body; head conical |
F. proboscidaria (Fig. |
Species accounts
Fiorinia externa, second-instar female, Alleghany Co., Glade Creek, North Carolina, November 22, 2019, on Abies fraseri, A. Bartlett, (2019-6449). Abbreviations: a) antenna; b) anterior spiracle; c) microduct with sclerotized orifice; d) large gland spine; e) small microduct; f) enlargement of pygidium.
Fiorinia externa
Field characteristics
First-instar exuviae barely touching second-instar exuviae. Distinct indentation formed between attachment of first- and second-instar exuviae. Second-instar exuviae narrow, parallel sided, and elongate; longitudinal ridge absent or weakly developed. Second-instar exuviae reddish brown anteriorly and light brown to yellow posteriorly. Posterior end of adult female within second-instar exuviae rounded (Suppl. material
First-instar nymph
Described in
Second-instar female
Median lobes slender, narrower than medial lobule of second lobe, not projecting beyond medial lobule of second lobes. With five pairs of marginal macroducts. Swelling of body margin adjacent to macroduct usually pointed. With four large gland spines on margin of each side of body from abdominal segments II–V; usually without small gland spine on each side of abdominal segment VI; with small gland spines on margin or submargin of abdominal segment I. With one microduct on each side of head. Longitudinal line of microducts present submarginally on venter of II–V, normally with one microduct on each side of each segment. Cicatrices absent.
Second-instar male
Three duct clusters on each side of body; posterior cluster composed of several small ducts and two communal ducts. Five longitudinal lines of microducts on venter of abdomen (one medial, two mediolateral, two submarginal). Cluster of small microducts with sclerotized orifice laterad of anterior spiracle. Fewer than five large-sized gland spines on each side of body between anterior and posterior spiracles. Antennae each with one enlarged seta.
Florida collection records
All records are on Christmas trees imported from states outside of Florida. This species is not established in Florida, and its common host, Abies fraseri, also does not occur naturally in Florida. It has been found on imported Christmas trees in the following localities in Florida: Broward Co., Miramar, November 20, 2013, on Abies fraseri, S. Alspach (2013-8494); Broward Co., Davie, December 10, 2013, on Abies fraseri, S. Beidler (2013-8906); Citrus Co., Inverness, December 4, 2013, on Abies fraseri, S. Jenner (2013-9766); Hamilton Co., White Springs, December 11, 2012, on Abies fraseri, H. Randolphs (2012-9239); Hillsborough Co., Tampa, November 20, 2012, on Abies fraseri, T. Streeter (2012-8844); Marion Co., Ocala, December 2, 2013, on Abies fraseri, S. Wayte (2013-8755); Monroe Co., Tavernier, November 28, 2012, on Abies fraseri, J. Farnum (2012-8924); Volusia Co., Port Orange, November 27, 2017, on Abies fraseri, K. Coffey (2017-4496).
Fiorinia externa, second-instar male, Alleghany Co., Glade Creek, North Carolina, November 22, 2019 on Abies fraseri, A. Bartlett, (2019-6449). Abbreviations: a) anterior spiracle; b) microduct with sclerotized orifice; c) large microduct; d) large gland spine; e) small gland spine; f) small microduct; g) enlargement of pygidium; h) large microduct; i) enlargement of communal duct; j) enlargement of portion of duct cluster.
Specimens examined for description and diagnosis
Alleghany Co., Glade Creek, North Carolina, November 22, 2019 on Abies fraseri, A. Bartlett, 5 2nd ♀, 5 2nd ♂, 10 ad ♀ (2019-6449), Alleghany Co., Laurel Springs, North Carolina, December 8, 2020 on Abies fraseri, L. Milton, 10 ad ♀ (2020-4778).
Other specimens examined from USNM
Japan, Kobe, Arboretum, May 8, 2006, on Tsuga ? sieboldii, S. Lyon 7 2nd ♀ (0606537). United States, Connecticut, Danbury, September 7, 1944, on hemlock, S.W. Bromley 3 1st ♀ (JOH 07-77); Connecticut, Fairfield Co., New Canaan, November 3, 1950, on Nordman fir, S.W. Bromley 1 1st ♀, 18 2nd ♀, 20 ad ♀. New York, Nassau Co., Oyster Bay, May 17, 1947, on hemlock, B.F. Maker 2 1st ♂ (JOH 10-77); New York, Suffolk Co., Brookhaven, November 25, 1985, on leaves of hemlock, T. Kowalsick (ek-01-86); Pennsylvania, Radnor, July 26, 1946, on hemlock, S.W. Bromley 1 1st ♀ (JOH 08-77).
Fiorinia fioriniae
Field characteristics
First-instar exuviae overlapping second-instar exuviae. Without indentation formed between attachment of first- and second-instar exuviae. Second-instar exuviae oval, convex marginally; yellow to light brown; longitudinal ridge conspicuous. Posterior end of adult female within second-instar exuviae rounded. Heavily infested leaves with slight white secretion.
First-instar nymph
Described in
Second-instar female
Median lobes broad, equal to or wider than medial lobule of second lobe, projecting ca. same or slightly less than medial lobule of second lobes. With five pairs of marginal macroducts. Swelling of body margin adjacent to macroduct usually rounded. With four large gland spines on margin of each side of body from abdominal segments II–V; usually without small gland spine on each side of abdominal segment VI; with small gland spines on margin or submargin of abdominal segment I. With three microducts on each side of head. Longitudinal line of microducts present submarginally on venter of abdominal segments II–VI, normally with one microduct on each side of each segment. Cicatrices absent.
Second-instar male
Submargin of abdominal segments II–VI with scattered small-sized macroducts, not in clusters; communal ducts absent. Medial longitudinal line of microducts absent. Cluster of small microducts with sclerotized orifice laterad of anterior spiracle absent. Fewer than five gland spines on each side of body between anterior and posterior spiracles. Antennae each with several enlarged setae.
Notes
The single specimen collected with identified adult females of this species is unusual and may not be the second-instar male of this species. U.S. populations of Fiorinia fioriniae have been reported to be parthenogenetic (
Florida collection records
Brevard Co., Melbourne, February 22, 1984, on Phoradendron sp., F.A. Smith (1984-2933, 3014) (2 slides); Brevard Co., Sharpes, January 19, 1972, on Callistemon sp., H.C. Levan (1972-005–008) (4 slides); Broward Co., Dania, January 4, 1966, on Howea sp., J.W. Shirah (1966-7369); Broward Co., Dania, June 26, 1981, on Mangifera indica, M. McDonald (1981-1606) (2 slides); Broward Co., Dania, August 24, 2011, on Persea sp., G. Azone (2011-5990); Broward Co., Davie, April 3, 1962, on Ilex sp., D.P. McLean (1962-2896) (2 slides); Broward Co., Davie, October 12, 1978, on Camellia sp., R. Gaskalla (1978-2879) (2 slides); Broward Co., Coral Springs, October 5, 2011, on Persea americana, L. Charlton (2011-7789) (2 slides); Broward Co., Fort Lauderdale, February 5, 1968, on Callistemon sp., D.C. Clinton (1968-2883) (2 slides); Broward Co., Fort Lauderdale, February 6, 1970, on Callistemon sp., D.C. Clinton (1970-2878) (2 slides); Broward Co., Fort Lauderdale, May 2, 1974, on Sabal sp., J.A. Reinert (1974-2915) (2 slides); Broward Co., Fort Lauderdale, November 13, 1979, on Callistemon citernus, K. Tyson (1979-7500) (2 slides); Broward Co., Fort Lauderdale, February 25, 1988, on Howea forsteriana, J. McCluskie (1985-2925) (2 slides); Broward Co., Fort Lauderdale, February 25, 1988, on Manilkara roxburghiana, J. Hickey (1988-005); Broward Co., Fort Lauderdale, January 8, 1984, on Manilkara roxburghiana, J. Hickey (1984-3287) (3 slides); Broward