Two-sex life table and predation rate of Cybocephalus flavocapitis Smith (Coleoptera: Cybocephalidae) reared on Aulacaspis yasumatsui Takagi (Hemiptera: Diaspididae), in Taiwan

Development, survival, and fecundity of the predator, Cybocephalus flavocapitis Smith, reared on its host Aulacaspis yasumatsui Takagi, were studied at 24 °C, 70 ± 10% RH, and photoperiod of 12:12 (L:D) h under laboratory conditions. Developmental periods for egg, larval, and pupal stages were 6.4, 12.1 and 14.3 d, respectively. The mean longevity of adults was 85.1 d. Raw life history data of all individuals were pooled and analyzed according to the age-stage, two-sex life table theory. The intrinsic rate of increase (r) was 0.05 d^? 1, the finite rate of increase (λ) was 1.05 d^? 1, the net reproductive rate (R₀) was 29.1 offspring, and the mean generation time (T) was 63.3 d. When we incorporated the age-stage variability of predation rate with the two-sex life table, our results indicated that the predation rates of larvae and adults were 743 eggs and 1595 scales, respectively. The net predation rate of C. flavocapitis is 1874 scales. Additionally, we compared life table results of C. flavocapitis with those of a previous life table study on Cybocephalus nipponicus because both species are currently used for the biological control of A. yasumatsui in Taiwan.     

Effect of temperature on the development and consumption of Phaenochilus kashaya (Coleoptera: Coccinellidae), a predator of the cycad aulacaspis scale,Aulacaspis yasumatsui

The cycad aulacaspis scale (CAS), Aulacaspis yasumatsui Takagi (Hemiptera: Diaspididae), is a serious pest of ornamental cycads in the southeastern United States. In Florida, CAS was first reported in Miami in 1996 and is now present in 43 Florida counties. Chemical control of CAS can be effective, but it is expensive, and insecticides must be frequently and regularly applied. Recent surveys of natural enemies in Thailand identified a new potential biological control agent of CAS. The objective of this study was to examine the temperature requirements and consumption rates of Phaenochilus kashaya Giorgi and Vandenberg (Coleoptera: Coccinellidae). Survival, developmental time and number of scales consumed by P. kashaya were measured at four constant temperatures (20, 25, 30 and 35°C). In addition, adult longevity and consumption were recorded at three constant temperatures (20, 25 and 30°C). Phaenochilus kashaya completed development to adulthood at temperatures ranging from 20 to 30°C. The highest survival rate (48%) was obtained at 25°C, and the accumulated degree-days for this species was 714. Phaenochilus kashaya is a voracious predator of CAS, consuming 380 scales during the larval stage and an estimated 4700 scales during the adult stage. Adult longevity varied from 59 days at 30°C to 220 days at 20°C for females, and 31 days at 30°C to 148 days at 20°C for males. The intrinsic rate of increase (r _m) was 0.34 at 20°C, which is high compared with other coccinellid species. According to this study, P. kashaya has the potential to be an effective biological control agent of the CAS in Florida.     

Host range of Cybocephalus flavocapitis and Cybocephalus nipponicus,two potential biological control agents for the cycad aulacaspis scale,Aulacaspis yasumatsui

Host-range and host-specificity tests were performed with Cybocephalus flavocapitis T. R. Smith (Coleoptera: Cybocephalidae) and Cybocephalus nipponicus Endrödy-Younga (Coleoptera: Cybocephalidae), two biological control candidates against the invasive cycad aulacaspis scale, Aulacaspis yasumatsui Takagi (Hemiptera: Diaspididae). Seventeen native scale species plus the invasive A. yasumatsui scale were tested in growth chambers using no-choice tests and hosts suitable for each of the two predatory beetles. The results revealed that the two Cybocephalus beetles, one imported species from Thailand and one native species, both fed on relatively similar scale prey species. Additionally, the adult beetles of these two species oviposited only on Diaspididae scales. The results showed that both Cybocephalus beetles may share a similar host niche in Taiwan.     

Key to Species of Aulacaspis (Hemiptera: Diaspididae) Intercepted at the Republic of Korea Ports of Entry

Six species of aulacaspis scales, Aulacaspis crawii (Cockerell), Aulacaspis madiunensis (Zehntner), Aulacaspis neospinosa Tang, Aulacaspis spinosa (Maskell), Aulacaspis tubercularis Newstead and Aulacaspis yasumatsui Takagi, have been taken in quarantine at the republic of Korea ports of entry between 1996 and 2005. We have also found the specimens of A. yasumatsui Takagi on Cycas plants from Malaysia and Taiwan and provide a key with illustrative photographs to identify intercepted specimens of Aulacaspis.     

