Research Article
Print
Research Article
A new species of Aulacaspis and a revived combination of Diaspididae (Hemiptera, Coccomorpha) from China
expand article infoMinmin Niu‡, Bo Cai§, Jiufeng Wei‡
‡ Shanxi Agricultural University, Taigu, China
§ Post-Entry Quarantine Station for Tropical Plant, Haikou, China

Corresponding author: Jiufeng Wei ( wjfeng@nwsuaf.edu.cn )

Academic editor: Takumasa Kondo
© 2023 Minmin Niu, Bo Cai, Jiufeng Wei.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation: Niu M, Cai B, Wei J (2023) A new species of Aulacaspis and a revived combination of Diaspididae (Hemiptera, Coccomorpha) from China. ZooKeys 1174: 301-314. https://doi.org/10.3897/zookeys.1174.105851
ZooBank: urn:lsid:zoobank.org:pub:B4B54CF2-322D-4B8B-9C5D-89B1CFA063A5
Open Access

Abstract

A new species of armored scale insect, Aulacaspis fanjingshanensis sp. nov. is described and illustrated based on adult female specimens collected on Rosaceae plants in China. A key to the Aulacaspis species known from Guizhou Province of China is provided. Our molecular study suggests that Aulacaspis schizosoma (Takagi, 1970) is not a true member of the genus Aulacaspis; the genus Superturmaspis Chen, 1983 is revived and A. schizosoma is transferred to it as Superturmaspis schizosoma (Takagi, 1970), revived combination, based on a molecular phylogeny.

Key words

Armored scale insect, molecular phylogenetic analysis, new record, pest, taxonomy

Introduction

Diaspididae, the largest family of Coccomorpha with more than 2700 species (García Morales et al. 2016), includes important economic pests and is one of the most invasive insect groups in the world.

The genus Aulacaspis Cockerell, 1893 belongs to the Chionaspidina, within the tribe Diaspidini, in the subfamily Diaspidinae (Normark et al. 2019). They are generally found living in groups on the host plants and suck the host plants’ sap through their piercing mouthparts. At high infestation levels of the pests, the leaves and branches of the host plants often wither, or even die, seriously affecting the yield and quality of the host plant, causing greater economic losses.

One hundred and fifty-one species of Aulacaspis have been recorded in the world and distributed worldwide (García Morales et al. 2016). The genus was established by Cockerell in 1893 based on Aspidiotus rosae Bouché, 1833, as the type species. This genus has been accepted by other scholars for more than 20 years (Kuwana 1926; Ferris 1936). Currently, the body shape of Aulacaspis mainly includes two types: the ‘rosae-type’ and the ‘vitis-type’ (Takagi 2012). And arrangement and number of dorsal and marginal ducts and median lobes are considered important morphological taxonomic features of Aulacaspis.

Guizhou Province in China has a subtropical monsoon climate, with rich species resources (http://invest.guizhou.gov.cn/tzgz/tzgk/). Hitherto, seven Aulacaspis species are currently recorded in Guizhou Province, namely A. crawii (Cockerell), A. guiyangensis Tian & Xing, A. longanae Chen, Wu & Su, A. paralonganae Tian & Xing, A. saigusai Takagi, A. yabunikkei Kuwana and A. zunyiensis Wei (Easton and Pun 1999; Hua 2000; Wei et al. 2016; García Morales et al. 2016; Tian and Xing 2022).

Aulacaspis was placed in Chionaspidina by Takagi (1999), within the tribe Diaspidini, in the subfamily Diaspidinae. Normark et al. (2019) found the non-monophyly of a genus-level phylogeny and taxonomy of Chionaspidina, including the Aulacaspis and Chionaspis Signoret, 1868.

Recently, a new species of Aulacaspis was discovered in Guizhou Province of China, and it is described and illustrated herein. And molecular study revealed the need for the resurrection of the genus Superturmaspis Chen, which was synonymized with Aulacaspis by Takagi (1985, 1999).