Co., Fort Lauderdale, December 26, 2003, on Leucospermum sp., G. Farina (2003-6693); Broward Co., Fort Lauderdale, June 4, 2004, on Laurus nobilis, F.W. Howard (2004-4142) (3 slides); Broward Co., Hollywood, February 19, 1979, on Persea americana, R. Gaskalla (1978-3264) (2 slides); Broward Co., Hollywood, June 1986, on Howea forsteriana, D. Fenster (1986-008) (3 slides); Broward Co., Hollywood, December 5, 1997, on Ravenea rivularis, M.S. Quintanilla (1997-2912) (3 slides); Broward Co., North Lauderdale, May 28, 1981, on Dicytosperma album, D. Clinton and J. Aubry (1981-2895, 2904) (2 slides); Broward Co., Tamarac, March 21, 2012, on Persea americana, C. Millan (2012-1986); Broward Co., on unknown host, June 4, 2004, on Laurus nobilis, F.W. Howard (2004-4142-301); Charlotte Co., Punta Gorda, August 9, 2007, on Camellia japonica, D. Renz (2007-5759); Collier Co., Naples, August 28, 2013, on Palmae, R. Nanneman (2013-6323); Duval Co., Nocatee, April 4, 1978, on Persea americana, L.J. Chambliss (1978-3288) (2 slides); Escambia Co., Pensacola, November 3, 1988, on Prunus angustifolia, G. Corbitt and R. Burns (1988-2899); Glades Co., Moore Haven, October 4, 2006, on Celtis laevigata, L. Richards (2006-7213); Hendry Co., Devils Garden, November 20, 2014, on Persea palustris, M. Terrell (2014-788) (2 slides); Highlands Co., April 28, 1975, on Camellia sp., R.F. Denno, J.A. Davidson, D.R. Miller (1975-2886); Highlands Co., on unknown host, July 24, 1987 on Persea borbonia, R. Payne (1987-2988) (2 slides); Highlands Co., Lake Placid, November 1970, on Camellia sp. J.A. Weidhaas (1970-3265, 3854) (2 slides); Hillsborough Co., Sun City, November 14, 1994, on Phoradendron leucarpum, M. Runnals (1994-2918, 3924) (2 slides); Hillsborough Co., Tampa, April 10, 1964, on Sabal sp., S.A. Fuller (1964-2901); Hillsborough Co., Tampa, March 25, 1983, on Hedera sp., E.R. Simmons (1983-2998, 3943) (2 slides); Miami-Dade Co., Homestead, June 9, 1979, on Manilkara roxburghiana, P. Chobrda (1979-011); Indian River Co., Vero Beach, December 16, 1970, on Callistemon sp., R.H. Kendrick (1970-012–016) (5 slides); Lake Co., Tavares, September 9, 2012, on Hedera sp., M. Sellers (2012-6901); Lee Co., Sanibel Island, April 4, 1978, on Zamia sp., R. Driggers (1978-2936); Leon Co., Tallahassee, February 3, 1916, on Camellia sp. A.C.M. (1916-017); Leon Co., Tallahassee, February 3, 1916, on Camellia sp., A.C.M. (1916-2850) (4 slides); Leon Co., Tallahassee, October 31, 1919, on Camellia sp., P.F. Robertson (1919-021); Levy Co., Bronson, January 4, 2011, on Camellia sp., W. Bailey (2011-29); Lucie Co., Fort Pierce, January 16, 1980, on Dracaena sp., E.W. Campbell (1980-2944); Lucie Co., Lakewood Park, July 16, 1980, on Acoelorrhaphe wrightii, E.W. Campbell (1980-483, 2937) (2 slides); Madison Co., Greenville, March 12, 1993, Ilex sp., F. Bennett (1993-2903); Manatee Co., Oneco, December 17, 1987, on Callistemon sp., A. Waters (1987-014); Manatee Co., Snead Island, April 3, 1991, on Schinus sp., Runnals M. (1991-2922); Manatee Co., Snead Island, April 3, 1991, on Schinus sp. Runnals M. (1991-006–007) (2 slides); Marion Co., Weirsdale, December 22, 1985, on Hedera canarensis, F.J. McHenry (1985-2832, 2894) (2 slides); Martin Co., Hobe Sound, April 21, 1980, on Persea americana, E.W. Campbell (1980-2959); Martin Co., Hobe Sound, June 9, 1981, on Callistemon viminalis, S. Hakala (1981-418); Martin Co., Jensen Beach, September 27, 1978, on Dictyosperma sp., E.W. Campbell (1978-7502) (2 slides); Martin Co., Palm City, February 8, 2012, on Magnolia virginiana, L. West (2012-833) (2 slides); Martin Co., Palm City, October 17, 2012, on Persea palustris, L. West (2012-7964) (2 slides); Martin Co., Palm City, September 1, 1977, on Magnolia sp., E.W. Campbell (1977-2890, 2884) (2 slides); Martin Co., Stuart, January 31, 1978, on Eugenia sp., E.W. Campbell (1978-0955, 3283) (2 slides); Martin Co., Stuart, November 17, 1978, on Myrica sp., E.W. Campbell (1978-021); Miami-Dade Co., Big Cypress National Preserve, February 16, 1978, on Magnolia sp., A. Harmon and D. Martinelli (1978-2910); Miami-Dade Co., Big Cypress National Preserve, February 16, 1978, on Magnolia virginiana, A. Hamon and D. Martinelli (1978-006–007) (2 slides); Miami-Dade Co., Coral Gables, August 13, 2010, on Gymnanthes lucida, K. Griffiths (2010-4926); Miami-Dade Co., Florida City, November 19, 1986, on Mimusops roxburghiana, L.D. Howerton (1986-962); Miami-Dade Co., Florida City, November 19, 1986, on Mimusops roxburghiana, L.D. Howerton (1986-2914, 2980) (2 slides); Miami-Dade Co., Hialeah, March 28, 1979, on Callistemon viminalis, D. Stocks (1979-3382, 3869) (2 slides); Miami-Dade Co., Hialeah, January 1, 1980, on Callistemon sp., D. Stocks and W. James (1980-2952); Miami-Dade Co., Homestead, January 24, 1962, on Macadamia sp., R.J. McMillan (1962-3276) (2 slides); Miami-Dade Co., Homestead, October 16, 1962, on Melaleuca sp., J.H. Knowles (1962-2897) (2 slides); Miami-Dade Co., Homestead, February 2, 1969, on Persea sp., D.O. Wolfenbarger (1969-491) (2 slides); Miami-Dade Co., Homestead, March 28, 1969, on Persea sp. D.O. Wolfenbarger (1969-026–031) (4 slides); Miami-Dade Co., Homestead, February 2, 1978, on Hedera sp., W.E. Wyles (1978-488); Miami-Dade Co., Homestead, February 27, 1978, on Hedera sp., W.E. Wyles (1978-7522) (5 slides); Miami-Dade Co., Homestead, June 8, 1979, on Manilkara roxburghiana, P. Chobrda (1979-0146) (4 slides); Miami-Dade Co., Homestead October 3, 1979, on Persea americana, W.E. Wyles (1979-2928); Miami-Dade Co., Homestead, September 11, 2007, on Persea americana, B. Saunders (2007-6958); Miami-Dade Co., Homestead, July 31, 2018, on Gymnanthes lucida, W. Mazuk (2018-4092) (2 slides); Miami-Dade Co., Kendall, February 24, 1989, on Camellia sp., W. Francillon (1989-2855) (2 slides); Miami-Dade Co., Miami, April 22, 1966, on Chamaedora sp., C.F. Dowling (1966-2876) (2 slides); Miami-Dade Co., Miami, June 6, 1967, on Chamaerops sp., J.S. Sloan (1967-2907) (2 slides); Miami-Dade Co., Miami, October 26, 1967 on Callistemon viminalis, J.F. Dillon (1967-3305) (2 slides); Miami-Dade Co., Miami, October 26, 1967, on Callistemon viminalis, J.F. Dillon (1967-038); Miami-Dade Co., Miami, March 3, 1969, on Howea sp., J.F. Dillon (1969-493, 3060) (2 slides); Miami-Dade Co., Miami, March 7, 1969, on Howea sp., J.F. Dillon (1969-3267) (4 slides); Miami-Dade Co., Miami, September 5, 1969, on Macadamia ternifolia, J.F. Dillon (1969-045–047) (3 slides); Miami-Dade Co., Miami, January 22, 1975, on Callistemon viminalis, D. Sager (1975-2885) (3 slides); Miami-Dade Co., Miami, July 27, 1978, on Mangifera sp., M. Corman (1978-3308) (2 slides); Miami-Dade Co., Miami, April 5, 1979, on Kigelia pinnata, P. Chobrda (1979-2913); Miami-Dade Co., Miami, November 5, 1979, on Callistemon sp., H. VonWald (1979-3003); Miami-Dade Co., Miami, April 1, 1980, on Callistemon viminalis, G. Webster and E. Pena (1980-014); Miami-Dade Co., Miami, April 1, 1980, on Callistemon viminalis, G. Webster and E. Pena (1980-2881); Miami-Dade Co., Miami, April 2, 1980, on Santalum album, H. Von Wald and C. Dowling (1980-2935) (2 slides); Miami-Dade Co., Miami, April 3, 1981, on Persea americana, K. Martin (1981-016 –020) (5 slides); Miami-Dade Co., Miami, March 3, 1981, on Macadamia ternifolia, W. James (1981-2934) (2 slides); Miami-Dade Co., Miami, March 25, 1981, on Macadamia ternifolia, W. James (1981-2984); Miami-Dade Co., Miami, February 10, 1982, on Diospyros lotus, H. VonWald (1982-017–019) (3 slides); Miami-Dade Co., Miami, April 8, 1982, on