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Article

Dynamics of the Adult Litchi Stink Bugs, Tessaratoma papillosa (Drury) (Hemiptera: Tessaratomidae), on Four Urban Tree Species in Taipei City

by
Meng-Hao Hsu
1,
Yu-Ping Tsai
2,3 and
Liang-Jong Wang
2,*
1
Taimalee Research Center, Taiwan Forestry Research Institute, Ministry of Agriculture, Taitung County 963001, Taiwan
2
Taiwan Forestry Research Institute, No. 67, 6 Floor, Sanyuan St., Zhongzheng Dist., Taipei City 10079, Taiwan
3
Department of Horticulture and Landscape Architecture, National Taiwan University, Taipei City 10617, Taiwan
*
Author to whom correspondence should be addressed.
Forests 2025, 16(4), 601; https://doi.org/10.3390/f16040601
Submission received: 5 February 2025 / Revised: 28 February 2025 / Accepted: 27 March 2025 / Published: 29 March 2025
(This article belongs to the Section Forest Health)
Browse Figures
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Abstract

:
A two-year study, starting at the beginning of 2018, in Taipei City, was conducted to investigate the population dynamics of adult litchi stink bugs, Tessaratoma papillosa (Drury), in the trees on roadsides and in urban greenspaces, namely, Litchi chinensis Sonn., Dimocarpus longan Lour., Sapindus mukorossi Gaertn. and Koelreuteria henryi Dümmer, and deduce the relationship between the population dynamics and the photoperiod or climatic factors. The winter diapause of the adults might play an important role in the population dynamics and affect their movement among various host species. However, we also found that a warm winter and early-summer rainy season may affect the population size of stink bugs. There were significantly more adults in 2018 than in 2019 when comparing the periods from March to December of the two years. In a comparison of the four tree species, there were significantly more stink bugs on the longan trees than on the other trees, especially during winter. The results of this study might allow decision-makers to apply better control strategies based on the correct timing and targeting of tree species of this invasive pest in Taiwan.

1. Introduction

The litchi stink bug, Tessaratoma papillosa (Drury), is a major pest on litchi and longan trees in China and Southeast Asia [1,2,3,4]. Probably through multiple introductions, this invasive species has spread over low-elevation areas of Taiwan since 2009 [5,6,7]. The life cycle of litchi stink bugs is more than one year, and they can move among various host plants [8,9]. The species appears on the plants of agricultural and non-agricultural areas, and it has become a concern to the general public in Taiwan. It not only affects the fruit production of litchi and longan trees, but also causes ulcers on human skin [5,10] from the secretions of the scent glands of the stink bugs [11], which possibly approach from nearby common urban trees, Koelreuteria henryi Dümmer [5].
According to the official annual reports [12,13], Taiwan has held several inter-ministerial meetings to enforce coordination of nationwide monitoring and effective control measures for the mitigation of agricultural production loss and for the appeasement of complaints from worried urban dwellers due to the public health issues caused by these stink bugs. The health issues include the noxious odor from the stink bugs and the lesions caused by their caustic secretions. The symptoms of lesions were similar to those in the case reports from Brazil by other species of stink bugs [14]. In Taiwan, many complaints were about accidental discoveries of green eggs stuck on laundry, windows, walls, etc., by gravid female bugs intruding into households during oviposition season [15]. The local city governments advised caution and recommended staying clear of litchi stink bugs, even suggesting that victims should go to dermatologists immediately for proper care. Moreover, several control measures were taken against the bugs, including pesticide sprays, the release of egg-parasitic wasps, and egg removal with government financial incentives [5,6,10,13,15,16].
The Taiwanese rain tree, K. henryi, is a species endemic to Taiwan and prevalently planted along roadsides and in greenspaces for appreciating the beauty of the yellow flowers and subsequent reddish fruits in autumn and early winter [15,16,17]. Based on the statistics of the Taipei City Government, nearly 9% of the 94,829 registered street trees are K. henryi (accessed on 4 February 2025, https://geopkl.gov.taipei), and the nymphs of litchi stink bugs were found on their new green shoots during spring and early summer [15,16]. The longan, Dimocarpus longan Lour., is usually not considered a street tree, but it is a popular shade tree in private gardens or greenspaces beside buildings. The people under the shade of a blooming longan tree could frequently become irritated by the volatile malodor from litchi stink bugs. The two trees mentioned above, Taiwanese rain trees and longans, are ubiquitous in Taipei City and other lowlands of Taiwan [15]. The litchi tree, Litchi chinensis Sonn., and the soapberry tree, Sapindus mukorossi Gaertn., are rather sporadically distributed in Taipei City; however, there are many plantations of these trees in the central and southern lowlands of Taiwan [18,19]. Sapindus mukorossi used to be a source of ingredients for traditional cleansing products [19], but currently, it is planted in parks or at roadsides as an ornamental tree for a scenic view of winter foliage turning yellow. Both litchis and longans are evergreens, while S. mukorossi and K. henryi are deciduous. This difference in the morphological and physiological characteristics between the tree species may affect the overwintering of litchi stink bugs and their dispersal as well as population dynamics.
In the past, most of the research on the population dynamics or monitoring of the litchi stink bug focused on only one of the crops, either litchis or longans [6,20,21,22,23]. Investigation and research on a single host plant cannot encompass the whole picture of the population dynamics of the pest. Therefore, in this study, four main host plant species belonging to the family Sapindaceae in the non-agricultural district of Taipei City were investigated monthly throughout a year and again the next year. Finally, we aimed to pinpoint the key environmental factors of the population dynamics and discuss the relationship between the factors, litchi stink bugs, and host plants.