Materials and methods

Taxonomy

Samples of plants infested by the new species described in this study were collected in Fanjing Mountain (Guizhou Province, China). Permanent slide mounts of adult females from the samples were made according to Henderson (2011). Illustrations of adult females of the new species were drawn based on the type-series of slide-mounted specimens, each showing an overview of the dorsum on the left side and the venter on the right. Enlarged details of the significant features are illustrated around the body, not in direct proportion to each other. The morphological terminology and measurements in the descriptions follow those of Miller and Davidson (2005). All measurements are given in micrometres (µm). Measurements were made using the measurement tools NIS-Elements D. The type-series of the new species is deposited in the Insect Collection of Shanxi Agricultural University, Taigu, Shanxi Province, China (SXAU) and the Entomological Museum, Northwest A&F University, Yangling, Shaanxi, China (NWAFU).

Molecular phylogenetic analysis

DNA extraction and amplification

Each specimen was subjected to a joint molecular/morphological preparation protocol that resulted in genomic DNA from a single specimen and a permanent slide-mount of its cuticle (Normark et al. 2019). We used the single adult whole bodies for DNA extraction from each collection site. The total DNA was extracted using the Qiagen DNeasy Blood & Tissue kit (Qiagen, Valencia, California, USA), stored at 4 °C in the refrigerator. Other parts of the extraction follow the manufacturer’s protocols of the Qiagen kit. Polymerase chain reaction (PCR) was performed to amplify regions of the nuclear protein-coding gene elongation factor-1 alpha (EF-1α) the D2 and D3 expansion segments of the large subunit ribosomal RNA (28S). The PCR reactions contained a total of 25 μl, which included 12.5 μl 2×Taq MasterMix (with dye) (Coolaber, Beijing, China), 8.5 μl distilled water, 1 μl of each primer and 2 μl DNA template, and a negative control was included for all reactions. To amplify the fragment, we used the T100TM Thermal Cycler (BIO-RAD Laboratories, Inc., Hercules, California) to execute the cycling profiles given in Table 1.

Table 1.
Download as
CSV
XLSX

PCR primers and annealing temperatures, from Gruwell et al. (2007), Andersen et al. (2010) and Normark et al. (2019). Before thermal cycling begins, all PCR reactions start with a single 2-minute denaturation at 95 °C. Each subsequent cycle consists of a 30 s denaturation at 95 °C, a 1-minute annealing step with temperature given below, and a 2-minute extension at 72 °C. After thermal cycling is completed, all PCR reactions end with a single 10-minute extension at 72 °C. Primer sequences are given from 5' to 3'.

Gene Region Forward Primer Reverse Primer Annealing temperature profile
28S 28s_s3660 28s_a335 51 °C, 35 cycles at 48 °C
GAG AGT TMA ASA GTA CGT GAA AC TCG GAR GGA ACC AGC TAC TA
EF-1α EF-1α(a) (amplification/sequencing) GAT GCT CCG GGA CAY AGA G EF2rod (amplification/sequencing) ATG TGA GCG GTG TGG CAA TCC AA 58–42 °C, -2 °C/3 cycles + 11 cycles @ 42 °C

