Latitudinal clines in life-history traits of the cabbage beetle,  Colaphellus bowringi: showing a stepwise pattern

Latitudinal clines in life-history traits of the cabbage beetle, Colaphellus bowringi: showing a stepwise pattern | Research Square window.SnipcartSettings = { analytics: { enabled: false } }; (function() { var accessVector = localStorage.getItem('access_vector') || ''; window.dataLayer = window.dataLayer || []; if (accessVector) { window.dataLayer.push({ user: { profile: { profileInfo: { snid: accessVector } } } }); } })(); (function(w,d,s,l,i){w[l]=w[l]||[];w[l].push({'gtm.start':new Date().getTime(),event:'gtm.js'});var f=d.getElementsByTagName(s)[0],j=d.createElement(s),dl=l!='dataLayer'?'&l='+l:'';j.async=true;j.src='https://www.googletagmanager.com/gtm.js?id='+i+dl;f.parentNode.insertBefore(j,f);})(window,document,'script','dataLayer','GTM-K279D39R'); Browse Preprints In Review Journals COVID-19 Preprints AJE Video Bytes Research Tools Research Promotion AJE Professional Editing AJE Rubriq About Preprint Platform In Review Editorial Policies Our Team Advisory Board Help Center Sign In Submit a Preprint Cite Share Download PDF Research Article Latitudinal clines in life-history traits of the cabbage beetle, Colaphellus bowringi: showing a stepwise pattern Lili Huang, Fangsen Xue, Jianjun Tang, Haimin He This is a preprint; it has not been peer reviewed by a journal. https://doi.org/ 10.21203/rs.3.rs-6116757/v1 This work is licensed under a CC BY 4.0 License Status: Published Journal Publication published 09 Dec, 2025 Read the published version in Oecologia → Version 1 posted 4 You are reading this latest preprint version Abstract Studying the latitudinal cline in life-history traits is crucial for understanding how organisms adapt to seasonal environments and for predicting their potential responses to climate change. In this study, we systematically examined the life-history traits of the cabbage beetle Colaphellus bowringi collected from six sites spanning a 21º latitudinal range. Our results demonstrated that post-diapause female body weight and fecundity decreased in a stepwise manner with increasing latitude, consistent with the converse Bergmann’s rule. This pattern was also found in pupal and adult weight of their offspring. Larval development time increased while growth rate decreased in a stepwise manner with increasing latitude, indicating cogradient variation. We further found that these stepwise changes are associated with voltinism. Specifically, multivoltine populations exhibited one set of life-history trait pattern, bivoltine populations another, and univoltine populations yet another, collectively forming a stepwise pattern. Additionally, male pupae experienced significantly greater weight loss during metamorphosis compared to female pupae, resulting in lower sexual size dimorphism (SSD) in pupae than in adults. This suggests that sex-specific weight loss during metamorphosis mediates SSD. In summary, our study provides a comprehensive example of insect life-history evolution, particularly in the empirical study of stepped variation patterns. These findings enhance our understanding of latitudinal variation in life-history traits. Latitudinal cline Body weight Development time Growth rate Fecundity Figures Figure 1 Figure 2 Figure 3 Figure 4 Figure 5 Figure 6 Introduction Development time, body size (or body weight), growth rate and fecundity are four critical life-history traits in insects that significantly influence various fitness components of individuals (Nylin and Gotthard 1998 ; Rhainds and Fagan 2010 ). During the non-breeding season, the shorter development time typically reduces the risk of mortality before breeding (Sibly and Calow 1986 ). In contrast, during the breeding season, shortened development times can increase the number of generations per year, thereby enhancing population intrinsic growth rates (Lewontin 1965 ). In most cases, high growth rates are advantageous, as herbivorous insect females often select host plants that support rapid larval development (Thompson 1980) or orient towards sites with optimal sunlight exposure for oviposition (Kadej et al. 2018 ). Female body size is usually positively correlated with fecundity, where larger females tend to produce more eggs (Honěk 1993, Nylin and Gotthard 1998 ). Male body size is positively associated with mating success and survival ability (Blanckenhorn 2000 ). Fecundity serves as a species-specific indicator of reproductive potential and can be used to predict future population dynamics (Figueiredo et al. 2008 ; Stillwell and Fox 2009 ). For widely