Life table and heat tolerance of Acyrthosiphon pisum (Hemiptera: Aphididae) in subtropical Taiwan

The effect of temperature on the life table of Acyrthosiphon pisum reared on Pisum sativum was evaluated under laboratory conditions using temperatures of 10, 15, 20, 25, 30, and 35°C. The development time of juvenile A. pisum decreased with increasing temperature (from 21.3 days at 10°C to 4.7 days at 35°C). Adult longevity also decreased with increasing temperature (from 53.2 days at 10°C to 2.3 days at 35°C). Interestingly, 70% and 25% of A. pisum nymphs reared at 30°C and 35°C, respectively, successfully developed into adults. These temperatures have previously been considered unsuitable for A. pisum development. However, adult aphids reared at 30°C and 35°C failed to reproduce. Linear regression analysis revealed that the lower development threshold of A. pisum was 153.1 degree‐days above 1.9°C. Maximal average reproductive capability was observed at 10°C for A. pisum adults, with each adult producing more than 120 nymphs. The intrinsic rate of increase (r_m) of A. pisum increased from 0.124/day at 10°C to 0.337/day at 25°C, whereas opposite trends were observed for the net reproductive rate (R₀) and the mean generation time (GT). At 20°C and 25°C, the intrinsic rate of increase of A. pisum was significantly higher than at 10°C and 15°C (P < 0.0001), indicating that 20°C and 25°C are within the optimal range for the growth of A. pisum, and that 30°C is beyond the upper threshold limit for reproduction, which involves a temperature range that is narrower than that of the survival range (upper limit is unknown, but above 35°C).     

A molecular phylogenetic study of armoured scale insects (Hemiptera: Diaspididae)

Abstract.  Armoured scale insects are economically important parasites of woody plants and grasses. They are promising subjects for the evolutionary study of physiology (no complete gut), genetics (chimerism, paternal genome elimination, frequent parthenogenesis) and coevolution (with host plants, parasitoids, Septobasidium fungi, endosymbiotic bacteria). Little phylogenetic work has been accomplished with armoured scales, and uncertainty surrounds their classification. Here, we report the phylogenetic results of Bayesian and parsimony analyses of 705 base pairs of Elongation Factor 1α and 660 base pairs of 28S from eighty-nine species of armoured scale insects, representing forty-seven genera and five tribes in the subfamilies Diaspidinae and Aspidiotinae, together with two outgroups. 28S was aligned based on a secondary structural model. Our results broadly corroborate the major features of the existing classification, although we do not find perfect monophyly of any of the traditionally recognized subfamilies or tribes. The subfamily Aspidiotinae is paraphyletic with respect to the subfamily Diaspidinae. Diaspidinae consists of two main clades that only roughly correspond to the tribes Lepidosaphidini and Diaspidini. Diaspidini is nearly monophyletic, except that it includes a single aspidiotine species. Other members of the tribe Aspidiotini form a clade, except that the clade includes a single species of Leucaspidini and excludes Maskellia and Pseudaonidia . Our results weakly support the hypothesis that the most recent common ancestor of the Diaspididae had adult females that were permanently enclosed within the derm of the second instar (the pupillarial habit) and had diploid adult males that eliminated their paternal genomes during spermatogenesis (late paternal genome elimination).     

DNA barcodes for two scale insect families, mealybugs (Hemiptera: Pseudococcidae) and armored scales (Hemiptera: Diaspididae)

Although DNA barcode coverage has grown rapidly for many insect orders, there are some groups, such as scale insects, where sequence recovery has been difficult. However, using a recently developed primer set, we recovered barcode records from 373 specimens, providing coverage for 75 species from 31 genera in two families. Overall success was >90% for mealybugs and >80% for armored scale species. The G·C content was very low in most species, averaging just 16.3%. Sequence divergences (K2P) between congeneric species averaged 10.7%, while intra-specific divergences averaged 0.97%. However, the latter value was inflated by high intra-specific divergence in nine taxa, cases that may indicate species overlooked by current taxonomic treatments. Our study establishes the feasibility of developing a comprehensive barcode library for scale insects and indicates that its construction will both create an effective system for identifying scale insects and reveal taxonomic situations worthy of deeper analysis.     