Sequence analysis and molecular systematics

We used PhyloSuite v.1.2.3 (Zhang et al. 2020) for molecular phylogeny and tree‐based analyses. The concatenated dataset (the full dataset) contained all 56 taxa. These taxa had both the 28S and EF1α gene fragments, although some individuals were missing one gene fragment. Some genes were obtained from NCBI (Table 2), originating from Morse and Normark (2006) and Normark et al. (2019). Each gene fragment was analyzed independently in order to look for congruence between the loci, and to verify the vertically inherited nature of the endosymbiont sequences. Maximum likelihood and Bayesian analyses were run on the dataset, as well as on the individual gene fragments themselves. The concatenated datasets were partitioned as follows: 28S; EF1α 1st, 2nd, and 3rd codon positions. This partition scheme was used in both the Maximum likelihood and Bayesian analyses. Each partition was examined in ModelFinder (Kalyaanamoorthy et al. 2017) to determine the most appropriate model on DNA evolution. Maximum likelihood phylogenies were inferred using IQ-TREE v.2.2.0 (Nguyen et al. 2015) under the TIMe+I+G4+F model for 5000 ultrafast (Minh et al. 2013) bootstraps. Bayesian Inference phylogenies were inferred using MrBayes v.3.2.7a (Ronquist et al. 2012) under SYM+I+G model (2 parallel runs, 1999997 generations), in which the initial 25% of sampled data were discarded as burn-in.

Table 2.
Download as
CSV
XLSX

Taxa included in the phylogenetic analysis.

Species name Isolate 28S/GenBank accession numbers EF-1α/GenBank accession numbers
Aonidiella aurantii D3778A KY219911.1 KY221741.1
Aspidiotus destructor D0496A KY219108.1 KY221360.1
Aulacaspis alisiana D2435B KY219609.1 –
D2435A KY219608.1 –
D2434B KY219606.1 –
D2434C KY219607.1 –
Aulacaspis crawii D3360B KY219853.1 –
Aulacaspis difficilis D0375A KY219047.1 KY221320.1
D0375E KY219048.1 KY221321.1
D2474B KY219626.1 KY221577.1
Aulacaspis spinosa D376A DQ145367.2 DQ145479.1
Aulacaspis distylii D0384A KY219057.1 KY221329.1
D0384C KY219058.1 –
Aulacaspis rosae D0395C KY219081.1 KY221343.1
D0395A GQ325451.1 GQ403823.1
D1433A KY219387.1 KY221462.1
D0395B GQ325452.1 GQ403824.1
Aulacaspis rosarum D1804D KY219405.1 KY221468.1
Aulacaspis tubercularis D1180B KY219359.1 KY221454.1
D1180C KY219360.1 –
D0225C KY218955.1 KY221261.1
D0225D KY218956.1 KY221262.1
D2952A KY219759.1 KY221643.1
D1180A KY219358.1 KY221453.1
Aulacaspis vitis D4354A KY219974.1 KY221782.1
Aulacaspis yabunikkei D2472A KY219622.1 –
Aulacaspis yasumatsui D0242A KY218963.1 KY221269.1
D1092C KY219318.1 –
D5050B KY220038.1 –
D0992A KY219297.1 KY221440.1
D1093A KY219319.1 –
D0307A KY219014.1 KY221302.1
D1833A KY219407.1 –
D0304A KY219011.1 KY221301.1
D0304C KY219013.1 –
D0304B KY219012.1 –
Chionaspis americana D2427A KY219604.1 KY221572.1
D2181A GQ325453.1 GQ403945.1
D0833D KY219245.1 –
D0833A GQ325454.1 GQ403869.1
Chionaspis etrusca D0606A KY219146.1 KY221382.1
D0687A GQ325455.1 GQ403852.1
D0687B KY219175.1 –
D0687C KY219176.1 –
D0687D KY219177.1 KY221398.1
D0687E KY219178.1 KY221399.1
D0810B KY219234.1 KY221418.1
D0811A KY219235.1 KY221419.1
D2528A KY219649.1 –
Chionaspis gleditsiae D0932A KY219276.1 JX677928.1
Chionaspis kosztarabi D1950A KY219471.1 KY221509.1
Chionaspis nyssae D1109B KY219325.1 KY221445.1
D0939A KY219278.1 KY221436.1
Chionaspis ortholobis D2713B KY219704.1 –
Chionaspis salicis D2903A KY219746.1 KY221633.1
D0662A GU349105.1 GU349853.1
D0662B KY219174.1 –
Chionaspis wistariae D0386A GU349092.1 GU349840.1
D0386B KY219061.1 –
Duplachionaspis displicata D3506B KY219861.1 KY221706.1
Duplachionaspis divergens D0309E KY219016.1 –
D0309C GQ325478.1 GQ403821.1
D1124B KY219336.1 KY221446.1
D2863A KY219741.1 –
D1125B KY219337.1 –
D0309B KY219015.1 KY221303.1
Duplachionaspis sicula D0623C KY219156.1 KY221389.1
D0623B KY219155.1 KY221388.1
D0623A GU349091.1 GU349839.1
Duplachionaspis spartinae D2909A KY219747.1 KY221634.1
D2442A KY219612.1 –
Megacanthaspis leucaspis D0401A KY219088.1 KY221347.1
D0401E KY219089.1 –
Pinnaspis aspidistrae D2535A KY219653.1 –
D2537A KY219654.1 –
D2939A KY219753.1 KY221638.1
Pinnaspis hikosana D0388D KY219065.1 –
Pinnaspis strachani D0248A KY218966.1 KY221272.1
D0248C KY218967.1 KY221273.1
Pseudaulacaspis pentagona D0372A KY219042.1 KY221316.1
Superturmaspis schizosoma – OQ917119 OQ869250
– OQ917120 OQ869251