distributed insect species, clinal variations in life-history traits have been documented in a variety of insect species. The term “cline” was introduced by Huxley ( 1938 ), refers to an observable gradient reflecting continuous changes in the biological characteristics of a species over extensive geographical areas (Koch 1986 ). Clines serve as valuable tools for tracking interactions between climate variables and organisms (Mayekar et al. 2022 ). Two primary patterns describe clinal variation in development time and growth rate: cogradient and countergradient variation (Conover et al. 2009 ). Cogradient variation characterizes geographic patterns where genetic and environmental influences on phenotypic expression act synergistically in the same direction (Falconer 1990 ). For instance, Barton et al. ( 2014 ) observed that low latitude populations of the common brown butterfly, Heteronympha merope exhibited faster growth and development rates compared to higher latitude populations. Countergradient variation describes geographic patterns where genetic and environmental influences on phenotypes oppose each another (Falconer 1990 ). Kivelä et al. ( 2011 ) reported that larval development time decreased with increasing latitudinal gradient in four species of geometrid moths: Cabera exanthemata , Cabera pusaria, Chiasmia clathrata and Lomaspilis marginata . Zeng and Zhu ( 2014 ) found that nymphal development in the Cricket Velarifictorus micado decreased with increasing latitude. Blankenhorn et al. (2018) noted that high-latitude population of the yellow dung fly Scathophaga stercoraria had faster growth rates and short development time than low-latitude populations. Kojima et al. ( 2020 ) observed a strong positive correlation between growth rate and latitude in the univoltine Japanese rhinoceros beetle Trypoxylus dichotomu s. Fu et al. ( 2022 ) reported that high-latitude populations of the Asian corn borer, Ostrinia furnacalis , showed significantly shorter larval developmental times compared to low-latitude populations. According to Conover et al. ( 2009 ), among 15 insect species studied, 13 exhibited countergradient variation, while only two species— Drosophila melanogaster (James et al. 1995 ) and Drosophila subobscura (Gilchrist et al. 2004 ) showed cogradient variation. This suggests that countergradient variation is more prevalent than cogradient variation in insects. However, the underlying mechanisms driving these patterns remain unclear. Both Bergmann’s cline and converse Bergmann’s cline have primarily been utilized to characterize clinal variation in body size (Blanckenhorn and Demont 2004 ; Shelomi 2012 ). According to Bergmann’s rule, body size exhibits a positive correlation with latitude (Atkinson and Sibly 1997 ), whereas the converse Bergmann rule posits a negative correlation between body size and latitude (Mousseau 1997 ). These two clines have been observed in 234 insect species, with 123 studies documenting Bergmann clines and 111 studies reporting converse Bergmann clines (Blanckenhorn and Demont 2004 ). Subsequently, converse Bergmann clines have been identified in various species, including the beetle Paropsis atomaria (Schutze and Clarke 2008 ), the generalist grasshopper, Melanoplus femurrubrum (Parsons and Joern 2014 ), the swallowtail Sericinus montelus (Zheng et al. 2015 ), the univoltine damselfly Lestes spons (Sniegula, et al. 2016 ), the cabbage beetle Colaphellus bowringi (Tang et al. 2017 ) and the dung fly Sepsis fulgens (Roy et al. 2018 ). In the Asian corn borer, O. furnacalis , populations at high-latitudes exhibited significantly greater body weight compared to those at lower latitudes (Fu et al. 2022 ), consistent with Bergmann’s rule. Given that fecundity generally increases with female body size (Honek 1993), it is reasonable to expect that species exhibiting body size clines would also display clinal variations in fecundity. Fecundity has been shown to increase with latitude in D. melanogaster (Schmidt et al. 2005 )d furnacalis (Tu et al. 2012 ), aligning the Bergmann’s rule, while it decreased with increasing latitude in the spruce budworm Choristoneura fumiferana (Harvey 1983 ), in the forest tent caterpillar Malacosoma disstria (Parry et al. 2001 ), in the evergreen bagworm, Thyridopteryx ephemeraeformis (Rhainds and Fagan 2010 ) and in green-veined white butterfly Pieris napi (Günter et al. 2020 ), following the converse Bergmann’s rule. To date, only a limited number of studies have simultaneously examined all four traits when investigating life history variation along a latitudinal gradient. Building upon