2. Materials and Methods

This study was carried out in Taipei City, Taiwan at the locations of three roadsides, three riverbanks, three parks, and other areas with lines of four single or mixed main host plants, that is, Litchi chinensis Sonn., Dimocarpus longan Lour., Sapidus mukorossi Gaertn., and Koelreuteria henryi Dümmer, that might be infested by Tessaratoma papillosa (Drury). Table 1 shows the locations and selected trees for monthly investigation of adult litchi stink bugs. Before investigation, the specimens of adult T. papillosa were collected as voucher specimens and identified by L. J. Wang, an insect taxonomist in Taiwan. The researchers in the present study were trained and could recognize and identify T. papillosa in the field without difficulty.
One hundred and fourteen trees of four species belonging to the family Sapindaceae in the non-agricultural district of Taipei City were investigated monthly in the daytime over two years, from January 2018 to December 2019. The occurrences of adult T. papillosa on L. chinensis (n = 21), D. longan (n = 30), S. mukorossi (n = 33), and K. henryi (n = 30) were recorded via the naked eye or with a telescope (Pentax Papilio II, 8.5 × 21, RICOH IMAGING Company, LTD., Tokyo, Japan), and aided with a flashlight (MT14, Ledlenser GmbH & Co.KG, Solingen, Germany) at high luminosity (up to 1000 lumens) when necessary. Only adults were recorded because their sizes were large enough to be spotted from under the tree. The distance of observation was from 0 to approximately 8 m, depending on the tree height. In the meantime, we recorded the number of adults in reproductively premature or mature phases, and the number of mating individuals within 20 min per tree. Therefore, approximately 10~15 trees were investigated in a day, and we conducted the sampling twice a week. An investigation of all 114 trees could be completed monthly. We could distinguish the premature adults from mature ones by their cuticle colors, residual white wax, and behaviors (Figure 1 and Figure 2).
Monthly mean temperatures, rainfall, rainy days, sunshine hours, and the day lengths of the middle of the month (photoperiod) were obtained from the Central Meteorological Administration for analysis of the correlation of the population dynamics of the insects with meteorological factors. For detailed analysis, the daily minimum temperatures of the winter months including January and February, and the daily mean temperatures of May were collected. The data of the means of the bugs on four tree species and correlation were analyzed using the software, SPSS Statistics for Windows version 17.0 (Released 2008, SPSS Inc., Chicago IL, USA), which is suitable for biostatistics [24]. We compared the difference in the number of adult T. papillosa between two survey years by the paired sample t-test. One-way analysis of variance was performed to compare the numbers of adult bugs among the four main host plant species, then Fisher’s least significant difference (LSD) post hoc test at a 95% confidence level was used to compare means. The correlation analyses were used to test the relationships between the numbers of T. papillosa and each climatic factor separately. Five climatic factors, including photoperiods, mean temperature, rainfall, rainy days, and sunshine hours, were analyzed in this study. Moreover, paired sample t-tests were applied to examine the temperature differences between the two years in the specific periods of time.