Taxonomy

Aulacaspis Cockerell

Aulacaspis Cockerell, 1893: 180.

Type species

Aspidiotus rosae Bouché: by subsequent designation by Newstead 1901: 168.

Generic diagnosis

The following diagnosis is taken from Miller and Davidson (2005) and Takagi (2012). Female scale. White, circular, exuviae located on anterior end. Male scale. White, long, and narrow, exuviae located on anterior end. Adult female. Body elongate; derm membranous except for the margin of pygidium; prosoma generally swollen or wider than metathorax and abdomen, slightly squared in most species. Cephalothorax. Antennae each with a seta. Anterior spiracles usually with a cluster of trilocular pores, posterior spiracles with or without associated trilocular pores. Dorsal ducts present or absent on prosoma, scattered. Pygidium. Usually with three pairs of lobes (rarely with two or four pairs). Median lobes (L1) well developed, sunken into or protruding apex of pygidium, much larger than lobules of lateral lobes, zygotic basally or not, without marginal setae between lobes. In general, L1 are divided into two types depending on feeding site: bark-type, where individuals occur on bark and L1 protrudes at the end of the pygidium; and leaf-type, on leaves and L1 is sunken into the end of pygidium. Second lobes (L2) much smaller than L1, bilobed, divided into inner lobule and outer lobule, outer lobule usually smaller than inner lobule. Third lobes (L3) smaller than L2, bilobed, outer lobule smaller than inner lobule. Fourth lobes (L4) present in some species and usually represented by serrations along the body margin. Gland spines. Marginal gland spines developed, present on lateral of abdominal segment II and III; usually single on abdominal segments V–VIII, but in some species two or more. Marginal gland spines becoming shorter to conical on anterior segments; in some species they are called gland tubercles. Ducts. Dorsum with double-barred ducts. Marginal macroducts of pygidium usually larger than dorsal macroducts. Dorsal macroducts forming submedial and submarginal rows on abdominal segment and pygidium, sometimes occurring in two sizes. Dorsal ducts on the lateral margins of the second and the third abdominal segments. Ventral microducts scattered. Anal opening situated at the center of the pygidium, small. Perivulvar disc pores in five groups.

Remarks

As in other members of the subtribe Chionaspidina, Aulacaspis species have median lobes often joined by a zygosis; without setae or gland spines or marginal macroducts between median lobes (Miller and Davidson 2005; Takagi 2012). Aulacaspis is distinguished from other genera, especially from Chionaspis and Myrtaspis Takagi, 1999 by lacking lateral macroducts and gland spines on abdominal segment I and on the thorax, present in these locations on Chionaspis and Myrtaspis (Takagi 1999).