Eucalyptus sp., P. Perun (1982-223, 2853) (2 slides); Miami-Dade Co., Miami, April 8, 1982, on Eucalyptus sp., P. Perun (1982-2853); Miami-Dade Co., Miami, November 15, 1985, on Howea forsteriana, D. Chalot (1985-3009) (2 slides); Miami-Dade Co., Miami, September 19, 29, 1986, on Persea americana, D. Storch (1986-021–023, 3091) (3 slides); Miami-Dade Co., Miami, January 17, 2001, Manilkara roxburghiana, E. Putland (2001-189) (2 slides); Miami-Dade Co., Miami, March 14, 2002, on Persea americana, L. Davis (2002-870); Miami-Dade Co., Miami, August 14, 2007, on Manilkara roxburghiana, O. Garcia (2002-5912); Miami-Dade Co., Miami, May 11, 2012, on Laurus nobilis, M. Figueroa (2012-3720) (2 slides); Miami-Dade Co., North Beach, January 20, 1981, on Amyris elemfera, E.W. Campbell and R. Kendrick (1981-2947) (2 slides); Miami-Dade Co., Opa Locka, October 5, 1977, on Callistemon sp., M. Corman (1977-3268, 3278) (2 slides); Miami-Dade Co., Opa Locka, May 22, 1978, on Persea americana, J. Hilderbrandt (1978-3313) (2 slides); Miami-Dade Co., West Miami, November 22, 1977, on Camellia sp., D. Martinelli (1977-2888, 2898) (2 slides); Monroe Co., Little Torch Key, April 10, 2018, on Bidens alba, P. Corogin, J. Hayden, E. Talamas, B. Danner, J. Farnum (2018-1780); Orange Co., Apopka, Jan 10, 2001, on Ravenea rivularis, K. Gonzalez (2001-116) (2 slides); Orange Co., Belle Isle, January 20, 2006, on Garcinia livingstonei, T. Williams (2006-268); Orange Co., Orlando, February 2013, on Theaceae, A. Puppelo (2013-1127); Orange Co., Orlando, May 27, 2008, on Magnolia virginiana, A. Puppelo (2008-3337); Orange Co., Winter Garden, October 31, 2008, on Machilus thunbergii, G. Warden (2008-7478); Orange Co., Zellwood, March 7, 2019, Laurus nobilis, K. Gonzalez (2019-1006) (2 slides); Palm Beach Co., Boca Raton, May 19, 1982, on Chamaerops humilis, D.C. Clinton (1982-487) (3 slides); Palm Beach Co., Boynton Beach, October 10, 1973, on Ficus sp., K. Geyer (1973-3284) (3 slides); Palm Beach Co., Delray Beach, May 12, 1978, on Diospyros sp., K.C. Stolley (1978-2852) (2 slides); Palm Beach Co., Boynton Beach, June 7, 1978, on Mimusops sp., K. Stolley (1978-3307); Palm Beach Co., Boynton Beach, September 27, 1991, on Camellia japonica, E. Tannehill (1991-2906); Palm Beach Co., Boynton Beach, January 13, 1988, on Mimusops roxburghiana, D. Leone (1988-2880) (2 slides); Palm Beach Co., Boynton Beach, November 2, 1989, on Sisyrinchium solstitiale, E. Tannehill (1989-485, 3303) (2 slides); Palm Beach Co., Delray Beach, February 23, 1988, on Melaleuca sp., E. Tannehill and A. Hamon (1988-2851) Palm Beach Co., Jupiter, May 8, 2013, on Magnolia sp., L. West (2013-3217) Palm Beach Co., Lake Park, June 14, 1978, on Chrysalidocarpus lutescens, J. Bennet (1978-3567) (2 slides); Palm Beach Co., Lake Park, April 24, 1979, on Chrysalidocarpus lutescens, J.E. Bennet (1979-2920) (2 slides); Palm Beach Co., Lake Worth, March 7, 1978, on Kentia sp., J, Bennett (1978-3285); Palm Beach Co., Lake Worth, October 8, 1981, on Magnolia virginiana, J. Fellers and R. Buchholz (1981-026–027) (2 slides); Palm Beach Co., Lake Worth, July 13, 1995, on Chrysalidocarpus lutescens, Cook S.H., Clinton D.C. (1995-3024); Palm Beach Co., Lake Worth, July 13, 1995, on Chrysalidocarpus lutescens, S.H. Cook, D.C. Clinton (1995-3024); Palm Beach Co., Lake Worth, March 25, 2004, on Calophyllum inophyllum, L. Smith (2004-2103); Palm Beach Co., Pahokee, February 22, 1980, on Chrysalidocarpus lutescens, N. Miles and B. Walsh (1980-2955) (2 slides); Palm Beach Co., South Bay, November 14, 2018, on Magnolia virginiana, J. Farnum (2018-5955); Palm Beach Co., West Palm Beach, January 14, 1991, on Mangifera indica, R.T. Doll (1991-3566) (2 slides); Pinellas Co., Clearwater, January 11, 2013, on Persea palustris, W. Salway (2013-575) (2 slides); Pinellas Co., Indian Rocks, October 3, 1972, on Persea americana, K.C. Lowery (1972-2860); Pinellas Co., Largo, November 8, 1978, on Persea sp., P. Pullara (1978-3306); Pinellas Co., St. Petersburg, August 15, 1967, on Persea sp., C.K. Hickman (1967-2854) (3 slides); Pinellas Co., St. Petersburg, February 2, 2008, on Persea borbonia, M. Spearman (2004-724); Pinellas Co., St. Petersburg, May 28, 2009, on Persea borbonia, M. Spearman (2009-3632); Polk Co., Cypress Gardens, January 16, 1962, on Tetrapanax sp., J.N. Pott (1962-3266) (2 slides); Polk Co., Cypress Gardens, August 13, 1964, on Magnolia sp. W.P. Henderson (1964-2905) (2 slides); Polk Co., Lake Wales, October 25, 1962, on Ficus sp., Ralph E. Brown (1962-2857); Polk Co., Winter Haven, April 8, 1980, on Persea Americana, H.G. Schmidt (1980-2938) (2 slides); Polk Co., Winter Haven, July 26, 2018, on Laurus nobilis, J. Bryan (2018-4054); St. Lucie Co., Fort Pierce, March 23, 1978, on Paurotis sp., E.W. Campbell (1978-032–033) (2 slides); St. Lucie Co., Fort Pierce, January 17, 1979, on Persea sp., E.W. Campbell (1979-2893); St. Lucie Co., Fort Pierce, March 23, 1979, on Paurotis sp., E.W. Campbell (1979-2900); St. Lucie Co., Fort Pierce, February 24, 1984, on Bumelia tenax, K. Hibbard and E.W. Campbell (1984-492, 2948) (2 slides); St. Lucie Co., Fort Pierce, November 6, 1985, on Paurotis sp., K Hibbard and E.W. Campbell (1985-484, 2950) (2 slides); St. Lucie Co., Fort Pierce, March 13, 2003, on Phoradendron leucarpum, K. Hibbard (2003-927); St. Lucie Co., Fort Pierce, February 21, 2005, on Ilex cornuta, D. Vazquez (2005-4069); St. Lucie Co., Hutchinson, Isle, April 18, 1980, on Eugenia simpsonii, E.W. Campbell (1980-2990); St. Lucie Co., Port St. Lucie, May 17, 1978, on Persea borbonia, E.W. Campbell (1978-2858) (2 slides); St. Lucie Co., Port St. Lucie, February 20, 1980, on Persea borbonia, E.W. Campbell and R.H. Kendrick (1980-2989, 3008) (2 slides); St. Lucie Co., White City, May 30, 1980, on Persea borbonia, E.W. Campbell (1980-2961) (2 slides); Volusia Co., Allandale, March 16, 1983, on Hedera helix, J.N. Pott (1983-486) (2 slides); Volusia Co., Daytona Beach, August 16, 1984, on Howea forsteriana, J.N. Pott (1984-2994); Volusia Co., Holly Hill, March 15, 1956, on Chamaedorea sp., C.R. Roberts (1956-2877) (3 slides); Volusia Co., New Smyrna Beach, April 8, 1985, on Palm, J.N. Pott (19852942); Volusia Co., New Smyrna Beach, September 27, 1971, on Camellia sp., J.N. Pott (1971-2887, 2889) (8 slides); Walton Co., Walton, February 12, 1980, on Mangifera indica, E.W. Campbell (1980-2946) (2 slides).
Fiorinia fioriniae, second-instar male, Marion Co., Ocala, August 13, 2019 on Chamaerops humilis, T. Gordon, (2019-4546). Abbreviations: a) large microduct; b) large gland spine; c) large gland spine; d) large microduct; e) small gland c) small microduct; d) enlargement of pygidium; e) pores with S-shaped opening f) dorsal large microduct; g) dorsal large microducts; h) antennae each with several enlarged seta.
Specimens examined for description and diagnosis
Marion Co., Ocala, August 13, 2019 on Chamaerops humilis, T. Gordon, 5 2nd ♀, 5 2nd ♂, 10 ad ♀ (2019-4546).
Other material examined from USNM
Mexico: July 11, 1988, on Mangifera indica, S. Sanner 6 2nd ♀ (El Paso 032924). Peru: May 7, 1977, on Mangifera indica, R. Narkaus 5 2nd ♀ (Los Angeles 19190); August 21, 1972, on Camellia sp., E.B. Lee 1 1st ♀, 4 2nd ♀, 3 ad ♀. Portugal: Azores, August 20, 1928, on Camellia sp., C.A. Davis 1 1st ♀ (at Providence, Rhode Island). United States: California, San Diego, San Diego Zoo, August 19, 2002, D. Kellum, J.F. Miller, D.R. Miller, on Camellia sp. 3 1st ♀, 18 2nd ♀, 7 ad ♀; Georgia, Camden Co., June 14, 1969, on Ruscus sp., R.J. Beashear 1 1st ♀.
