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The first data on the biology of Ips stebbingi Strohmeyer, 1908 (coleoptera: curculionidae: Scolytinae), a pest of himalayan blue pine

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INTRODUCTION

Bark beetles (Coleoptera: Curculionidae: Scolytinae) comprise a large subfamily of insects, although only a small percentage of more than 6000 bark beetle species found worldwide are capable of causing significant economic impacts (Wood and Bright 1992). Among the most aggressive and notorious of these insects are species of the genus Ips DeGeer (Postner 1974; Wood and Bright 1992; Pfeffer 1995). Ips is one of the better-known bark beetle genera because of several species that are destructive to conifer forests and plantations (Chararas 1962; Furniss and Carolin 1992). Epidemic Ips populations can destroy thousands of hectares of forest (Furniss and Carolin 1992). In addition, some aggressive species, like Ips pini (Say) and Ips paraconfusus Lanier, kill the tops of healthy pine trees, which increase their susceptibility to attack by Dendroctonus spp. (Furniss and Carolin 1992).

According to faunistic estimates, Ips genus is composed of 37 species distributed in North America, Europe, Asia and introduced into Australia and Africa (Cognato 2015). Fourteen Ips species occur in the Palaearctic region (Knizek 2011), of which only eight are endemic to Asia showing remarkable variation of aggressiveness and host preference. Buhroo and Lakatos (2011) provided a detailed morphological key to three Himalayan Ips species and highlighted that these species can build up high population levels when there is excessive supply of breeding material and favourable environmental conditions. All three Himalayan species are phloeophagous and oligophagous (Wood and Bright 1992; Buhroo and Lakatos 2011) and like other conspecifics, they have evolved adaptations to exploit Pinaceae despite formidable defences that these tree species can mount (Franceschi et al. 2005).

The bark beetle, Ips stebbingi Strohmeyer, 1908, is distributed in the transition zone between the Palaearctic and the Oriental areas of Asia – Bhutan, China (Xinjiang), Uttar Pradesh, Himachal Pradesh, Kashmir, Pakistan and Afghanistan (Cognato 2015). Despite its aggressive habit of utilising Pinaceae host trees (Stebbing 1914; Wood and Bright 1992; Buhroo and Lakatos 2011), a little is known on the biology of I. stebbingi, which is a significant obstacle to success in terms of the development and application of pest management programme in the field conditions (Khanday et al. 2018).

Based on the field and laboratory observations, this study presents the first detailed data on the mating behaviour, maturation feeding, gallery patterns, life cycle and seasonal distribution of I. stebbingi on Himalayan blue pine (Pinus wallichiana) in Kashmir.

MATERIAL AND METHODS
Field studies

The biological data were obtained mainly in Nowpora village (33°61.078’N, 075°18.700’E and elevation, 5920 ft.) in district Anantnag, Jammu and Kashmir. Based on the preliminary surveys, severe bark beetle infestations were observed in the study area, inhabited by almost 30–40-year-old blue pine trees. The sampling procedure for monitoring the I. stebbingi activity was adopted as per the methods of Buhroo and Lakatos (2007). By using trap logs and other infested branches of standing trees (Beaver 1967; Buhroo and Lakatos 2007; Khanday and Buhroo 2015), observations of the various life stages were made once or twice weekly throughout the experimental period. These trap logs were cut periodically from April to October each year from 2017 to 2018. Further information was obtained by careful removal of bark sections both in the field and in the laboratory. The eggs found in maternal galleries were exposed carefully and counted in their individual chambers. For measurement of galleries (both maternal and larval), a digital caliper scale was used (Khanday and Buhroo 2015). The correlation between the length of maternal galleries and the number of eggs deposited by beetle pests was worked out using the method of Zhang et al. (1992).

At the experimental site, the entrance holes that the newly emerged beetles started grooving were marked on one main branch and some twigs of a declining pine tree during May–June. After the marking dates, 15 beetle entries were dissected each day and examined for eggs to determine female maturation feeding period as per the method of Buhroo and Lakatos (2007).

Laboratory rearing

Laboratory rearing of I. stebbingi was accomplished by placing infested logs (30–50 cm long and 5–10 cm in diameter) in three rearing boxes of similar design made up of glass with dimensions of 75 × 35 × 40 cm, 45 × 35 × 35 cm and 55 × 35 × 35 cm. The top face of each box was fitted with white muslin cloth. Each box could be opened from the top to facilitate exchange of logs. After every month, cut branches (20–40 cm long and 2–8 cm in diameter) from the host tree (P. wallichiana) were placed in the rearing boxes to induce fresh attack, 5–10 days before the emergence of adults in every generation. This enabled continuous rearing and examination of beetle development. A few infested logs were also debarked at regular intervals (10 days) to study various stages of the beetle under bark. Measurements of various developmental stages including egg, larva, pupa and adult were recorded. The development process and duration of beetle life stages were recorded and compared with the field results as per our earlier methods (Buhroo and Lakatos 2007; Khanday and Buhroo 2015).