3. Results

3.1. Tessaratoma papillosa (Drury) on Different Tree Species

The two-year study in Taipei City shows that the peak of the monthly mean numbers of adult litchi stink bugs first appeared on the litchi trees in April 2018, and in the same month, the mean numbers of bugs increased drastically on the longan trees. Then, the mean number of bugs reached their peaks on Sapindus mukorossi Gaertn. in May, and on both Koelreuteria henryi Dümmer and the longan trees in June. In 2019, the sequence of the appearance of population peaks of litchi stink bugs on different tree species was the same as the previous year except for the longan trees, whose bug population drastically decreased after the peak in April (Figure 3A).
Table 2 indicated that all sampled longan trees (100% infestation) had been infested with adult litchi stink bugs for at least one month each year. Moreover, 86.67% of K. henryi and 81.82% of S. mukorossi had infestation for both years, respectively. However, approximately 40% of litchi trees were never infested with adult bugs. The comparison of the annual mean numbers of adult stink bugs is indicated in Table 3. There were significantly more bugs on the longan trees (mean ± SE = 55.73 ± 3.27 and 39.47 ± 10.42 bugs/tree in 2018 and 2019, respectively) than on other species of trees (mean = 6.57~25.1 bugs/tree) in the same year.

3.2. Adults in Reproductively Premature and Mature Phases

In the study, we noted that the first mating pair appeared in early spring or late winter (March 2018 and February 2019) (Figure 3C), and newly emerged adults began to appear in early summer (June 2018 and June 2019) (Figure 3D). The appearance of the first mating pair indicates the termination of the premature phase and resumption of the active, reproductively mature phase. The number of mating pairs peaked in April on the litchi and longan trees and soon declined in May (Figure 3C). Some adults of the population very likely moved to K. henryi and S. mukorossi, and mated there in May and June (Figure 3B,C; Table 3). The percentages of monthly recorded mating individuals on K. henryi were high even in late summer, for example, 85% and 77.27% in August 2018 and July 2019, respectively (Table 4). In terms of the percentage of mating individuals from the total number of recorded adults calculated for each tree species separately, the percentages of mating pairs in litchi were highest in both years, whereas the ones in longan were lowest (Table 4).
Two generations of adult bugs overlapped during the second half of a year; however, the number of adults on trees drastically declined after June (Figure 4A), probably mainly because of increased numbers of deaths in aging, reproductively mature adults. However, the number of newly emerged adults increased after June and that is clearly shown in the curves of the longan trees in Figure 3D. The second peak in the number of bugs on the longan trees in August 2019 (Figure 3A,D) mostly comprised adults in the reproductively premature phase. In terms of longan trees, the maximum percentage of monthly recorded premature adults in a year was 97.65% and 100% in December 2018 and November 2019, respectively. Most of the annually recorded premature adults on the 114 sampled trees were on the longan trees, i.e., 399 (78.39%) and 415 (87.74%) in 2018 and 2019, respectively (Table 5). In December 2018, 83 premature adults were detected under the leaves of all sampled longan trees, of which 35 individuals were found in one tree and stayed there until next January. In terms of the percentage of reproductively premature adults from the total number of recorded adults calculated for each tree species separately, the percentages of premature adults in longan were highest in both years, whereas the ones in deciduous tree species, including K. henryi and S. mukorossi, were lower (Table 5).