Fiorinia japonica
Field characteristics
First-instar exuviae overlapping second-instar exuviae. Without indentation or with slight indentation formed between attachment of first- and second-instar exuviae. Second-instar exuviae oval, convex marginally; medium to dark brown; longitudinal ridge inconspicuous. Posterior end of adult female within second-instar exuviae rounded. Heavily infested leaves with white secretion (Suppl. material
First-instar nymph
described in
Second-instar female
Median lobes broad, as wide as or wider than medial lobule of second lobe, projecting ca. same distance as or further than medial lobule of second lobes. With five pairs of marginal macroducts. Swelling of body margin adjacent to macroduct usually rounded. With four large gland spines on margin of each side of body from abdominal segments II–V; usually without small gland spine on each side of abdominal segment VI; with small gland spines on margin or submargin of abdominal segment I. With one microduct on each side of head. Longitudinal line of microducts present submarginally on venter of abdominal segment II or III–VI, normally with one microduct on each side of each segment. Cicatrices absent.
Second-instar male
One duct cluster on each side of body, composed of several small ducts and two communal ducts. Three longitudinal lines of microducts on venter of abdomen (one medial and two submarginal). Cluster of small microducts with sclerotized orifice laterad of anterior spiracle absent. Fewer than five gland spines on each side of body between anterior and posterior spiracles. Antennae each with one enlarged seta.
Fiorinia japonica, second-instar male Virginia, Chesterfield Co., Southside Nursey, July 27, 1974, on blue spruce, R. Sears. Abbreviations: a) antenna; b) anterior spiracle; c) large gland spine; d) large microduct; e) small microduct; f) small gland spine; g) enlargement of pygidium; h) enlargement of portion of duct cluster; i) large microduct; j) enlargement of communal duct.
Florida collection records
Fiorinia japonica has not been collected in Florida.
Specimens examined for description and diagnosis
Virginia, Chesterfield Co., Southside Nursey, July 27, 1974, on blue spruce, R. Sears, 5 2nd ♀, 5 2nd ♂.
Other material examined from USNM
China; “Hsifeushang” January 23, 1933, on Pinus sp., W.B. Wood 2 1st ♀ (JOH 55-76 F). Taiwan: Maruyama, near Taihoku, June 3, 1928, on Pinus thunbergi, R. Takahashi 1 1st ♂ (JOH 58-76); Taihoku, June 7, 1929, on Pinus sp., R. Takahashi 1 1st ♀ (JOH 54-76). United States: Virginia, Chesterfield Co., Southside Nursey, July 27, 1974, on blue spruce, R. Sears 2 1st ♀, 1 1st ♂, 5 2nd ♀, 5 2nd ♂, 4 ad ♀; Washington, D.C., August 27, 1991, on national Christmas tree, Horton 3 1st ♀, 4 2nd ♀, 3 ad ♀ (93-09742).
Fiorinia phantasma
Field characteristics
First-instar exuviae overlapping second-instar exuviae. Without indentation formed between attachment of first- and second-instar exuviae. Second-instar exuviae oval, convex marginally; light to dark brown, longitudinal ridge weakly developed. Posterior end of adult female within second-instar exuviae constricted and pointed (Suppl. material
First instar
Similar to F. fioriniae and F. proboscidaria in having gland spines on abdominal segment VI at least half as long as gland spine on segment VII. Fiorinia fioriniae and F. proboscidaria differ by having (characters in parentheses are those of P. phantasma): pattern of derm surrounding large duct on head serpentine (globular); inner apex of large duct on head flat (rounded or mushroom like).
Second-instar female
Median lobes broad, as wide as or slightly narrower than medial lobule of second lobe, projecting ca. same amount or slightly less than medial lobule of second lobes. With five pairs of marginal macroducts. Swelling of body margin adjacent to macroduct usually pointed. With three large gland spines on margin of each side of body from abdominal segments II–IV, without gland spine on abdominal segment VI; with small gland spines on margin or submargin of abdominal segment I. With three microducts on each side of head. Longitudinal line of microducts absent submarginally on venter of abdomen. Cicatrices absent.
Second-instar male
One duct cluster on each side of body, composed of several small ducts and one communal duct. Five longitudinal lines of microducts on venter of abdomen (one medial, two mediolateral, and two submarginal), medial line sometimes incomplete. Cluster of small microducts with sclerotized orifice laterad of anterior spiracle absent. Fewer than five gland spines on each side of body between anterior and posterior spiracles. Antennae each with one enlarged seta.
Adult female
Body tapering at segment III to narrow pygidium. With three or four pairs of dorsal macroducts on each side of body, ducts similar in shape and size to microducts. Projection between antennae with many spicules. Antennae close together, with distinct projection.
Florida collection records
Miami-Dade Co., Miami, March 1, 2018, on Phoenix canariensis, Olga Garcia (2018-789) (3 slides); Miami-Dade Co., Coral Gables, April 2, 2018, on Phoenix sp., J. Farnum (2018-1499) (2 slides); Miami-Dade Co., Pinecrest, April 2, 2018, on Phoenix sp., J. Farnum (2018-1487, 1489, 1491, 1492, 1496, 1500, 1503, 1504, 1524, 1525) (20 slides); Miami-Dade Co., Palmetto Bay, April 2, 2018, on Phoenix sp., J. Farnum (2018-1488, 1493, 1498, 1501) (8 slides); Miami-Dade Co., Pinecrest, May 22, 2018, on Phoenix sp., J. Farnum and J. Vergel (2018-2780, 2785, 2796) (6 slides); Palm Beach Co., Boynton Beach, January 23, on Cocos nucifera, L. Smith (2018-304) (2 slides); Miami-Dade Co., Coral Gables, Miami-Dade Co., Palmetto Bay, May 22, 2018, on Phoenix sp., J. Farnum and J. Vergel (2018-2783, 2788, 2790) (6 slides); May 22, 2018, on Dypsis lutescens, J. Miller, H. Mayer, M.Z.
Specimens examined for description and diagnosis
Palm Beach Co., Boynton Beach, October 30, 2019, on Wodyetia bifurcata, L. Smith, 5 1st (2019-5998); Palm Beach Co., Boynton Beach, April 6, 2020, on Ligustrum japonicum, L. Smith, 10 ad ♀ (2020-1365); Palm Beach Co., Boynton Beach, November 6, 2019, on Wodyetia bifurcata, 5 2nd ♀ (2019-6182); Philippines, June 28, 1996 on Plumeria sp., 2nd ♀ (SF023635); Miami-Dade Co., Miami, November 9, 2019, on Palmae, O. Garcia, 5 2nd ♂ (2019-6149); Palm Beach Co., Boca Raton, December 29, 2020, on Phoenix canariensis, L. Smith, 10 ad ♀ (2020-4958).
Fiorinia phantasma, First-instar nymph, Palm Beach Co., Boynton Beach, October 30, 2019, on Wodyetia bifurcata, L. Smith, (2019-5998). Abbreviations: a) large dorsal duct on head; b) small microduct; c) gland spine abdominal segment II with small projection; d) gland spine at abdominal segment VI with long projection; e) gland spine at abdominal segment VII with long projection.
Fiorinia phantasma (Cockerell & Robinson); adult female, Palm Beach Co., Boynton Beach, April 6, 2020, on Ligustrum japonicum, L. Smith (2020-1365). Abbreviations: a) detail of antennae and inter-antennal process; b) detail of anterior spiracle; c) microducts; d) marginal duct tubercles; e) marginal microduct; f) detail of pygidium; g) marginal macroduct on pygidium.
Fiorinia phantasma, second-instar female and second-instar female shed shin, Palm Beach Co., Boynton Beach, November 6, 2019, on Wodyetia bifurcata, (2019-6182). Abbreviations: a) antenna; b) anterior spiracle; c) microduct with sclerotized orifice; g) old second-instar female, Philippines, June 28, 1996 on Plumeria sp., 2nd ♀ (SF023635).
Fiorinia phantasma, second-instar male, Miami-Dade Co., Miami, November 9, 2019, on Palmae, O. Garcia, (2019-6149). Abbreviations: a) antenna; b) large microduct; c) small microduct; d) large gland spine; e) small macroduct; f) enlargement of communal duct ; g) enlargement of pygidium; h) large microduct; j) enlargement of part of duct cluster.