Photography

Photographs were taken during the field study using Canon PowerShot SX60 camera fitted with macro lens (Raynox MSN-505, 37 mm; Yoshida Industry Co., Ltd, Tokyo, Japan). Analysis of digital images was done using ImageJ analysis software (Version 1.38f, 2006). For morphometric description of beetle life stages, images were taken with a Leica DFC295 camera (Leica Microsystems GmbH, Wetzlar, Germany) attached to a Leica M205A stereozoom binocular microscope having Leica Automontage Software (version 4.10). Spatial information regarding sample site was recorded in the form of latitude and longitude with the help of handheld global positioning system (GPS; Garmin eTrex 10, India).

Statistical analyses

Statistical analyses were performed using OriginPro software (version 2015). The gallery pattern and developmental stages were analysed using descriptive statistics (mean and standard deviation [SD]). The larval instars were separated from each other by head capsule measurements (Dyar 1890).

RESULTS
Mating behaviour

After host tree selection, the male of I. stebbingi was observed to form an initial gallery known as nuptial chamber (Fig. 1). Many female beetles (two to six) successively entered the nuptial chamber through the tunnel made by the male and after copulation, excavated their individual egg galleries radiating away from the central chamber (Fig. 1).

Figure 1.

Initial gallery (nuptial chamber)

Maturation feeding

Bark dissections for female maturation feeding (Fig. 2) showed that no eggs were collected from beetle entries on the first and second days; only 4/15 from 3-day-old entries yielded eggs, 9/15 from 4-day-old entries contained eggs, while 13/15 from 5-day-old entries contained eggs, but all the 15 of 6-day-old beetle entries contained eggs. This indicated that newly emerged females mostly fed and oviposited in 4–6 days in the field. The feeding sites were then continued into the maternal galleries.

Figure 2.

Female maturation period of Ips stebbingi

Gallery patterns

The gallery patterns are shown in Fig. 3. The average length of maternal galleries is 10.76 (±3.30 SD) cm (Fig. 3A, Tab. 1). Larval galleries radiate out from the maternal galleries. The average length of larval galleries is 3.83 (±0.63 SD) cm (Fig. 3B, Tab. 1). The larval tunnels are initially perpendicular to the maternal gallery, then radiate in different directions and can intersect (Fig. 3B). These tunnels are closely packed with the wood excreta passed out by the larvae during feeding.

Figure 3.

A typical gallery pattern of Ips stebbingi excavated in Pinus wallichiana: maternal galleries (A), larval galleries (B)

Measurement of gallery length

Variable n Mean (cm) Standard deviation (cm)
Maternal gallery length 30 10.76 3.30
Larval gallery length 30 3.83 0.63

Note: n = number of observations.

Data based on the length of the maternal gallery and the numbers of eggs deposited indicate that there is a positive correlation (y = 2.1686x – 2.315, r = 0.67) between the two variables (Fig. 4). The number of eggs increases linearly with maternal gallery length.

Figure 4.

Relationship between the length of maternal galleries (cm) and eggs per gallery in Ips stebbingi

Developmental stages
Egg

After copulation, each female bores out her tunnel, gnaws out little notches at the side and places an egg in each (Fig. 5). These notches are not made so symmetrically as in the case of the monogamous bark beetle species, with the eggs being laid usually more on one side than on the other. On an average, 23.16 (±7.03 SD) eggs were laid per female (Tab. 2). The eggs were about 0.72 (±0.03 SD) mm in length and 0.48 (±0.02 SD) mm in width (Tab. 2). The eggs hatched after an incubation period of 8–18 days (Tab. 3).

Figure 5.

Egg of Ips stebbingi

Egg deposition and measurement of developmental stages of Ips stebbingi

Variable n Mean (mm) Standard deviation (mm)
Eggs deposited 30 23.16 (number) 7.03
Egg length 30 0.72 0.03
Egg width 30 0.48 0.02
Larval length 30 3.00 0.13
Larval width 30 1.35 0.04
Pupal length 30 2.37 0.08
Pupal width 30 1.02 0.06
Female length 30 4.08 0.07
Female width 30 1.78 0.04
Male length 30 3.74 0.05
Male width 30 1.72 0.08

Note: n = number of observations

Developmental duration of Ips stebbingi

Duration of generations (days)
Stage First generation Second generation Third generation
Egg 9–15 8–14 8–18
Larva 35–45 30–38 30–40
Pupa 20–26 18–24 21–28
Adult 60–70 45–60 140–175
Total 124–156 101–136 199–261
Larva

The larva on hatching was apodous, wrinkled, minute, cylindrical and creamy white in colour (Fig. 6). The mature larvae reached an average length of 3.00 (±0.13 SD) mm and a width of 1.35 (±0.04 SD) mm (Tab. 2). On the basis of the data recorded in the field, five larval instars were observed (Tab. 4). The expected head capsule width of each instar was also determined by Dyar’s ratio (Dyar 1890), which asserts that the growth ratio remains constant between the moults. Each instar was progressively longer (length) than the preceding instar. Development from hatching to the prepupal larvae took 30–45 days (Tab. 3).

Figure 6.

Larva of Ips stebbingi

Table 4.