Effects of pyriproxyfen on California red scale (Hemiptera: Diaspididae) development and reproduction

Two life stages of a laboratory colony of California red scale, Aonidiella aurantii (Maskell) (Hemiptera: Diaspididae), were exposed to 10 concentrations of pyriproxyfen to evaluate the effect of this insect growth regulator (IGR) on scale development and fecundity. First instars exposed to pyriproxyfen responded with mortality during the first and second molts. Second instars were more tolerant of pyriproxyfen than were first instars, indicating that growers should treat California red scale in the field before the first molt to achieve maximum efficacy. Male scales responded with an LC50 12-fold lower and an LC90 47-fold lower than was observed for female scales. Female scales that survived first instar exposure to pyriproxyfen experienced reduced fecundity with increasing pyriproxyfen concentration. Reduced fecundity was due to reduced survival of the females rather than sterility. Results from this study provide baseline California red scale susceptibility data for pyriproxyfen, and they suggest that 1 ppm pyriproxyfen can be used as a concentration that would discriminate between susceptible and resistant populations of scales. Field monitoring for incipient pyriproxyfen resistance in California red scale should be implemented in the San Joaquin Valley of California, and if resistance is detected, integrated resistance management strategies should be used to slow the progression of resistance.     

The unusual cover of the armoured scale Cryptaspidiotus barbusano (Lindinger) (Hemiptera: Diaspididae: Aspidiotinae) with comments on the scale covers of other Diaspididae

Cryptaspidiotus barbusano (Lindinger 1908) is an endemic species of the Canary Islands, only known on Apollonias barbujana (Lauraceae), a plant growing in the “Laurisilva” or laurel forest, in several Macaronesian islands. Recently C. barbusano was collected on Heberdenia excelsa (Myrsinaceae), a new host plant and family, in the Jardín Botánico Canario Viera y Clavijo, Las Palmas, Gran Canaria. The recent collection of C. barbusano gives us the opportunity to redescribe its unusual adult female scale cover. The truncate cone-shaped scale cover and its peculiar reflexed exit tunnel for crawlers, with microscopic details of this rare species, are described and illustrated. The shape of the exit tunnel is discussed and the scale cover is compared with scale covers of several other diaspidid species. Finally, this study shows that C. barbusano is not a pupillarial species as formerly described.     

Monitoring and mechanisms of organophosphate resistance in San Jose scale,Diaspidiotus perniciosus (Hemiptera: Diaspididae)

San Jose scale, Diaspidiotus perniciosus (Comstock), is a serious pest in Chilean apple tree orchards, and a number of organophosphate insecticides were used to control them for decades. Recently, control failures with these insecticides were reported and linked to insecticide resistance development. In this study, 40 San Jose scale field populations were collected and their susceptibility to two commonly used organophosphate insecticides, that is chlorpyrifos and methidathion, was assessed. The obtained bioassay data suggest moderate levels of resistance to both insecticides when compared to a reference susceptible strain. The highest resistance ratio (RR) detected for chlorpyrifos and methidathion was 31‐fold and 11‐fold, respectively. The bioassay results suggest the occurrence of a significant cross‐resistance between both compounds. Biochemical measurements revealed a role for esterases in conferring resistance to organophosphates, but not modified acetylcholinesterase. The spatial spread and extend of insecticide resistance were also evaluated. Our result shows that no autocorrelation can be assumed, and then, insecticide resistance is caused by random factors.     

Effect of papaya trunk angle on infestation by white peach scale,Pseudaulacaspis pentagona (Hemiptera: Diaspididae)

Two papaya (Carica papaya L.) seedlings growing in one planting hole often results in angular or non-vertical growth of the trees. Data on trunk angularity, or leaning, (deviation from the vertical line of reference) and white peach scale, Pseudaulacaspis pentagona Targioni-Tozzetti (Hemiptera: Diaspididae), densities on paired papaya trees were collected approximately 1 year after infestation of a papaya field was discovered. Paired trees showed a significantly higher degree of leaning than single trees. White peach scale densities were significantly higher on tree trunks with a greater departure from vertical in paired comparisons. Therefore, paired tree planting practices may facilitate the development of economic infestations of white peach scale populations in papaya orchards.     