3.3. Relationship Between Population and Photoperiod or Climate

Although data from simple linear correlation or difference tests may not be enough to show conclusive results for the influence of climate factors on the seasonal population dynamics, we would still like to display our findings based on the two consecutive years. Although there was no significant difference between the mean numbers of adult bugs in the two years (mean ± SE = 273.42 ± 79.00 and 208.00 ± 55.16 bugs/tree in 2018 and 2019, respectively; df = 11, t = −1.99, p = 0.07) as a whole, there was a significant difference between the mean numbers of adult bugs when comparing the same periods from March to December of the two years (mean ± SE = 320.70 ± 87.54 and 229.70 ± 64.30 bugs/tree in 2018 and 2019, respectively; df = 9, t = 2.69, p = 0.03). To verify the key factors that affected the population dynamics of T. papillosa, we examined the monthly recorded population dynamics of adult litchi stink bugs with photoperiod, and climatic factors including temperature, rainfall, rainy days, and sunshine hours, as depicted in Figure 4. We found that each of the two annual patterns of the population dynamics of the litchi stink bugs significantly showed a similar trend with the photoperiod (r = 0.87 and r = 0.81 for 2018 and 2019, respectively). However, in terms of the four climatic factors analyzed in this study, there was no consistency when comparing them between the two years (for correlation coefficients, see Table 6). For example, the correlation coefficient between the population dynamics of adult T. papillosa recorded monthly and the rainfall was −0.27 without statistical significance in 2018; however, the coefficient was 0.59 with significance in 2019 (Table 6).
We noticed some divergence between the two lines of the monthly mean temperatures shown in Figure 4C. For an in-depth analysis, we examined the daily minimum temperatures (Figure 5). There was a significant difference between the temperatures of the winter months of the two years (mean ± SE = 14.05 ± 0.42 and 16.40 ± 0.24 °C in 2018 and 2019, respectively; df = 58, t = −4.46, p < 0.001). It means that winter (from January 1 to February 28) 2018 was significantly colder than winter 2019. In fact, there were two cold surges (minimum temperature < 10 °C) in 2018, while there were none in 2019 (Figure 5). We suppose that the significantly warmer winter may be the cause of early termination of diapause, hence the increased numbers of reproductively mature adults on the longan trees (see the yellow line of Figure 3B) as well as the recorded adults on all sampled trees (see the black line of Figure 4A) in the first two months of 2019.
The second divergence of the lines, depicted in Figure 4C, of monthly mean temperatures appeared in May, and a significant difference can be pinpointed when carrying out a detailed analysis of daily mean temperatures of the same months between the two years (mean ± SE = 28.19 ± 0.45 and 24.99 ± 0.55 °C in May 2018 and May 2019, respectively; df = 30, t = 6.46, p < 0.001). We also observed a divergence in the lines in May between the two years presented in Figure 4A (number of adults on all sampled trees), and the sharp decline in the mean number of bugs on the longan trees recorded in 2019 (Figure 3A). Moreover, there was significantly higher rainfall during the months around the early-summer rainy season (plum rain), i.e., from April to July, in 2019 than that during the same period in 2018 (mean ± SE = 102.93 ± 33.55 and 327.43 ± 74.24 mm in 2018 and 2019, respectively; df = 3, t = −3.91, p = 0.03) (Figure 4D).