Other specimens examined from USNM
Grenada, Calivingy Island, March 2012, on Phoenix dactylifera, S.W. Evans (E-2012-2099); Guam, Tamuning, June 4, 1984, on Cocos nucifera, R. Muniappan; Hawaii: Oahu, Kapahulu area, March 27, 2009, on Ligustrum sp., M. Ramadan (0904651); Hawaii, Hilo ?, date ?, on Pittosporum sp., B. Kumashiro 5 2nd ♀,5 2nd ♀; Philippines, April 7, 1965, on Cocos nucifera, J.I. Mason; Philippines, November 15, 1971, on palm leaf, R.F. Goodall (Seattle 8910); Philippines, March 13, 1975, on Mangifera indica, M. Yoshinaga (Hawaii 28847); Philippines, August 26, 1975, on Cocos nucifera, A. Buchanan (LA 015303); Philippines, November 30, 1975, on palm leaf, Ozuka & Richardson (Hawaii 32858); Philippines, June 8, 1977, on palm leaf, Tamiya (Hawaii 39130); Philippines, August 17, 1977, on Areca sp., J. Sato (Honolulu 42721); Philippines, October 19, 1978, on Areca sp., Takeda, (Honolulu 43108); Philippines, November 4, 1978, on palm leaf, Jodoi, (Honolulu 42704); Philippines, May 26, 1981, on Tamarindus indica, D.O. Wienhee & F.G. Walisen (Seattle 17304); Philippines, May 12, 1985, on Philippines, Clausenia anisum, C. Dollopf (Chicago 009312); Philippines, June 28, 1996, on Plumeria sp. (SF 023635); Philippines, October 19, 1978, on Areca sp., Takeda (Honolulu 43108); Taiwan, May 10, 1981, on leaf, J.L. Levitt (Seattle 17328); Taiwan, April 8, 1988, on Ficus sp., V. McDonald (JFK 100609); Thailand, August 31, 1982, on leaf, G. Hinsdale (Anchorage 017131); Thailand, July 19, 1985, on Murraya koenigii, J. Alabu (LA 052785); Thailand, November 15,1987, on Areca sp., J. Elridge (Atlanta 003471); Thailand, March 20, 2003, on Arecaceae, F. Hadded; Thailand, March 27, 2006, on palm, M. Hanzlik (ANC 060060); Vietnam, April 26, 2007, on Cocos sp., A. Coronel (LA 207977 CA); Vietnam, October 26, 2012, on unknown host, D. Gregory (SF 1301449).
Notes
We examined four paratype slides of F. coronata Williams & Watson from Guadalcanal, Solomon Islands deposited in the
We also examined a paratype slide of F. phantasma in the same collection, but it is in such poor condition that only half of the pygidium is useful for diagnosis. It is impossible to even find the posterior spiracles, let alone microducts between them. In addition, the holotype of F. phantasma deposited in The Natural History Museum, London (NHMUK) was loaned to and examined by one of us (DL). It also was in poor condition; microducts close to anterior and posterior spiracles and in prepygidial abdominal segments were not visible. All of the examined specimens of F. phantasma that were in good condition had easily discernable microducts between the posterior spiracles.
Fiorinia pinicola
Field characteristics
First-instar exuviae overlapping second-instar exuviae. With indentation formed between attachment of first- and second-instar exuviae. Second-instar exuviae oval, convex marginally; medium to dark brown; longitudinal ridge conspicuous. Posterior end of adult female within second-instar exuviae rounded. Heavily infested leaves with white secretion (Suppl. material
First-instar nymph
Described in
Second-instar female
Median lobes broad, as wide as or wider than medial lobule of second lobe, projecting ca. same amount as medial lobule of second lobes. With five pairs of marginal macroducts. Swelling of body margin adjacent to macroduct usually pointed. With four large gland spines on margin of each side of body from abdominal segments II–V; usually without small gland spine on each side of abdominal segment VI; with small gland spines on margin or submargin of abdominal segment I. With one microduct on each side of head. Longitudinal line of microducts present submarginally on venter of abdominal segments II–VI, normally with one microduct on each side of each segment. Cicatrices absent.
Fiorinia pinicola, second-instar female, Hong Kong, December 1895, on Pinus sinensis, A. Koebele (1529) mounted from type material. Abbreviations: a) antenna; b) anterior spiracle; c) small microduct with sclerotized orifice d) small microduct e) enlargement of pygidium. Note the blank blotches on the body margin of the pygidium enlargement.
Fiorinia pinicola, second-instar male, Hong Kong, China, December 1895, on Pinus sinensis, A. Koebele (1529) mounted from type material. Abbreviations: a) antenna; b) large gland spine; c) large microduct; d) small microduct; e) small gland spine; f) enlargement of pygidium; g) enlargement of communal duct ; h) small microduct; i) large microduct; j) enlargement of part of duct cluster.
Notes
We have been unable to find characters that consistently separate second-instar females of F. japonica and F. pinicola. The swelling of the body adjacent to the abdominal macroducts is usually pointed in F. pinicola and is usually rounded in F. japonica, but we have too few specimens to understand the possible variation in this character.
Second-instar male
One duct cluster on each side of body, composed of several small ducts and two communal ducts. Three longitudinal lines of microducts on venter of abdomen (one medial and two submarginal). Cluster of small microducts with sclerotized orifice laterad of anterior spiracle absent. Five or more gland spines on each side of body between anterior and posterior spiracles. Antennae each with one enlarged seta.
Specimens examined for description and diagnosis
China, Hong Kong, December 1895, on Pinus sinensis, A. Koebele 2nd ♀, ♂ (1529); United States, California, Los Angeles Co., Los Angeles, August 11, 2020, on Podocarpus macrophyllus, N. Ellenrieder, 3 2nd ♀, 2 ad ♀ (2020-3174).
Other specimens examined from USNM
China, Hong Kong, December 1895, on Pinus sinensis, A. Koebele (1529) mounted from type material, 2 2nd ♀, 2 2nd ♂, 2 ad ♀; Japan, Yokohama, Yamashita-cho, October 15, 1941, on Pittosporum tobira, K. Soto 1 1st ♀, 2 2nd ♀ (Yokohama 199); Japan, November 2, 1977, on Podocarpus sp., 1 1st ♀, 7 ad ♀; United States, California, Orange Co., October 2002, on Pittosporum sp., H. Mitchell 1 1st ♂ embryo, 1 1st ♀ embryo, 7 2nd ♀, 6 ad ♀.
Fiorinia proboscidaria
Field characteristics
First-instar exuviae overlapping second-instar exuviae. Without indentation formed between attachment of first- and second-instar exuviae. Second-instar exuviae oval, convex marginally or parallel sided; light to medium dark brown; longitudinal ridge conspicuous and thick. Posterior end of adult female within second-instar exuviae rounded. Heavily infested leaves with white secretion.
First-instar nymph
Similar to F. fioriniae and F. phantasma in having gland spines on abdominal segment VI at least half as long as gland spine on segment VII. Fiorinia fioriniae differs by having (characters in parentheses are those of P. proboscidaria): inner apex of large duct on head flat (rounded or mushroom like). Fiorinia phantasma differs by having (characters in parentheses are those of F. proboscidaria): pattern of derm surrounding large duct on head globular (serpentine); inner apex of large duct on head rounded or mushroom like (flat).
Adult female
Process between antennae without spicules, often clubbed. Head conical. Antennae close together. Macroducts usually 3–5 on each side of pygidium, thin, longer than wide, resembling microducts. Gland spines barely projecting from body margin. Gland tubercles nearly continuous along body margin from head to abdominal segment III. Microducts in medial areas of prepygidial segments dorsally and ventrally. Lateral margin of head with cluster of circular tubercles possibly representing eye.
Notes
There are a number of species with processes between the antennae;
Second-instar female
Median lobes broad, as wide as or wider than medial lobule of second lobe, projecting ca. same amount or slightly less than medial lobule of second lobes. With four pairs of marginal macroducts. Swelling of body margin adjacent to macroduct usually rounded. With three large gland spines on margin of each side of body from abdominal segments II–IV; usually with small gland spine on each side of abdominal segments V and VI; without small gland spines on abdominal segment I. With three microducts on each side of head. Longitudinal line of microducts present submarginally on venter of abdominal segments II–VI, normally with 1–5 microducts on each side of each segment. Small lobular projections on anterior of head sometimes present. Cicatrix present on dorsal submargin of abdominal segment I.
Second-instar male
Two duct clusters on each side of body, anterior cluster without communal duct, posterior cluster composed of communal duct without associated smaller ducts. Normally, three longitudinal lines of microducts on venter of abdomen (one medial and two submarginal) occasionally with two medially forming four longitudinal lines. Without cluster of small microducts with sclerotized orifice laterad of anterior spiracle. Fewer than five gland spines on each side of body between anterior and posterior spiracles. Antennae each with one enlarged seta.
Fiorinia proboscidaria, First-instar nymph, Putnam Co., Crescent City, October 2, 2019, on Citrus sp., D. Rigby, M. Cain, (2018-5548). Abbreviations: a) large duct on head; b) dorsal microduct; c) short gland spine on abdominal segment II; d) long gland spine on abdominal segment VI; e) long gland spine on abdominal segment VII.
Fiorinia proboscidaria, adult female, Hillsborough Co., Tampa, December 16, 2013, on Citrus sp., J. Hoffman (2013-9087). Abbreviations: a) conical head with protrusion, antennae: b) anterior spiracle; c) gland tubercle; d) gland spine with small dermal protrusion; e) small microduct; f) enlargement of pygidium; g) microduct; h) circular tubercles in invagination.
Fiorinia proboscidaria, second-instar male, Putnam Co., Crescent City, October 2, 2019, on Citrus sp., D. Rigby, M. Cain, (2018-5548). Abbreviations: a) small microduct; b) large gland spine; c) enlargement of pygidium; d) small macroduct; e) large microduct; f) enlargement of part of duct cluster; g) enlargement of communal duct.