The effect of the armored scale,Rhizaspidiotus donacis (Hemiptera: Diaspididae), on shoot growth of the invasive plant Arundo donax (Poaceae: Arundinoideae)

The effect of feeding by the armored scale, Rhizaspidiotus donacis (Leonardi, 1920) (Hemiptera: Diaspididae) on the growth of the plant Arundo donax L. (Poaceae) was evaluated under field conditions in its native range. The study was designed to evaluate the impact of R. donacis, a candidate agent for biological control of A. donax which is invasive in arid riparian ecosystems of the Southwestern USA and Mexico. The study was carried out at five A. donax sites in the Province of Alicante, Spain, differing in altitude and climate. At each site, 30 infested lateral shoots were selected and 15 were randomly treated monthly with imidacloprid insecticide. Shoot lengths were measured monthly over a 1-year period in a comparative growth analysis. Shoots infested with R. donacis had an over 2-fold reduced growth rate as compared to treated shoots. Growth of shoots varied by site, and the effect of R. donacis on growth was most pronounced in the late spring, when mature females produced first instar scale crawlers. The impact of R. donacis on A. donax growth under field conditions in the native range, combined with its narrow host specificity, indicate that R. donacis is a promising candidate for biological control of A. donax in North America and other areas invaded by this weed.     

不同温度下康氏粉蚧实验种群生命表

为了探索温度对康氏粉蚧Pseudococcus comstocki发育和繁殖的影响, 经室内实验, 建立了康氏粉蚧在17℃,20℃,23℃,26℃,29℃和32℃下的实验种群生命表。结果表明： 在17~29℃温度范围内, 随着温度的升高康氏粉蚧的世代发育历期缩短, 在17℃下雌雄世代的发育历期最长（分别为121.27 d和89.64 d）, 而29℃时仅分别为52.10 d和36.01 d,且其温度与发育速率的关系符合 Logistic模型。雌虫世代的发育起点温度为8.69℃, 有效积温为1 020.90日·度; 雄虫世代的发育起点温度为10.27℃, 有效积温为659.04日·度。在26℃时, 康氏粉蚧的世代存活率最高(81.94%), 种群趋势指数(I)也最高(322.27), 有利于种群增长; 而在17℃和29℃, 康氏粉蚧的世代存活率分别为64.85%和67.23%, 种群趋势指数(I)分别为69.91和42.19, 且32℃时康氏粉蚧1龄若虫生长停滞。说明高温和低温都不利于康氏粉蚧的种群增长, 且高温的影响大于低温。     

Identification of the sex pheromone of the invasive scale Acutaspis albopicta (Hemiptera: Diaspididae), arriving in California on shipments of avocados from Mexico

As a result of relaxation of importation restrictions ordered by the Animal and Plant Health Inspection Service of the U.S. Department of Agriculture, shipments of fresh avocados from Mexico began entering California year-round in 2007, despite the fact that these shipments were heavily infested with a number of exotic and potentially invasive armored scale species that are not thought to be present in California. Here, we report the identification of the sex pheromone of one of these species, Acutaspis albopicta (Cockerell), from a quarantine colony of these insects initiated from specimens collected from commercial shipments of Mexican avocados. The compound was identified as [(1S,3S)-2,2-dimethyl-3-(prop-1-en-2-yl)cyclobutyl)]methyl (R)-2-methylbutanoate, and was similar in structure to the pheromones of several other scale and mealybug species. In laboratory bioassays, the pheromone was highly attractive to male scales in microgram doses. The pheromone will provide a very sensitive and selective tool for detection of the scale to try and prevent its permanent establishment in California.     

Biology and prey range of Cryptognatha nodiceps (Coleoptera: Coccinellidae), a potential biological control agent for the coconut scale, Aspidiotus destructor (Hemiptera: Diaspididae)

Cryptognatha nodiceps Marshall is an important natural enemy used in biological control programmes against Aspidiotus destructor (Signoret), an introduced pest of coconut in many countries. In order to increase our understanding of C. nodiceps, the present study to quantify aspects of the developmental, reproductive and feeding biology under constant environmental conditions (26±2°C, 55-65% RH) was carried out. In addition, methods for culturing the scale and the beetle were developed. The average duration of development was 19.2±0.1 days for males and 19.6±0.2 days for females. Female longevity and lifetime fecundity was 35.6±5.1 days and 141.0±35.6 eggs, respectively. Life table statistics were as follows: reproductive rate, 9.99, intrinsic rate of increase, 0.09, finite rate of increase, 1.1, doubling time, 7.8 days and generation time, 41.1 days. Studies on feeding biology showed that C. nodiceps was oligophagous. The coccinellid fed and reproduced on prey species in two families, Diaspididae and Coccidae. Adults also fed on the coconut whitefly Aleurodicus cocois (Curtis) (Aleyrodidae) but no reproduction occurred on this prey. Aphididae, Psyllidae and Pseudococcidae were not fed upon.     