4. Discussion

According to Saulich and Musolin (2012) [25], the majority of pentatomid species (Pentatomidae) overwinter as adults. Such a distribution of the overwintering stages approximately corresponds to the situation observed in the entire order, where species diapausing as adults clearly prevail. This is also the case of our study on Tessaratoma papillosa (Drury), which mainly hides in longan trees to overwinter [8]. The start and termination of the reproductive diapause of insects are supposed to affect the monthly recording of the population dynamics. She and Pan (1993) [26] observed that the annual population dynamics of T. papillosa always consisted of the alternate appearance of two stages, that is, the reproductively active stage with a large population size followed by the reproductive diapause stage with a low population density. They also tried to apply five grades of ovary development by dissection of female adult T. papillosa to predict whether an outbreak would occur in the following litchi crop season. In this study, the relationship of seasonal population dynamics of this bug with photoperiod or some climatic factors was based on only two years of data; therefore, it is too early to draw any decisive conclusions for this invasive pest. Moreover, many other environmental factors such as natural enemies, plant phenology, typhoons, etc., could be important factors that would greatly affect the population dynamics. Especially, the photoperiodic regulation of insect seasonal cycles is much more complicated and should be verified by special experiments such as dissection of both sexes to determine the development of reproductive organs and fat bodies [27]. Moreover, premature adults probably tend to leave the original host plants, especially litchi, Koelreuteria henryi Dümmer, and Sapindus mukorossi Gaertn., wherein they emerged from the last nymphal stage. Aggregation of premature adults could be detected in winter under the leaves of some longan trees. In January 2019, we discovered a case of more than 936 adults staying under the leaves of a longan tree with no distinct odor in central Taiwan. On the same tree, seven litchi stink bugs were found inactive under one leaf (unpublished data, Hsu). Beside litchi orchards in southern China, premature adults were found on nearby miscellaneous trees, especially longan trees [8]. Based on a study in a mixed orchard of litchi and longan trees in central Taiwan, Wu et al. (2021) [28] pointed out that significantly more adults were recorded on the longan trees than on the litchi trees during winter. Dissection of the adult stink bugs collected in January and March revealed remarkable developments of their ovaries and testes from premature to mature [28].
According to one study of longan trees, litchi stink bugs started to mate in February, peaked during March and April, and then the number of mating pairs drastically declined in May [6]. The results resemble those of our study; however, Wu et al. (2020) [6] also indicated that no mating activity was observed for eight months after May. In our study, the number of mating pairs on S. mukorossi and K. henryi peaked in May or June 2018 and 2019. The differences in the findings may be due to the investigation sites; however, most importantly, the movement of the adult bugs among various host plant species with distinct phenology such as blooming or other different morphological and physiological characteristics between deciduous and evergreen trees should be considered. Therefore, the relationship between possible factors and the population dynamics of the polyphagous insects may be better comprehended only when investigating all the main species of sympatric host plants in the same period, since a coevolved phenology might exist between the insect and its host plants.
The size of the population may be impacted by the rainfall, especially continuous rainfall such as that in the early summer rainy season in 2019 of our study. The population dynamics investigated in Hainan, China in 2009 [29] had the similar result of a large number of stink bugs dying rigidly on the litchi trees because of the infestation of entomogenous fungi after a large amount of precipitation. We reiterate that, although the size of the population was impacted by precipitation, it did not alter the pattern or trend of the dynamics. Therefore, there were no consistencies in the relationship between the monthly population dynamics of the litchi stink bugs and the yearly fluctuation in rainfall or the number of rainy days, when compared to those between 2018 and 2019. Typhoons can destroy street trees and usually arrive with a large amount of precipitation; however, only one typhoon made landfall on the southern tip of Taiwan and none hit Taipei City or nearby northern Taiwan during our investigation. Therefore, the storm’s effect on the insect population in street trees might be negligible in the study.
In our study, we found that the minimum temperature of the winter is probably an additional key factor in the population dynamics of the stink bugs. It may lengthen the diapause because of the arrival of cold surges, and this is the case in 2018 of our study, or it may terminate the winter diapause early and enter into the reproductively mature phase due to a warm winter, as in 2019. According to Menzel and Waite (2005) [30] and Chang et al. (1997) [18], cool temperatures stimulate shoot initiation and promote flowering in litchi plants. Therefore, it may merit further analysis of the relationship between the population dynamics and the blooming of the four host plant species.
Although a recent study has revealed some value of the implications for utilization as functional food ingredients [31], in Taiwan, this invasive insect is still considered a notorious pest. The results of this study provide great strategic value for decision-making in relation to timing and target plants for better control of this invasive pest in Taiwan. After the population peak in June, the adults probably have a tendency to leave the original host plants, especially Litchi chinensis Sonn., S. mukorossi, and K. henryi, while adult aggregation can be discovered on some longan trees during winter. Furthermore, the peak number of adults and mating pairs on K. henryi, which is one of the most popular street trees in Taiwan, always appeared after the other three species of host plants and was still abundant in summer.

5. Conclusions

Timing and identifying the target plants are critical factors for applying effective control measures against this invasive pest in Taiwan. The results of this study could provide valuable information for decision-making for better control of Tessaratoma papillosa (Drury) on the trees in the city as well as in orchards. Through simultaneous investigations of the four main sympatric host plants in this study, the issues of the polyphagous and monovoltine insect, T. papillosa, can be better addressed, including the relationship between the population and some possible factors as well as the movement between different host plant species. However, many factors were not analyzed in this study including typhoons, natural enemies, plant phenology, extreme climate, etc. Therefore, further research and the collection of more data from longer periods of time are necessary for a precise interpretation of the mechanism of seasonal dynamics.

Author Contributions

Conceptualization, L.-J.W.; Methodology, M.-H.H., Y.-P.T. and L.-J.W.; Software, Y.-P.T.; Validation, M.-H.H. and L.-J.W.; Formal analysis, M.-H.H. and Y.-P.T.; Investigation, M.-H.H. and L.-J.W.; Resources, M.-H.H. and L.-J.W.; Data curation, M.-H.H. and Y.-P.T.; Writing—original draft, M.-H.H.; Writing—review & editing, M.-H.H. and L.-J.W.; Visualization, M.-H.H.; Supervision, L.-J.W.; Project administration, M.-H.H.; Funding acquisition, M.-H.H. All authors have read and agreed to the published version of the manuscript.