Aulacaspis fanjingshanensis sp. nov.

https://zoobank.org/9A000A90-B394-4409-A75D-F846C571BDAC
Figs 1–4

Material examined

Holotype female: China, Guizhou Province, Tongren city, Fanjing Mountain, on leaves of an undetermined plant of the family Rosaceae, coll. Wei Jiufeng and Niu Minmin, 8.viii.2015 (at SXAU). Paratypes female: CHINA, same data as holotype (at SXAU; a total of 2 adult females, 2 slides each containing 1 adult females).

Figures 1–4. 

Aulacaspis fanjingshanensis Niu & Wei, sp. nov., adult female 1 body 2 anterior spiracle 3 posterior spiracle 4 margin of pygidium.

Description of slide-mounted adult females

Prosoma. Prosoma with anterior margin rounded and with sides nearly parallel; peribuccal scleroses well formed. Antennae separated by 60–67 µm; each antenna with 1 seta. Anterior spiracles each associated with 24–27 trilocular disc pores; posterior spiracles each with 3–7 disc pores. Pygidial lobes. With 3 pairs of lobes; L1 well developed, largely sunken into apex of pygidium, base of each L1 with a mesad linear extension onto ventral derm, these extensions separated from each other by a narrow space; L1 elongate and divergent, widest separation distance 19–22 µm, divergent mesal margins minutely serrate, lobe apices blunt or rounded. Setae absent from between median lobes. L2 and L3 well developed, bilobulate. Marginal macroducts, of two-barred type, nearly as long as L1, absent between L1, one present between L1 and L2, two present between L2 and L3, two present on the abdominal segment V. Dorsal macroducts on pygidium and abdominal segments shorter than marginal macroducts; of two-barred type, arranged segmentally in submedian and submargin rows; submarginal dorsal macroducts present on abdominal segment III to V: 4–9 on segment III, 1–6 on segment IV, 2–5 on segment V; submedian dorsal macroducts present on segment III to VI, on III and IV being divided into segmental and infrasegmental series: front row 4, rear row 3–4 on segment III, front row 4–5, rear row 1–3 on segment IV, 2–5 on segment V, 2–3 on segment VI. Dorsal microducts present on abdominal segment II to head: 5 on forehead, 2–4 on the position of mouthparts, 3–4 on the submargin of prothorax, 5–6 on the submargin of mesothorax, 1 on the middle area of mesothorax, 4 on the submargin area of metathorax, 3 on the submedian area of segment I, 5 on the submargin area of segment I, 2–3 on the submedian area of segment II, 0–2 on the submargin area of segment II. Lateral ducts few, 5–7 in total, present between abdominal area of segments II and III, 2–4 on segment II, 2–5 on segment III, smaller than dorsal ducts present on abdominal segments and pygidium. Ventral microducts scattered on pygidium, few. Marginal gland spines on each side numbering 2‒4 on segment IV, and 1 on V; also, with 1 lateral to each pygidial lobe. Each side with lateral gland spines numbering 5‒7 on segment II, and 6–13 on segment III. Perivulvar pores in five groups, with 10–17 in the median group, 20–26 in each anterolateral group and 27–35 in each posterolateral group.

Host

Rosaceae sp.

Etymology

The specific epithet is formed by a combination of Fanjing Mountain, the type locality, and the Latin “-ensis”, meaning “from”.

Distribution

China (Guizhou Province).

Remarks

Aulacaspis fanjingshanensis sp. nov. is similar to other species in the Yabunikkei complex (A. yabunikkei Kuwana, 1926, A. shirodamo Takagi, 2014 and A. neolitseae Takagi, 2014) by the form of its body shape, but can be distinguished from A. yabunikkei by the following characteristics of the adult female (character-states of A. yabunikkei in brackets): (1) dorsal microducts present on abdominal segment II to head (absent); (2) marginal macroducts of abdominal segment VII scarcely or only a little extending anteriorly beyond bases of these extensions (extending); and (3) median lobes separated from each other by a narrow space (united basally).