Identification of Armored Scales (Hemiptera: Diaspididae) on Bamboos in Korea

This work provides an identification key to adult females of six species and four genera of armored scales including a new record, Kuwanaspis hikosani (Kuwana) that may be found on bamboos (Poaceae: Arundinaria spp., Phyllostachys spp., Pseudosasa spp., and Sasa spp.) in Korea.     

A new species of armored scale, Mycetaspis ailynaomi (Hemiptera, Diaspididae, Aspidiotinae), associated with Mammea americana L. (Malpighiales, Calophyllaceae) from Puerto Rico

A new species of armored scale, Mycetaspis ailynaomi Dones and Evans is described and illustrated from specimens collected on mamey (Mammea americana) from Puerto Rico. A key to the species of Mycetaspis is provided.     

Phylogenetic congruence of armored scale insects (Hemiptera: Diaspididae) and their primary endosymbionts from the phylum Bacteroidetes

Insects in the sap-sucking hemipteran suborder Sternorrhyncha typically harbor maternally transmitted bacteria housed in a specialized organ, the bacteriome. In three of the four superfamilies of Sternorrhyncha (Aphidoidea, Aleyrodoidea, Psylloidea), the bacteriome-associated (primary) bacterial lineage is from the class Gammaproteobacteria (phylum Proteobacteria). The fourth superfamily, Coccoidea (scale insects), has a diverse array of bacterial endosymbionts whose affinities are largely unexplored. We have amplified fragments of two bacterial ribosomal genes from each of 68 species of armored scale insects (Diaspididae). In spite of initially using primers designed for Gammaproteobacteria, we consistently amplified sequences from a different bacterial phylum: Bacteroidetes. We use these sequences (16S and 23S, 2105 total base pairs), along with previously published sequences from the armored scale hosts (elongation factor 1alpha and 28S rDNA) to investigate phylogenetic congruence between the two clades. The Bayesian tree for the bacteria is roughly congruent with that of the hosts, with 67% of nodes identical. Partition homogeneity tests found no significant difference between the host and bacterial data sets. Of thirteen Shimodaira-Hasegawa tests, comparing the original Bayesian bacterial tree to bacterial trees with incongruent clades forced to match the host tree, 12 found no significant difference. A significant difference in topology was found only when the entire host tree was compared with the entire bacterial tree. For the bacterial data set, the treelengths of the most parsimonious host trees are only 1.8-2.4% longer than that of the most parsimonious bacterial trees. The high level of congruence between the topologies indicates that these Bacteroidetes are the primary endosymbionts of armored scale insects. To investigate the phylogenetic affinities of these endosymbionts, we aligned some of their 16S rDNA sequences with other known Bacteroidetes endosymbionts and with other similar sequences identified by BLAST searches. Although the endosymbionts of armored scales are only distantly related to the endosymbionts of the other sternorrhynchan insects, they are closely related to bacteria associated with eriococcid and margarodid scale insects, to cockroach and auchenorrynchan endosymbionts (Blattabacterium and Sulcia), and to male-killing endosymbionts of ladybird beetles. We propose the name "Candidatus Uzinura diaspidicola" for the primary endosymbionts of armored scale insects.     

Discovery of cryptic species among North American pine‐feeding Chionaspis scale insects (Hemiptera: Diaspididae)

Cryptic species are present in many animal groups and they may be best detected through large sample sizes collected over broad geographic ranges. Fine‐scale local adaptation has been hypothesized to occur in armoured scale insects (Hemiptera: Diaspididae) and a consequence of this process may be multiple cryptic species. We estimate species diversity of pine‐feeding Chionaspis scale insects across North America by inferring species boundaries using genealogical concordance across allele genealogies of two nuclear loci and one mitochondrial locus. Our ingroup sample includes 366 individual insects from 320 localities and 51 host species within the Pinaceae. We also conducted a morphological survey of all insect specimens and assigned them as either one of the two currently recognized pine‐feeding species, Ch. heterophyllae, Ch. Pinifoliae, or with undescribed morphology. Using maximum likelihood allele genealogies in a majority‐rule consensus to assess congruence, we conservatively detect ten species in this group. Most of these species are robust to alternative methods of genealogical inference (Bayesian inference, maximum parsimony) and a few are robust to the use of strict consensus to assess congruence. Species show both narrow and more widespread ranges where almost half of the individuals sampled belong to a single very widespread polyphagous species. © 2011 The Linnean Society of London, Biological Journal of the Linnean Society, 2011, 104 , 47–62.