Funding

This study was supported by the granted projects No. 107AS-10.3.1-FI-G1 and No. 108AS-10.3.1-FI-G1 from the Taiwan Forestry Research Institute.

Data Availability Statement

The data presented in this study are available on request from the corresponding author.

Acknowledgments

We thank Yueh-Lin Yang for her assistance with the field investigation and appreciate her comments to improve the earlier version of the draft.

Conflicts of Interest

The authors declare no conflicts of interest. The funders had no role in the design of the study; in the collection, analyses, or interpretation of data; in the writing of the manuscript; or in the decision to publish the results.

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Figure 1. Adults in the reproductively premature phase of Tessaratoma papillosa. (A) A newly emerged adult fed on a pedicel bearing a young longan fruit, preparing for overwintering. (B) During winter, inactive adults could be detected under longan leaves individually. Their ventral bodies were covered by rather intact, thick layers of white wax. (C) Sometimes, several adults could be spotted under one longan leaf. The dark brown cuticles of the adults in this phase look brand new and are devoid of black marks.
Figure 1. Adults in the reproductively premature phase of Tessaratoma papillosa. (A) A newly emerged adult fed on a pedicel bearing a young longan fruit, preparing for overwintering. (B) During winter, inactive adults could be detected under longan leaves individually. Their ventral bodies were covered by rather intact, thick layers of white wax. (C) Sometimes, several adults could be spotted under one longan leaf. The dark brown cuticles of the adults in this phase look brand new and are devoid of black marks.
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Figure 2. Adults in the reproductively mature phase of Tessaratoma papillosa. In this phase, adults are active and vigilant. (A) On Koelreuteria henryi, a mating female displayed a defensive response by releasing a jet of fluid from the anus when an investigator was approaching. (B) On Dimocarpus longan, a mating female ejected volatile secretion like a spray of machine-gun bullets from the openings of metathoracic scent glands when disturbed by a rod. The cuticles of mature adults are orange and become more exposed due to less residual wax remaining with age. (C) Black marks were apparent on the body of an aged male feeding on a branch of K. henryi. (D) A female laid eggs on a leaf of K. henryi.
Figure 2. Adults in the reproductively mature phase of Tessaratoma papillosa. In this phase, adults are active and vigilant. (A) On Koelreuteria henryi, a mating female displayed a defensive response by releasing a jet of fluid from the anus when an investigator was approaching. (B) On Dimocarpus longan, a mating female ejected volatile secretion like a spray of machine-gun bullets from the openings of metathoracic scent glands when disturbed by a rod. The cuticles of mature adults are orange and become more exposed due to less residual wax remaining with age. (C) Black marks were apparent on the body of an aged male feeding on a branch of K. henryi. (D) A female laid eggs on a leaf of K. henryi.
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Figure 3. The monthly mean number of Tessaratoma papillosa on four different trees, including Litchi chinensis, Dimocarpus longan, Sapindus mukorossi and Koelreuteria henryi, from January 2018 to December 2019. (A) Average number of adults per tree. (B) Average number of adults in reproductively mature phase per tree. (C) Average number of mating individuals per tree. (D) Average number of adults in reproductively premature phase per tree.
Figure 3. The monthly mean number of Tessaratoma papillosa on four different trees, including Litchi chinensis, Dimocarpus longan, Sapindus mukorossi and Koelreuteria henryi, from January 2018 to December 2019. (A) Average number of adults per tree. (B) Average number of adults in reproductively mature phase per tree. (C) Average number of mating individuals per tree. (D) Average number of adults in reproductively premature phase per tree.