Florida collection records
Hillsborough Co., Tampa, December 16, 2013, on Citrus sp., J. Hoffman (2013-9087) (7 slides); Hillsborough Co., Tampa, November 19, 2014, on Citrus sp., M. Briceno (2014-859) (6 slides); Hillsborough Co., Tampa, October 23, 2014, on Citrus sp., M. Briceno (2014-7431) (2 slides); Hillsborough Co., Tampa, October 30, 2014, on Citrus sp., M. Briceno (2014-7574) (7 Slides); Hillsborough Co., Valrico, September 17, 2018, on Citrus × paradisi, P. Barker (2018-4907) (2 slides); Putnam Co., Palatka, October 25, 2018, on Ilex cornuta, M. Cain, T. Wright, and C. Hall (2018-5664) (2 slides); Santa Rosa Co., Gulf Breeze, January 3, 2014, on Citrus sp., M. Anderson (2014-46) (3 slides).
Specimens examined for description and diagnosis
Flagler Co., Palm Coast, June 18, 2020, on Citrus sp., M. Cain, 5 2nd ♀, (2020-2353); Hillsborough Co., Tampa, December 16, 2013, on Citrus sp., J. Hoffman 5 ad ♀ (2013-9087); Manatee Co., Bradenton, January 6, 2021, on Citrus sp., P. Kumar, 10 ad ♀ (2021-67); Pinellas Co., Palm Harbor, September 16, 2019, on Citrus sp. B. Rose, 2 ad ♀ (2019-5124); Putnam Co., Crescent City, October 2, 2019, on Citrus sp., D. Rigby, M. Cain, 5 1st (2018-5548); Putnam Co., Crescent City, October 2, 2019, on Citrus sp., D. Rigby, M. Cain, 5 2nd ♂ (2018-5548).
Material examined from USNM
China, Hong Kong, September 2, 1980, on Podocarpus sp., J. Dooley 1 2nd ♂, 1 prepupa, 4 ad ♀ (Los Angeles 25002). Martinique, February 8, 2000, on Citrus aurantifolia, K. Stewart 1 2nd ♀, 1 ad ♀ (St. Thomas 010770).
Fiorinia theae
Field characteristics
First-instar exuviae overlapping second-instar exuviae. Without indentation between attachment of first- and second-instar exuviae. Second-instar exuviae oval, convex marginally; light gray to nearly black; longitudinal ridge conspicuous. Posterior end of adult female within second-instar exuviae rounded. Heavily infested leaves with extensive white secretion (Suppl. material
First-instar nymph
Described in
Second-instar female
Median lobes broad, equal to or wider than width of medial lobule of second lobe, projecting ca. same amount or slightly less than medial lobule of second lobes. With four pairs of marginal macroducts. Swelling of body margin adjacent to macroduct usually rounded. With three large gland spines on margin of each side of body from abdominal segments II–IV; usually with small gland spine on each side of abdominal segments V and VI; without small gland spines on abdominal segment I. With three microducts on each side of head. Longitudinal line of microducts present submarginally on venter of abdominal segments II–VI, normally with 1–5 microducts on each side of each segment. Small lobular projections anteriorly on head sometimes present. Cicatrix present on dorsal submargin of abdominal segment I.
Notes
We have been unable to find characters that consistently separate second-instar females of F. proboscidaria and F. theae.
Second-instar male
Two duct clusters on each side of body, anterior cluster without communal duct, posterior cluster composed of two communal ducts without associated smaller ducts. Five longitudinal lines of microducts on venter of abdomen (one medial, two mediolateral and two submarginal). Without cluster of small microducts with sclerotized orifice laterad of anterior spiracle (sometimes with one duct present). Fewer than five gland spines on each side of body between anterior and posterior spiracles. Antennae each with one enlarged seta. Some specimens with small protrusions that are remnants of legs.
Florida collection records
Alachua Co., Alachua, October 1, 2012, Camellia sp., C. Jones, (2012-7479) (2 slides); Alachua Co., Gainesville, February 10, 1965, on Camellia japonica, A.E. Graham (1965-0345); Alachua Co., Gainesville, October 16, 1973, on Citrus mitis, F. Collins (1973-3065); Alachua Co., Gainesville, October 25, 1979, on Citrus sp., R.I. Sailer (1979-1513); Alachua Co., Gainesville, March 20, 1991, on Ilex sp., F. Bennet (1991-2953); Alachua Co., Gainesville, March 20, 1991, on Ilex sp., F. Bennett (1991-001–002) (2 slides); Alachua Co., Gainesville, July 19, 1991, on Ilex sp., F. Bennett (1991-0383) (3 Slides); Alachua Co., Gainesville, December 3, 1991, on Ilex sp., F. Bennet (1991-002–003) (2 slides); Alachua Co., Gainesville, October 16, 1992, Ilex sp., F. Bennett (1992-001); Alachua Co., Gainesville, April 15, 1999, on Ilex sp., D. Strosnider (1999-383) (2 slides); Alachua Co., Gainesville, July 2011, on Ilex cornuta, Shirley Vogel (2011-4847); Alachua Co., Gainesville, February 2012, on Camellia sasanqua, D. Feiber (2012-1006) (3 slides); Alachua Co., Gainesville, January 8, 2013, on Illicium floridanum, M. Frank (2013-102) (2 slides); Alachua Co., Gainesville, December 1, 2013, on Camellia sp., T. Harris (2013-8692); Alachua Co., Hawthorne, February 26, 1971, on Aucuba japonica, E.W. Holder (1971-2976) (5 slides); Baker Co., Macclenny, March 14, 1968, on Ilex latifolia, H.W. Collins (1968-351) (2 slides); Baker Co., Macclenny, October 21, 1975, on Ilex cornuta, C. Webb (1975-411); Bay Co., Panama City, March 8, 1978, on Euonymussp., A.E. Graham (1978-003); Brevard Co., Grant, June 15, 1962, on Camellia sp., H.C. Levan (1962-0388); Citrus Co., Hernando, August 15, 1979, on Ilex cornuta, R.H. Phillips (1979-1604) (2 slides); Collier Co., Naples, November 13, 2012, on Ilex sp., S. Krueger (2012-8615) (2 slides); Collier Co., Naples, November 13, 2012, on Ilex sp., S. Krueger (2012-8618) (2 slides); Dixie Co., Old Town, November 19, 1979, on Euonymus sp., F. McHenry (1979-1664) (3 slides); Dixie Co., Suwannee, May 22, 1978, on Citrus nobilis, A.E. Graham and A. Hamon (1978-1601) (3 slides); Duval Co., Jacksonville, February 17, 1981, on Ilex cassine, H. Collins (1981-0352) (2 slides); Duval Co., Jacksonville, October 13, 1981, on Citrus sinensis, G. Virgona (1981-0460, 3056) (3 slides); Duval Co., Jacksonville, January 25, 2005, on Camellia japonica, J. Smith (2005-464); Duval Co., Jacksonville, October 26, 2010, on Ilex vomitoria, J. Brambila (2010-6593); Duval Co., Jacksonville, October 15, 2012, on Camellia sp., K. Theriault (2012-7841–7842) (4 slides); Duval Co., Jacksonville, October 25, 2012, on Citrus sp., K.Coffey, Lisa Hassell (2012-8102) (2 slides); Duval Co., Jacksonville, April 15, 2013, on Ilex sp., K. Theriault (2013-2501) (2 slides); Duval Co., Jacksonville, March 26, 2014, on Camellia sasanqua, L. Hassel (2014-2025) (2 slides); Escambia Co., Pensacola, March 10, 1991, on Camellia sp., F.D. Bennett (1991-0425, 3010) (2 slides); Flagler Co., Bunnell, January 11, 2012, on Ilex cornuta (2012-369) (4 slides); Gadsden Co., Chattahoochee, December 13, 1990, on Camellia sp., F. Bennett (1990-011–013) (3 slides); Gadsden Co., Quincy, January 28, 2005, on Ilex sp., B. Cecil (2005-901); Gadsden Co., Quincy, March 20, 2012, on Poncirus sp., M. Bentley (2012-1944) (2 slides); Hillsborough Co., Brandon, May 29, 1986, on Ilex sp., J. Felty (1986-0453); Hillsborough Co., Tampa, March 13, 2018, on Citrus × paradisi, M. Briceno (2018-1035) (3 slides); Indian River Co., Vero Beach, August 26, 2013, on Ilex sp., J. Kennedy (2013-6273); Jefferson Co., Monticello, November 29, 1973, on Citrus sp., W.H. Pierce (19730348) (2 slides); Lake Co., Clermont, February 7, 2012, on Camellia sp., H. Alred (2012-793) (3 slides); Lake Co., Eustis, January 22, 1965, on Camellia sp., A.L. Bentley (1965-1655) (4 slides); Lake Co., Eustis, April 29, 2010, on Ilex sp., M. Sellers (2010-2438) (2 slides); Lake Co., Eustis, May 3, 2018, on Camellia japonica, M. Sellers (2018-2334) (2 slides); Leon Co., Tallahassee October 13, 1919, on Camellia japonica, P.F. Robertson (1919-1619) (2 Slides); Leon Co., Tallahassee, August 8, 1976, on Citrus × paradisi, S. Beidler (1976-0423) (3 slides); Leon Co., Tallahassee, October 2, 1978, on Euonymus sp., Q. Anglin (1978-3073) (3 slides); Leon Co., Tallahassee, December 13, 1991, on Camellia japonica, F. Bennett (1991-0445) (3 slides); Leon Co., Tallahassee, April 6, 2015, on Camellia sp., M. Bentley (2015-1687) (3 slides); Madison Co., Pinetta, October 22, 1985, on Poncirus trifoliata, J. Thomas (1985-1596); Manatee Co., Duette, March 2, 2005, on Ilex cornuta, K. Pippenger (2005-1077); Manatee Co., Oneco, October 19, 1923, on Camellia japonica, D.F. Schwarts (1923-1594) (2 slides); Marion Co., Citra, March 2, 2007, on Ilex sp., F. McHenry (2007-1335); Marion Co., Citra, March 2, 2007, on Ilex sp., F. McHenry (2007-1335) (2 slides); Marion Co., November 15, 2010, on Camellia sp. (2010-7091); Marion Co., Reddick, November 15, 2010, on Camellia sp., S. Wayte (2010-7091); Martin Co., Jensen Beach, April 23, 2014, on Ilex opaca, L. West (2014-2807) (2 slides); Martin Co., Stuart, December 13, 1979, on Raphiolepis umbellata, R. Gaskalla (1979-2966) (3 slides); Miami-Dade Co., Miami, December 6, 2012, Ilex sp., O. Garcia (2012-9157) (2 slides); Miami-Dade Co., Surfside, March 1, 2018, on Citrus sp., O. Garcia (2018-788) (2 slides); Nassau Co., Fernandina Beach, November 7, 2017, on Citrus sp., R. Leahy (2017-4261) (2 slides); Nassau Co., Yulee, September 21, 2012, on Ilex sp., R. Traya (2012-7169) (2 slides); Nassau Co., Yulee, March 14, 2013, on Ilex vomitoria, R. Traya (2013-1586) (2 slides); Nassau Co., Yulee, July 25, 2016, on Ilex cornuta, R. Traya (2016-3604); Orange Co., April 30, 2002, on Citrus reticulata, L. Brown (2002-1644); Orange Co., Apopka, January 30, 1990, on Ilex vomitoria, C. Murphy (1990-0353) (4 slides); Orange Co., Apopka, November 17, 1998, on Camellia japonica, L. Wilber (1998-3011) (5 slides); Orange Co., Apopka, April 20, 2002, on Citrus reticulata, L. Brown (2002-1644) (2 slides); Orange Co., Apopka, March 8, 2012, on Ilex sp., K. Gonzalez (2012-1575) (2 slides); Orange Co., Orlando, February 3, 1977, on Camelliasp., D.A. Graddy (1977-1514) (6 slides); Orange Co., Orlando, April 19, 2010, on Citrus reticulata, L. Russe (2010-2055); Orange Co., Orlando, February 16, 2012, on Ilex sp., R. Lopez (2012-1048) (3 slides); Orange Co., Orlando, February 2013, on Theaceae, A. Puppelo (2013-1127); Orange Co., Orlando, August 20, 2014, on Camellia japonica, T. Lyons (2014-5856) (2 slides); Orange Co., Pine Castle, February 5, 1962, on Camelliasp., A.C. Crews (1962-0333) (2 slides); Orange Co., Maitland, July 30, 1970, on Camellia japonica, E.R. Simmons (1970-1516) (2 slides); Palm Beach Co., West Palm Beach, October 9, 1979, on Persea americana, N. Miles (1979-029–032) (4 slides); Palm Beach Co., West Palm Beach, October 17, 2012, on Citrus sp., M. Clark (2012-7966) (2 slides); Pasco Co., Lutz, November 3, 2011, on Citrus sinensis, L. Osbeck (2011-8420) (2 slides); Pasco Co., Odessa, December 21, 2011, on Ilex cornuta (2011-9392); Pinellas Co., Clearwater, November 15, 1962, on Senecio confusus, C.E. Bingaman (1962-1623); Pinellas Co., Gulfport, December 14, 1977, on Citrus aurantifolia, K. Hickman (1977-0370) (2 slides); Pinellas Co., Oldsmar, May 5, 2005, on Citrus sp., D. Albritton (2005-2337); Pinellas Co., Palm Harbor, February 2012, on Citrus maxima, G. Campani and J. Hawk (2012-1112) (3 slides); Pinellas Co., Palm Harbor, November 16, 2010, on Citrus limon, J. Brownstein (2010-7134); Pinellas Co., Safety Harbor, April 11, 2012, on Syzygium jambos, L. Alston (2012-2672) (2 slides); Pinellas Co., St. Petersburg, May 4, 1979, on Citrus reticulata, K. Hickman (1979-0469) (2 slides); Pinellas Co., St. Petersburg, March 25, 2010, on Illicium floridanum, G. Bernard (2010-1506); Pinellas Co., Tarpon Springs, February 18, 2012, on Citrus sp., K. Edgerton (2012-1039) (2 slides); Pinellas Co., Tarpon Springs, March 3, 2015, on Cinnamomum camphora, B. Rose (2015-928) (2 slides); Polk Co., Haines City, January 22, 2013, on Ilex sp., S, Distelberg (2013-0328) (3 slides); Polk Co., Winter Haven, December 12, 1960, on Camellia japonica, A.C. McAulay and V.K. Norton (1960-0374); Polk Co., Winter Haven, September 9, 1963, on Gardenia sp., L.H. Heeb (1963-1653); Polk Co., Winter Haven, December 2, 2013, Camellia sp., C. Gibbard (2013-8729); Putnam Co., East Palaka, August 1, 2018, on Citrus sp., M. Cain (2018-4101) (2 slides); Putnam Co., Pomona Park, December 12, 1968, on Euonymus americanus, A.E. Graham (1968-0454) (6 slides); Santa Rosa Co., Bagdad, January 29, 1970, on Eurya japonica, R.W. Albritton (1970-1654) (7 slides); Santa Rosa Co., March 5, 2012, on Camellia japonica, M. Anderson (2012-1534); Seminole Co., Longwood, July 30, 1970, Camellia japonica, E.R. Simmons (1970-3068); Seminole Co., Oviedo, January 31, 2013, on Cleyera japonica, J. Krok (2013-640) (2 slides); Seminole Co., Oviedo, March 13, 2013, on Ilex opaca, J. Krok (2013-1691); Seminole Co., Sanford, July 16, 2012, on Myrtaceae, J. Krok, (2012-5308) (2 slides); St. Johns Co., St. Augustine, May 24, 2013, on Ilex sp., K. Theriault (2013-3667) (2 slides); St. Lucie Co., Ft. Pierce, February 21, 2005, on Ilex cornuta, D. Vazquez (2005-4069) (3 slides); Sumter Co., Bushnell, January 12, 2012, on Camellia sp., H. Alred (2012-377) (3 slides); Sumter Co., Bushnell, May 7, 2010, on Camellia sasanqua, H. Alred (2010-2501); Sumter Co., Center Hill, April 17, 2012, on Camellia japonica, H. Alred (2012-2752) (4 slides); Suwannee Co., Brandford, November 22, 2010, on Camellia japonica, W.W. Bailey (2010-7236); Suwannee Co., Live Oak, December 22, 2004, on Camellia japonica, Wayne Bailey (2004-8133); Suwannee Co., Live Oak, February 19, 2008, on Camellia japonica, W. Wayne Bailey (2008-209); Suwannee Co., Live Oak, February 28, 2012, on Ilex cornuta, D. Ruseell-Hughes and K. Collins (2012-1341) (3 slides); Suwannee Co., Live Oak, February 28, 2012, on Ilex cornuta, K. Collins (2012-1329) (2 slides); Suwannee Co., Live Oak, July 24, 2012, on Aquifoliaceae, D. Russell-Hughes (2012-5483) (2 slides); Suwannee Co., Live Oak, February 19, 2013, Camellia sp., W. Wayne Bailey (2013-1082); Taylor Co., November 18, 2010, on Camellia japonica, W. Wayne Bailey (2010-7155); Taylor Co., Perry, March 6, 1978, on Citrus limon, Q. Anglin (1978-0397); Taylor Co., Perry, March 6, 1978, on Citrus limon, Q. Anglin (1978-034); Taylor Co., Perry, March 6, 1978, on Citrus limon, Q. Anglin (1978-035); Taylor Co., Perry, March 8, 1979, on Euonymussp., Q. Anglin (1979-037); Taylor Co., Perry, March 8, 1979, on Euonymussp., Q. Anglin (1979-1599); Taylor Co., Perry, February 7, 2007, on Camellia japonica, Wayne Bailey (2007-748); Taylor Co., Perry, November 18, 2010, on Ilex cornuta, W.W. Bailey (2010-7155); Taylor Co., Steinhatchee, May 24, 2010, on Ilex cornuta, W. Wayne Bailey (2010-2993); Volusia Co., Daytona Beach, FL, March 14, 1963, on Malpighiasp., J.N. Pott (1963-2963) (2 slides); Volusia Co., Orange Mills, October 6, 1964, on Fortunellasp., A.E. Graham (1964-0470) (16 slides); Volusia Co., Holly Hill, November 16, 1971, on Fortunellasp., J.N. Pott (1971-0366) (2 slides); Volusia Co., Daytona Beach, November 23, 1976, on Ilex cornuta, J.N. Pott (1976-0378) (4 slides); Volusia Co., Ormond Beach, April 18, 2008, on Ilex cornuta, K. Coffey (2008-2272) (3 slides); Volusia Co., Edgewater January 13, 2012, on Ilex cornuta (2012-263) (3 slides).