The stable diagnostic characters that distinguish A. fanjingshanensis sp. nov. from A. shirodamo and A. neolitseae are very few. The differences are the presence of dorsal microducts that are found from the head region down to abdominal segment II and median lobes separated from each other by a narrow space.

Key to adult female Aulacaspis Cockerell from Guizhou Province of China

1 Abdominal segment I with dorsal microducts or macroducts 2
– Abdominal segment I without dorsal microducts or macroducts 3
2 Abdominal segment I with dorsal microducts Aulacaspis fanjingshanensis sp. nov.
– Abdominal segment I with dorsal macroducts A. crawii (Cockerell)
3 Abdominal segment VI without dorsal macroducts A. zunyiensis Wei
– Abdominal segment VI with dorsal macroducts 4
4 Abdominal segment II with submedial dorsal macroducts 5
– Abdominal segment II without submedial dorsal macroducts 7
5 Prosomatic tubercles prominent, prosoma angular A. guiyangensis Tian & Xing
– Prosomatic tubercles not developed, prosoma suborbicular 6
6 Lateral gland spines many, numbering about 30–45 on each side A. paralonganae Tian & Xing
– Lateral gland spines few, numbering about 10–25 on each side A. longanae Chen, Wu & Su
7 Submarginal and submedial dorsal macroducts all arranged in irregularly double or triple rows A. saigusai Takagi
– Submedial dorsal macroducts arranged in double rows A. yabunikkei Kuwana

Superturmaspis schizosoma (Takagi, 1970), revived combination

Chionaspis schizosoma Takagi, 1970: 77.

Superturmaspis schizosoma (Takagi, 1970); Chen 1983: 86. Change of combination.

Aulacaspis schizosoma (Takagi, 1970); Takagi 1985: 49. Change of combination.

Semichionaspis schizosoma (Takagi, 1970); Tang 1986: 170. Change of combination.

Aulacaspis schizosoma (Takagi, 1970); Takagi 1999: 149. Revived combination.

Material examined

China, Zhejiang Province, Taohua Island in Zhoushan Islands, on Lauraceae leaves, coll. Niu Minmin, 31.v.2017 (at NWAFU).

Remarks

We confirmed the non-monophyly of the genus-level relationships and taxonomy of Chionaspidina, including the genera Aulacaspis and Chionaspis (Normark et al. 2019). The same level of strong statistical support was observed for the species in our two molecular phylogenetic trees (ML and BI trees, Figs 6, 7). On those phylogenetic trees, Superturmaspis schizosoma comprises a clade separate from the one that includes the genera Aulacaspis, Chionaspis, Duplachionaspis, Pinnaspis and Megacanthaspis (In ML tree, the bootstrap value is 52, and in BI tree, the posterior probability is 1).

Our phylogenetic study has demonstrated that the type species of Aulacaspis and Chionaspis (A. rosae and C. salicis) are more closely related to each other than either is to Superturmaspis schizosoma. Hence, A. schizosoma was wrongly assigned to Aulacaspis (Figs 6, 7), although the species Superturmaspis schizosoma (Takagi, 1970) is morphologically very close to Aulacaspis, sharing very similar characters of the pygidial margin and a similar distribution of dorsal ducts, but differing by being widest across the mesothorax, body of adult female is top form in outline, pronounced contracted segmentation on lateral margins which contains the cephalothorax and abdominal segments preceding the pygidium.