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Figure 4. Comparison of (A) Monthly recorded adult Tessaratoma papillosa on all sampled trees (four tree species, n = 114) with (B) photoperiod (day length of the middle of the month), (C) temperature, (D) rainfall, (E) rainy days, and (F) sunshine hours.
Figure 4. Comparison of (A) Monthly recorded adult Tessaratoma papillosa on all sampled trees (four tree species, n = 114) with (B) photoperiod (day length of the middle of the month), (C) temperature, (D) rainfall, (E) rainy days, and (F) sunshine hours.
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Figure 5. Daily minimum temperature in Taipei City in the winters of 2018 and 2019 from January 1 to February 28. The arrival of a continental air mass brings cold and wet weather to Taiwan with the temperature dropping to 14 °C. Moreover, according to the Central Weather Bureau, a cold surge warning will be issued as mercury dips to 10 °C.
Figure 5. Daily minimum temperature in Taipei City in the winters of 2018 and 2019 from January 1 to February 28. The arrival of a continental air mass brings cold and wet weather to Taiwan with the temperature dropping to 14 °C. Moreover, according to the Central Weather Bureau, a cold surge warning will be issued as mercury dips to 10 °C.
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Table 1. Locations of the trees (n = 114) in Taipei City, Taiwan, selected for monthly investigation of adult Tessaratoma papillosa from January 2018 to December 2019.
Table 1. Locations of the trees (n = 114) in Taipei City, Taiwan, selected for monthly investigation of adult Tessaratoma papillosa from January 2018 to December 2019.
Survey SitesDetailsNo. Tree *GPS
Intersection of Chende
and Jiangtan Roads, Shulin		A sidewalkK16-2025.084, 121.524
Zhishanyan, Section 1 of
Zhicheng Road, Shulin		A sidewalkS1-1825.101, 121.533
Section 7 of Civic
Boulevard, Nangang		A sidewalkS19-3325.051, 121.591
Sanjiao Ferry Pier,
Shulin		The right bank of the Keelung RiverK1-12,
D4-8		25.081, 121.517
Shuangchi Riverside Park,
Shulin		The left bank of the Weishuangchi RiverK21-23,
D9-16, L1-6		25.099, 121.524
The bank along Wenlin
North Road, Shulin		The right bank of the Weishuangchi RiverK24-30,
D17-27		25.100, 121.521
Chiang Kai-Shek
Residence, Shulin		A city park on the foot of the hillL7-2125.093, 121.532
Hualing Park,
Shulin		A small greenspace in a local communityK13-1525.082, 121.520
Jiantan Park,
Shulin		A small greenspace on the foot of the hillD1-225.079, 121.525
Jiantanshan,
Shulin		A trail on a hillD3,
D28-30		25.080, 121.526
* No. Tree: L, D, S, and K mean four species of trees, including Litchi chinensis (n = 21), Dimocarpus longan (n = 30), Sapindus mukorossi (n = 33), and Koelreuteria henryi (n = 30), respectively. The selected trees were numbered for monthly investigation.
Table 2. The annual infestation of adult Tessaratoma papillosa on four different tree species in 2018 and 2019.
Table 2. The annual infestation of adult Tessaratoma papillosa on four different tree species in 2018 and 2019.
Tree Speciesn a20182019
% Trees Infested b% Trees Infested b
K. henryi3086.67%86.67%
S. mukorossi3381.82%81.82%
L. chinensis2161.90%57.14%
D. longan30100.00%100.00%
a Number of trees analyzed. b Infested means the sampled tree had been detected with adult litchi stink bugs for at least one monthly investigation in a year. For example, nine litchi trees were never found infested in 12 monthly investigations of 2019, % tree infested = (21 − 9)/21 = 57.14%.
Table 3. Comparisons of the occurrence of adult Tessaratoma papillosa on four different tree species in 2018 and 2019.
Table 3. Comparisons of the occurrence of adult Tessaratoma papillosa on four different tree species in 2018 and 2019.
Tree Speciesn adf20182019
Mean b ± SEFpLSDMean b ± SEFpLSD
(1) K. henryi30325.10 ± 5.15 b6.37 ***0.0005(4) > (1)
(4) > (2)
(4) > (3)		16.47 ± 3.14 b4.40 **0.006(4) > (1)
(4) > (2)
(4) > (3)
(2) S. mukorossi3321.76 ± 3.99 b19.85 ± 3.67 b
(3) L. chinensis216.57 ± 3.02 b7.76 ± 2.79 b
(4) D. longan3055.73 ± 3.27 a39.47 ± 10.42 a
a Number of trees analyzed. b Number of insects recorded per tree annually; means were calculated for all inspected trees. The inside column means followed by the same letter do not differ significantly at p > 0.05 (one-way ANOVA followed by an LSD test). ** p < 0.01, *** p < 0.001.