previous studies (Tang et al. 2017 ), this study extends the investigation into the latitudinal clines in developmental time, body weight, growth rate and fecundity of the cabbage beetle C. bowringi . The specimens were collected from six sites spanning a 21º latitudinal range, with an expanded scope of study content and increased sample size. Examining the latitudinal clines in life-history traits is crucial for understanding how organisms adapt to seasonal environments and for predicting their potential responses to climate change. Materials and methods Life history of C. bowringi The cabbage beetle C. bowringi , is a leaf-feeding pest primarily affecting cruciferous plants and is widely distributed across China. This study examines six populations that exhibit diverse life histories. Populations from lower latitudes (LN and XS) demonstrate multivoltine annual life history (Xue et al. 2002 a; Lai et al. 2008 ). In the field, these populations display two distinct infestation peaks, one in spring with a single generation and another in autumn with three generations. Both populations undergo an aestivating and hibernating imaginal diapause in the soil. Temperature-dependent short-day response are observed, where shorter day lengths coupled with higher temperature lead to non-diapause development, while longer daylengths coupled with higher temperature induce diapause (Xue et al. 2002 b; Lai et al. 2008 ). Regardless of photoperiod, diapause is triggered when mean daily temperature drop to ≤ 18°C for LN population or ≤ 20°C for XS population. Populations from mid-latitudes (XY and TA) exhibit bivoltine annual life history (Lai et al. 2008 ; Tang et al. 2017 ). These populations also show two distinct infestation peaks, one in spring and another in autumn, each with a single generation. They undergo imaginal summer and winter diapause in the soil. Unlike lower latitude populations, these populations lack photoperiodic response, and all individuals are induced into diapause at temperatures below 25°C (Tang et al. 2017 ; Lai et al. 2008 ). Higher latitude populations (SY and HB) exhibit univoltine annual life history. Only one generation is produced in summer, and adults overwinter in the soil as diapausing adults. All almost individuals enter diapause at ≤ 28°C regardless of photoperiod (Lai et al. 2008 ; Chen et al. 2014 ; He et al. 2021 ). Lai et al. ( 2008 ) reported that under controlled conditions of 25°C and L12:D12 photoperiod, the incidence of diapause increased with increasing latitude, indicating a latitudinal cline in diapause incidence. Collection sites and insect culture In spring of 2022, naturally diapausing adults of C. bowringi were collected from six locations: Longnan County (LN, 24°9' N, 114°8' E, as LN population), Xiushui County (XS, 29°1' N, 114°4' E, as XS population), Xinyang County (XY, 31°48' N, 114°03' E, as XY population), Taian City (TA, 36°2' N, 117°1' E, as TA population), Shenyang City (SY, 41°48' N, 123°23' E) and Harbin City (HB, 45°8' N, 126°6' E) (see Fig. 1 ). Diapause adults from each population were transferred into glass bottles (diameter 18 cm; height 32 cm) containing soil, allowing them to burrow for dormancy The bottles were maintained under semi-natural conditions (a covered, open-air room with natural temperature and photoperiod fluctuations) at Jiangxi Agricultural University in Nanchang, Jiangxi Province (28°46' N, 115°59' E). In spring 2023, post-diapausing adults emerged from the soil, and males and females from each population were weighed using an electric balance (AUY120, SHIMADZU Corporation, Japan). Females and males were paired randomly in petri dishes (9.0 cm diameter, 2.0 cm height) lined with filter paper and fresh leaves of the potherb mustard ( Brassica juncea (L.) Czern. et Coss. var. multiceps Tsen et Lee) for mating and oviposition. The lifetime number of eggs laid by each pair was recorded (see Table S1 ). Eggs laid with the first 3 days were collected for subsequent experiments and checked daily until hatching. Experimental design Up hatching, larvae for each population were transferred to rearing boxes (16 × 11 × 5.5 cm) containing fresh potherb mustard leaves. Each box housed 80 larvae, and the boxes were cleaned daily with fresh leaves provided as needed. Each population had three to four groups, with each group consisting of 1–2 boxes. After reaching maturity, larvae were individually placed in cell culture plates with 12 holes (hole: diameter: 2.4 cm, height: 2 cm) for pupation and emergence. Pupation time and adult emergence times were observed every morning. For each individual, we recorded