Comparison of observed (mean) and expected values of head capsule widths (mm) of the larvae of Ips stebbingi

Pupa

Pupation took place at the end of the larval galleries in pupal chambers. The pupae were soft and creamy white measuring 2.37 (±0.08 SD) mm in length and 1.02 (±0.06 SD) mm in width (Fig. 7, Tab. 2). The pupal stage lasted for 18–28 days (Tab. 3). After pupation, the adults fed for a short time in the phloem before emerging from the pupal chamber by tunnelling straight through the bark over it. After emergence, adults flew to suitable trees to produce the next generation.

Figure 7.

Pupa of Ips stebbingi

Adult

Body is oblong, shining and blackish brown (Fig. 8A, 8B). Antennae and legs are piceous brown. Head is smooth, shining on vertex, which has few scattered fine punctures on it; the front portion is flat and roughly punctate, and there are two tubercles placed transversely on the face (Fig. 8C). Elytra is truncate posteriorly; the declivous portion is concave and dull, sometimes slightly shining and finely punctate towards the bottom. The sides are furnished with four teeth, of which the upper one (1) is very small and set at a distance from 2; 2 is close to 3 and is larger than 1; 3 is the largest with a swollen head to it. Tooth 4 is prominent and sharp and smaller than 3 and set farther from 3 than 3 is from 2 on one side and a less distant from the lower margin (Fig. 8D). Elytra laterally and edges of declivity are fringed with long and yellow spiny hairs. The adult females had an average body length of 4.08 (±0.07 SD) mm and width of 1.78 (±0.04 SD) mm, and the body length of male was 3.74 (±0.05 SD) mm and its width was 1.72 (±0.08SD) mm (Tab. 2). The adults lived for 45–70 days (Tab. 3).

Figure 8.

Adult of Ips stebbingi: habitus, in dorsal view (A); habitus, in lateral view (B); head, in frontal view (C); abdominal declivity (D)

Seasonal incidence

The results of the present observations showed that I. stebbingi overwintered in adult stages in their respective galleries from the last week of October (Fig. 9). The adults remained inactive throughout the winter and resumed their activity from the second week of March onwards. The species produced three generations per year in Kashmir (Fig. 9). The first generation lasted from the last week of March to August, having a total life span of 124–156 days (Tab. 3). The second generation lasted from the second week of June to October, having a total life span of 101–136 days, while the third overwintering generation took 199–261 days and extended from August to May of the following year (Tab. 3). Laboratory observations also confirmed three generations with a little difference in developmental process and duration of insect stages.

Figure 9.

Seasonal distribution of Ips stebbingi on Pinus wallichiana in Kashmir (E – egg, L – larva, P – pupa, A – adult)

DISCUSSION

In the present study, the detailed biology of I. stebbingi has been worked out in the pine forest ecosystem of Kashmir Himalaya. The species shows a polygamous mating system. The male initiates gallery construction by excavating a nuptial chamber in the phloem. While doing this, the male also emits aggregation pheromones attracting conspecifics of both sexes to intensify the attack and subsequent colonisation at the site. After mating, each female carves her own tunnel radiating outwards from the central nuptial chamber and lays its eggs in individual niches along the sides of the gallery. Data based on the length of the maternal gallery and the numbers of eggs deposited indicate that there is a positive correlation between the two variables. The number of eggs increases linearly with maternal gallery length. The decline of maternal gallery length and oviposition with increasing attack density can be partially due to the intraspecific competition. For the first time, this study described the duration of developmental passage of I. stebbingi through individual stages and determined the number of instars using the distribution of head capsule widths (Dyar 1890; Caltagirone et al. 1983). The galleries of I. stebbingi contain all developmental stages from egg to adult, whose general qualitative characteristics are similar to those described for other scolytines (Wood 2007; Jordal 2014). This species overwinters in the adult stage under the bark. In the spring, adults spend a period of time (maturation feeding) under the bark, often in their galleries before dispersal, and the newly emerged females mostly feed and oviposit in 4–6 days in the field.

The present study results show that I. stebbingi produces three generations per year in Kashmir in contrast to Ips typographus having mostly univoltine generation in Europe (Ohrn 2012). The bark beetle attacks have increased during the past few decades in the northwestern Himalaya as a result of poor forest management systems and climate change. Currently, the climate changes are predicted to significantly affect the frequency and severity of disturbances in the forest ecosystems (Bentz et al. 2010). Temperature can be the most crucial factor in controlling insect life in the temperate forests. In response to higher temperatures and drought, the bark beetles, in general, and I. stebbingi, in particular, may show improved performance in trees and have a synergistic negative effect on the pine forest as water shortage weakens the host defence. The increased temperature will also promote earlier emergence in spring and faster preimaginal development. Under adverse conditions of climate change (prolonged droughts, floods and storms), the forest stands are susceptible to increased stresses and damages mostly by bark beetles (McNichol et al. 2021).

We can expect higher population outbreaks of I. stebbingi doing more damage to a less-defensive host tree in a future warmer climate. The forest management system has to be planned in view of increased wind damages, severe drought and higher population levels of bark beetles in the Himalayan region.

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