Table 4. The numbers of mating Tessaratoma papillosa recorded on four tree species.
Table 4. The numbers of mating Tessaratoma papillosa recorded on four tree species.
Tree
Species		20182019
No.
of Mating
Individuals		% aMax. % of Monthly Records b (Month)% cNo.
of Mating
Individuals		% aMax. % of Monthly Records b (Month)% c
K. henryi21423.46%85.00% (August)28.42%17820.94%77.27%
(July)		36.03%
S. mukorossi21823.90%41.05%
(June)		30.36%29434.59%73.55% (May)44.89%
L. chinensis444.82%13.99%
(April)		31.88%829.65%14.73% (April)50.31%
D. longan43647.80%90.00% (Mar)26.08%29634.82%100.00% (February)25.00%
Total912100.00%-27.80%850100.00%-34.05%
a (Number of mating bugs found on one of the tree species ÷ total number of the mating bugs recorded on all sampled trees of the four tree species) in a year × %. b Maximum of (monthly recorded mating bugs on one of the tree species ÷ number of mating bugs found on all sampled trees in the same month) × %. c (Number of mating bugs found on one of the tree species ÷ total number of the bugs recorded on the sampled trees of the same tree species) in a year × %.
Table 5. Number of adult Tessaratoma papillosa in the reproductively premature phase recorded on four different tree species.
Table 5. Number of adult Tessaratoma papillosa in the reproductively premature phase recorded on four different tree species.
Tree
Species		20182019
No. of
Premature Adults		% aMax. % of Monthly Records b (Month)% cNo. of
Premature Adults		% aMax.% of Monthly Records b (Month)% c
K. henryi499.62%21.21%
(January)		6.51%245.07%12.28%
(July)		4.86%
S. mukorossi397.66%25.49%
(July)		5.43%163.38%8.92%
(September)		2.44%
L. chinensis224.32%28.57%
(February)		15.94%183.81%5.93%
(August)		11.04%
D. longan39978.39%97.65%
(December)		23.8%41587.74%100.00% (November)35.05%
Total509100.00%-15.51%473100.00%-18.95%
a (Number of premature adults found on one of the tree species ÷ total number of premature adults recorded on all sampled trees of the four tree species) in a year × %. b Maximum of (monthly recorded premature adults on one of the tree species ÷ number of premature adults found on all sampled trees in the same month) × %. c (Number of premature adults found on one of the tree species ÷ total number of bugs recorded on the sampled trees of the same tree species) in a year × %.
Table 6. The preliminary correlation analysis between the number of adult Tessaratoma papillosa recorded monthly on all sampled trees (n = 114) and photoperiods or climatic factors based on two-year data.
Table 6. The preliminary correlation analysis between the number of adult Tessaratoma papillosa recorded monthly on all sampled trees (n = 114) and photoperiods or climatic factors based on two-year data.
Yearn aPhotoperiodMean TemperatureRainfallRainy DaysSunshine Hours
2018120.87 b**0.66 b*−0.27 b−0.61 b*0.39 b
2019120.81 b**0.39 b0.59 b*0.69 b*−0.18 b
a Number of months analyzed. b Pearson’s correlation coefficient. * Correlation is significant at the 0.05 level (2-tailed). ** Correlation is significant at the 0.01 level (2-tailed).
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Hsu, M.-H.; Tsai, Y.-P.; Wang, L.-J. Dynamics of the Adult Litchi Stink Bugs, Tessaratoma papillosa (Drury) (Hemiptera: Tessaratomidae), on Four Urban Tree Species in Taipei City. Forests 2025, 16, 601. https://doi.org/10.3390/f16040601

AMA Style

Hsu M-H, Tsai Y-P, Wang L-J. Dynamics of the Adult Litchi Stink Bugs, Tessaratoma papillosa (Drury) (Hemiptera: Tessaratomidae), on Four Urban Tree Species in Taipei City. Forests. 2025; 16(4):601. https://doi.org/10.3390/f16040601

Chicago/Turabian Style

Hsu, Meng-Hao, Yu-Ping Tsai, and Liang-Jong Wang. 2025. "Dynamics of the Adult Litchi Stink Bugs, Tessaratoma papillosa (Drury) (Hemiptera: Tessaratomidae), on Four Urban Tree Species in Taipei City" Forests 16, no. 4: 601. https://doi.org/10.3390/f16040601

APA Style

Hsu, M.-H., Tsai, Y.-P., & Wang, L.-J. (2025). Dynamics of the Adult Litchi Stink Bugs, Tessaratoma papillosa (Drury) (Hemiptera: Tessaratomidae), on Four Urban Tree Species in Taipei City. Forests, 16(4), 601. https://doi.org/10.3390/f16040601

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