the development time from hatching to pupation and to adult emergence and weighed the pupa and adult. Growth rate, proportional weight loss during metamorphosis, and sexual size dimorphism (SSD) were calculated. Pupae were weighed the next day after pupation, and adults were weighed on the day of emergence after their elytra hardened. Growth rate was computed as ln pupal weight / larval development times (Gotthard et al. 1994 ). Weight loss ratio was calculated as l-(adult weight/pupal weight). The SSD was estimated using a sexual dimorphism index (Lovich and Gibbons 1992 ), defined as SSD = (size of the larger sex/size of the smaller sex)-1. Overall, 1811 offspring (903 females and 908 males) from six populations were raised to adulthood and weighed. All experiments were conducted in illuminated incubators (LRH-250-GS, Guangdong Medical Appliances Plant, Guangdong,China). The temperature was maintained at 22°C with a photoperiod of L15:D9 h. Under these conditions, all individuals were induced into adult diapause. Statistical analyses Statistical analyses were conducted using IBM SPSS Statistics 22.0. A linear mixed model (LMM) was utilized to assess the effects of population, sex, and their interaction on life-history traits. In this model, population and sex were treated as fixed factors, while rearing group was considered a random factor. The model was specified as: Trait ~ population * sex + (1|rearing group). Residual diagnostics confirmed that the assumptions of normality and homoscedasticity were met. To further investigate population-level differences identified by the LMM, one-way ANOVA followed by Tukey’s HSD post hoc tests were employed. Independent samples t-tests, with Welch’s correction for unequal variances, were used to compare traits between sexes. Linear regression analyses were conducted to examine the relationships between latitude and both fecundity and sexual size dimorphism (SSD). Results Body weight and fecundity in post-diapause mated females S ignificant differences were observed in body weight (F 5, 116 = 109.116, P < 0.001) and fecundity (F 5, 116 = 8.557, P < 0.001) among population. Both body weight and fecundity showed a stepwise decline with increasing latitude (Fig. 2 A), forming a distinct latitudinal gradient. Low-latitude populations maintained consistent body weight and fecundity levels, while mid- and high-latitude latitude populations showed decreasing trends. This pattern aligns with the converse Bergmann’s rule. A significant positive correlation was found between female body weight and fecundity (Fig. 2 B, see also Table S2 ). Progeny development time Larval development time was significantly influenced by population (Table 1 ), but not by sex or the interaction population between population and sex. Larval development time increased stepwise with increasing latitude (Fig. 3 A), exemplifying cogradient variation. In contrast, pupal development time was unaffected by population, sex, or their interactions (Table 1 , Fig. 3 B). Table 1 Results from a linear mixed-model analysis of fixed effects on larval time, pupal time, pupal weight, growth rate, adult weight and proportionate weight loss in Colaphellus bowringi in relation to population and sex Life-history traits Fixed factors df F -value P -value Larval time Population 5 776.439 < 0.001 Sex 1 0.305 0.581 Population × sex 5 0.416 0.838 Pupal time Population 5 1.848 0.100 Sex 1 0.031 0.861 Population × sex 5 0.175 0.972 Pupal weight Population 5 905.185 < 0.001 Sex 1 2969.607 < 0.001 Population × sex 5 24.300 < 0.001 Growth rate Population 5 1521.485 < 0.001 Sex 1 1129.139 < 0.001 Population × sex 5 3.552 0.003 Adult weight Population 5 760.767 < 0.001 Sex 1 3724.932 < 0.001 Population × sex 5 30.422 < 0.001 Proportionate weight loss Population 5 43.497 < 0.001 Sex 1 459.878 < 0.001 Population × sex 5 10.941 < 0.001 Progeny pupal weight and growth rate Pupal weight and growth rate were significantly affected by population, sex and their interactions (Table 1 ). Both Pupal weight and growth rate decreased stepwise with increasing latitude, adhering to the converse Bergmann’s rule or cogradient variation (Fig. 4 A and 4 B). Across all populations, female pupae exhibited significantly higher body weight and growth rate compared to male pupae (see Table S3 , P < 0.05). Progeny adult weight and weight loss Adult weight and weight loss were significantly influenced by population, sex and their interactions (Table 1 ). Both female and male adults showed a stepwise decline in weight with increasing latitude (Fig. 5 A), confirming a converse Bergmann’s cline. Male adults