We transferred the species back to Superturmaspis based on our molecular analysis. Superturmaspis was previously established by Chen, 1983, and included only one species, Superturmaspis schizosoma. The adult female specimens (Fig. 5) substantially agree with the type-series from Taiwan (Takagi 1970). The body of the adult female is robust at maturity, spinning top form in outline, widest across the mesothorax, pronouncedly contracted segmentation on lateral margins which contains the cephalothorax and abdominal segments preceding the pygidium, and marginal macroducts rather stout and long.

Figure 5. 

Superturmaspis schizosoma (Takagi, 1970), revived combination, adult female.

Figure 6. 

The ML tree based on the concatenated data of 28S and EF-1α. Bootstrap values (BV) are shown on each node. Orange shading represents the position of Superturmaspis schizosoma (Takagi, 1970) on the phylogenetic tree.

Figure 7. 

The BI tree based on the concatenated data of 28S and EF-1α. Posterior probabilities (pp) are shown on each node. Orange shading represents the position of Superturmaspis schizosoma (Takagi, 1970) on the phylogenetic tree.

Additional information

Conflict of interest

The authors have declared that no competing interests exist.

Ethical statement

No ethical statement was reported.

Funding

This work was supported by the National Natural Science Foundation of China (No. 32100370), Science and Technology Innovation Funds of Shanxi Agricultural University (2020BQ79), the Excellent Doctoral Award of Shanxi Province for Scientific Research Project (SXBYKY2021024) and Science and Technology Innovation Projects of Universities in Shanxi Province (2021L097), Research Project Supported by Shanxi Scholarship Council of China (2020-065), the Natural Science Foundation of Shanxi, China (202103021224132), Hainan Province Science and Technology Special Fund (ZDKJ2021016).

Author contributions

Data curation: MN. Funding acquisition: BC, MN. Investigation: MN. Resources: JW. Software: MN. Writing – original draft: MN. Writing – review and editing: MN, JW.

Author ORCIDs

Minmin Niu https://orcid.org/0009-0004-6506-4950

Data availability

All of the data that support the findings of this study are available in the main text.