Fiorinia theae, second-instar male, Alachua Co., Gainesville, October 9, 2019, January 24, 2020 on Ilex sp., M. Borden, D. Miller (2019-5696, 2020-287). Abbreviations: a) antenna; b) anterior spiracle; c) small gland spine; d) large microduct; e) large gland spine; f) remnants of legs; g) small microduct; h) enlargement of pygidium; i) small microduct; j) large microduct; j) enlargement of part of duct cluster; k) enlargement of communal duct.
Specimens examined for description and diagnosis
Little Rock, Arkansas, February 15, 1972, on Bradford Holly, 2nd ♀; Alachua Co., Gainesville, October 9, 2019, January 24, 2020 on Ilex sp., M. Borden, D. Miller 5 2nd ♂ (E2019-5696, E2020-287).
Discussion
The most recent study providing taxonomic keys of first-instar nymphs of Fiorinia species was published more than four decades ago (
Use of immature armored scales for identification is hampered by the fact that slide mounting protocols are tedious and laborious. Immature stages, especially first-instar nymphs, are very small, ca. 0.1–0.2 mm in length, and can easily be lost during the mounting process. We reexamined previously published mounting protocols (
One unexpected discovery during this project was that the morphology of second-instar males was more reliable for species recognition than any other instar, including the adult female. For example, we were unable to distinguish between second-instar females of F. proboscidaria and F. theae, but their second-instar males were easily separated using the number of communal ducts. Second-instar males of F. fioriniae are remarkably different from the same instar of all other species of Fiorinia found in the USA even though other instars are quite similar to one another.
Recently,
The final step was to compare the results of multigene molecular analyses of the Florida population, the Chinese population, and two Malaysian populations (D1184 and D1185). The results clearly show that these populations are the same species. The morphological differences suggested as diagnostic of F. yongxingensis are within the range of variation that occurs in F. phantasma. Therefore, we here treat F. yongxingensis as a junior synonym of F. phantasma.
We obtained 37 5’-COI barcodes representing nine Fiorinia species in this study. Overall, low intraspecific genetic distances and high interspecific genetic distances ranging from 9.1% to 15.2% between Fiorinia species emphasize the reliability of 5’-COI barcodes in molecular diagnostics of armored scale species. Our rapid slide-mounting protocol and the morphological keys to immatures and adults can provide time- and cost-effective diagnostics of Fiorinia species in the USA. However, for instances where specimens are damaged and cannot be mounted and where molecular diagnostics is the only option, barcodes will help to identify the species of Fiorinia. All of our DNA extractions are vouchered by permanently archived specimens in
Three populations of Fiorinia species (isolates D4674F, D4778A, and D4682A), collected from Lambir Hills National Park, Malaysia, September 26, 2013 from an undetermined host, identified as F. phantasma by BBN, were found to be genetically different from the F. phantasma populations from China, Florida and Malaysia. We reexamined the skins of the specimens used in our molecular analyses. The slides of isolates D4674F and D4778A are in poor condition and covered with fog, but we can see processes between the antennae and the shape of the pygidium, and they are consistent with the morphology of F. phantasma. The slide of isolate D4682A appears to have most characters of F. phantasma including the microducts between the posterior spiracles. This isolate is ca. 9% genetically distant from F. phantasma (based on COI) and is placed far from the subclade of F. phantasma (containing populations from China, Florida, and Malaysia) in the concatenated phylogenetic tree (Fig.
Recently phylogenetic analyses in
Field habitus of adult females, especially the character of the overlap between the first-instar and second-instar exuviae, was used for the first time in this study. For example, in the case of F. externa, the first-instar exuviae are barely touching the second-instar exuviae and form a distinct indentation between the attachment of the first- and second-instar exuviae (Suppl. material
Fiorinia japonica was eradicated from California and has been rediscovered three times since its first report in 1910 (
Conclusions
There are six main conclusions of our study. 1) The utilization of molecular barcodes is highly beneficial in diagnosing species of Fiorinia that occur in the USA. 2) The new keys in this study demonstrate that the USA species of Fiorinia can be identified using immature specimens. 3) Second-instar male morphology provided a reliable suite of characters for species-level identification. 4) Based on our comparative analysis of morphological characters and multigene molecular sequencing of specimens of F. phantasma and F. yongxinensis, it is clear that the latter is a junior synonym. 5) Of the different protocols tested for mounting immature specimens of Fiorinia, Hoyer’s mounting medium was the best for discerning delicate morphological characters but it was not desirable for permanent slide preparations. Balsam was the best for permanent mounts but did not provide the morphological clarity of Hoyer’s mounts. 6) The use of a mesh container in the process of mounting immatures is an effective method for preventing the loss of specimens. Overall, the use of the morphological and molecular data provides effective methods for early detection of new infestations and assists regulators in making control decisions.
Acknowledgements
This work was made possible, in part, by FY2019 United States Department of Agriculture Plant Protection Act Section 7721 funding. The authors thank Greg Hodges, Kate Fairbanks, Leroy Whilby, and Paul Skelly (Florida Department of Agriculture and Consumer Services, Division of Plant Industry) for administrating the grant funding. Participation by BBN was supported 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. Thanks to Susan Halbert and John Mcvay (FDACSDPI), for constructive reviews. The authors also thank all of our collaborators in this project including Lance Osborne (Department of Entomology & Nematology, Mid-Florida Research and Education Center, Institute of Food and Agricultural Sciences, University of Florida) and Amy Roda (Miami Laboratory, Plant Protection and Quarantine, Animal Plant Health Inspection Service, United States Department of Agriculture). This work would not be accomplished without their kind advice and field surveys. Collection in Lambir Hills National Park was conducted in collaboration with Takao Itioka (Kyoto University, Japan) and Geoff Morse (University of San Diego, California), in accordance with the Memorandums of Understanding signed between the Sarawak Forest Department (SFD, Kuching, Malaysia) and the Japan Research Consortium for Tropical Forests in Sarawak (JRCTS, Sendai, Japan) in December 2012. Thanks to Mohamad Shahbudin Sabki, Engkamat Lading, and Mohamad bin Kohdi, Paulus Meleng of SFD for help in obtaining research permission at the Lambir Hills National Park. We also thank Gillian W. Watson (The Natural History Museum, London) for helping us in tracing the type specimens of Fiorinia phantasma. DRM is grateful to the Division of Plant Industry including Paul Skelly and Greg Hodges for providing space and resources to conduct this research. Authors thank Natalia von Ellenrieder (California Department of Food and Agriculture) for sending us the F. pinicola sample. We thank technicians Chelsea Skojec, Gabi Ouwinga, and Lily Deeter (FDACS-DPI) for their help in mounting slides during this study. Authors also thank Gevork Arakelian (Los Angeles County Department of Agricultural Commissioner/Weights and Measures) for providing images of Fiorinia japonica and Lyle Buss (University of Florida) for providing images of Fiorinia fioriniae, F. theae, Kuwanaspis hikosani, Pseudaulacaspis cockerelli, P. pentagona, Poliaspis cycadis, and Odonaspis ruthae. We also acknowledge the United States National Collection of Scale Insects Photographs, USDA Agricultural Research Service especially for image # UGA5111048 of Fiorinia pinicola.
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Supplementary materials
Figures S1–S4
Data type: Phylogenetic trees and images (docx. file)
Explanation note: Figure S1. Comparison of field habitus of first-instar exuviae overlapping second-instar exuviae of seven Fiorinia species occurring in the USA. Figure S2. Maximum likelihood bootstrap consensus tree of the subtribe Fioriniina based on 28S, EF1-α, 5’-COI, 3’-COI, and COII. The clade highlighted in solid red indicates a monophyletic Fiorinia. Figure S3. Maximum likelihood bootstrap consensus tree of the subtribe Fioriniina based on 28S. The clade highlighted in red contains Fiorinia. Red dashed line indicates two Fiorinia sp. isolates placed with Pseudaulacaspis species. Dashed black lines indicate non-Fiorinia species placed in the Fiorinia clade. Bootstrap support values equal or greater than 50 are indicated on the tree. Figure S4. Neighbor-joining tree of Diaspididae 5’-COI barcodes. Terminal taxa are labeled to their narrowest identification-level. Numbers in parentheses after terminal taxa indicate how many sequences are represented in each cluster. The cluster of Fiorinia species is highlighted in red. Bootstrap support values greater than 75 are indicated on the tree. Nodes with 100 percent bootstrap support are indicated by a “*”.
Table S1
Data type: Sequences accession numbers (excel table)
Explanation note: Table S1. Taxa, isolates, and GenBank accessions used for phylogenetic analyses.
Table S2. PCR primers and thermocycling conditions.
Data type: PCR primers and thermocycling conditions (docx. file)
Explanation note: Table S2. PCR primers and thermocycling conditions. All PCR reactions were performed with 2-minute denaturation at 95⁰ C. Each subsequent cycle consists of a 30 second denaturation at 98⁰ C, a 30 second annealing step with temperature given below, and a 45 secs extension at 72⁰ C., end with a single 7 -minute extension at 72⁰ C. Primer sequences are given from 5’ to 3’.
Raw data
Data type: fas. file
Explanation note: Raw data file containing aligned sequences used in our phylogenetic analysis.