References

  • Andersen JC, Wu J, Gruwell ME, Gwiazdowski R, Santana SE, Feliciano NM, Morse GE, Normark BB (2010) A phylogenetic analysis of armored scale insects, based upon nuclear, mitochondrial, and endosymbiont gene sequences. Molecular Phylogenetics and Evolution 57: 992‒1003. https://doi.org/10.1016/j.ympev.2010.05.002
  • Bouché PF (1833) Naturgeschichte der Schädlichen und Nützlichen Garteninsekten und die bewährtesten Mittel zur Vertilgung der ersteren. Berlin In der Nicolaischen Buchhandlung, 176 pp. https://doi.org/10.5962/bhl.title.9692
  • Chen FG (1983) The Chionaspidini (Diaspididae, Coccoidea, Homoptera) from China. Science & Technology Publishing House, Sichuan Province, China, 175 pp.
  • Cockerell TDA (1893) Museum notes, Coccidae. Journal of the Institute of Jamaica 1: 180.
  • Easton ER, Pun WW (1999) Observations on twelve families of Homoptera in Macau, Southeastern China, from 1989 to the present. Proceedings of the Entomological Society of Washington 101(1): 99–105.
  • Ferris GF (1936) Contributions to the knowledge of the Coccoidea (Homoptera). II. (Contribution no. 2). Microentomology 1: 17–92.
  • García Morales M, Denno B, Mille DR, Miller GL, Ben-Dov Y, Hardy NB (2016) ScaleNet: a literature-based model of scale insect biology and systematics. http://scalenet.info [Accessed 22 November 2022]
  • Gruwell ME, Morse GE, Normark BB (2007) Phylogenetic congruence of armored scale insects (Hemiptera: Diaspididae) and their primary endosymbionts from the phylum Bacteroidetes. Molecular Phylogenetics and Evolution 44: 267‒280. https://doi.org/10.1016/j.ympev.2007.01.014
  • Henderson RC (2011) Diaspididae (Insecta: Hemiptera: Coccoidea). Fauna of New Zealand 66. Manaaki Whenua Press, Lincoln, Canterbury, 275 pp.
  • Hua LZ (2000) List of Chinese Insects (Vol. 1). Zhongshan University Press Guangzhou, China, 448 pp.
  • Kalyaanamoorthy S, Minh BQ, Wong TKF, von Haeseler A, Jermiin LS (2017) ModelFinder: Fast model selection for accurate phylogenetic estimates. Nature Methods 14(6): 587–589. https://doi.org/10.1038/nmeth.4285
  • Kuwana SI (1926) The diaspine Coccidae of Japan. IV. Genera Cryptoparlatoria, Howardia, Sasakiaspis [n. gen.] Diaspis, Aulacaspis, Pinnaspis and Prontaspis. Department of Agriculture and Commerce, Bureau of Agriculture. Injurious Insects and Pests, Japan 4: 1–44.
  • Miller DR, Davidson JA (2005) Armored Scale Insect Pests of Trees and Shrubs (Hemiptera: Diaspididae). Cornell University Press, Ithaca, 456 pp.
  • Minh BQ, Nguyen MA, von Haeseler A (2013) Ultrafast approximation for phylogenetic bootstrap. Molecular Biology and Evolution 30(5): 1188–1195. https://doi.org/10.1093/molbev/mst024
  • Nguyen LT, Schmidt HA, von Haeseler A, Minh BQ (2015) IQ-TREE: A fast and effective stochastic algorithm for estimating maximum-likelihood phylogenies. Molecular Biology and Evolution 32(1): 268–274. https://doi.org/10.1093/molbev/msu300
  • Normark BB, Okusu A, Morse GE, Peterson DA, Itioka T, Schneider SA (2019) Phylogeny and classification of armored scale insects (Hemiptera: Coccomorpha: Diaspididae). Zootaxa 4616(1): 1–98. https://doi.org/10.11646/zootaxa.4616.1.1
  • Ronquist F, Teslenko M, van der Mark P, Ayres DL, Darling A, Höhna S, Larget B, Liu L, Suchard MA, Huelsenbeck JP (2012) MrBayes 3.2: Efficient Bayesian phylogenetic inference and model choice across a large model space. Systematic Biology 61(3): 539–542. https://doi.org/10.1093/sysbio/sys029
  • Takagi S (1970) Diaspididae of Taiwan based on material collected in connection with the Japan-U.S. Cooperative Science Programme, 1965 (Homoptera: Coccoidea). Pt. II. Insecta Matsumurana 33: 1–146.
  • Takagi S (1985) The scale insect genus Chionaspis: A revised concept (Homoptera: Coccoidea: Diaspididae). Insecta Matsumurana New Series 33: 1–77.
  • Takagi S (1999) For a better understanding of Aulacaspis: The calcarata species group (Homoptera: Coccoidea: Diaspididae). Insecta Matsumurana New Series 55: 133–180.
  • Takagi S (2012) Atypical species of Aulacaspis (Sternorrhyncha: Coccoidea: Diaspididae). Insecta Matsumurana New Series 68: 17–115.
  • Tang FT (1986) The scale insects of horticulture and forest of China, Volume III. Shanxi Agricultural University Press Taigu, Shanxi, 305 pp.
  • Wei JF, Jing XP, Zhang HF (2016) A new species of Aulacaspis Cockerell, 1893 from China with a key to Chinese species (Hemiptera, Coccoidea, Diaspididae). ZooKeys 619: 13–24. https://doi.org/10.3897/zookeys.619.9399
  • Zhang D, Gao F, Jakovlić I, Zou H, Zhang J, Li WX, Wang GT (2020) PhyloSuite: An integrated and scalable desktop platform for streamlined molecular sequence data management and evolutionary phylogenetics studies. Molecular Ecology Resources 20(1): 348–355. https://doi.org/10.1111/1755-0998.13096
login to comment