experienced significantly greater weight loss than females (Fig. 5 B, see Table S3 ). For males, the highest weight loss occurred at low latitudes, followed by high and middle latitudes. For females, the weight loss gradually decreased with increasing latitude, indicating a latitudinal cline. The higher weight loss in low-latitudinal populations suggests that larger individuals lose more weight during metamorphosis compared to smaller ones. Sexual size dimorphism (SSD) SSD for both pupae and adults varied substantially with latitude (Fig. 6 A). The SSD index for pupae was lower than that for adults, particularly in low-latitude populations. Higher latitudes exhibited lower SSD indices compared to middle and low latitudes. Linear regression analysis revealed a negative correlation between SSD and Latitude (Fig. 6 B). Discussion Our study revealed for the first time that adult body weigh and fecundity in C. bowringi decreased with increasing latitude after diapause, and female body weight and fecundity were significantly positively correlated (Fig. 2 ). This latitudinal cline in body weight and fecundity is converse to Bergmann’s cline. Notably, the latitudinal clines in body weight and fecundity exhibited a stepwise decline pattern. Specifically, populations with multivoltine annual life history at low latitudes (LN and XS) showed similar body weight and fecundity. Similarly, mid-latitude populations (XY and TA) with bivoltine annual life history exhibited comparable body weight and fecundity, as did high-latitude populations (SY and HB) with univoltine annual life history. This stepwise pattern has only been reported in two other species (Shelomi 2012 ), suggesting that voltinism plays a crucial role in determining life-history patterns. Furthermore, the weight of the pupae and adults produced by post-diapause individuals also decreased in a stepwise manner with increasing latitude. (Fig. 5 A and Fig. 6 A). Additionally, we observed a latitudinal cline in larval development time and growth rate. Larval development time increased significantly in a stepwise manner with increasing latitude (Fig. 3 A), while growth rate decreased significantly in a stepwise manner (Fig. 4 B). Both variables exhibited cogradient variation, consistent with findings in H. merope , where low-latitude populations had faster growth and shorter development time compared to high-latitude populations (Barton et al. 2014 ). Unlike the C. bowringi , H. merope did not show latitudinal cline in body size, indicating that selection pressures on development time and body weight may follow different evolutionary pathways. We also found significant variation across populations in the relationship between larval development and body weight. Low-latitude populations (LN and XS) had the shortest development time and largest body weight while mid-latitude populations (XY and TA) exhibited intermediate values, and high-latitude populations (SY and HB) had the longest development time and smallest body weight. These findings align with previous studies (Tang et al. 2017 ; He et al. 2021 ). Generally, larval development time is positively correlated with body size, that is, animals with longer development time are larger at maturity (Nijhout et al. 2010 ) or take longer time to grow larger (Roff 2000 ; Stearns 1992 ), but this assumption does not hold when comparing populations from different regions. For instance, in O. furnacalis for the diapausing generation, the high-latitude populations exhibited significantly longer larval development time and lower body weight compared to low-latitude populations (Fu et al. 2022 ). The converse Bergmann’s cline (decreasing body size with increasing latitude) is typically attributed to season length effect (Blanckenhorn and Demont 2004 ). However, in our study, the converse Bergmann’s cline in C. bowringi is unlikely to be associated with gradual changes in seasonality. Reproductive activity in high-latitude SY and HB populations occurs during summer, providing sufficient time for larvae to develop without seasonal constraints. The significantly longer development times and small body weights in northern populations may instead be related to local climatic conditions rather than season length. The stepwise clines in development time and body weight in C. bowringi suggest genetic regional differences in life-history traits. Based on Horne et al. ( 2015 ), voltinism in arthropods is strongly correlated with body weight, with univoltine species often being larger than multivoltine species. However, this rule does not apply to C. bowringi . In this study, the multivoltine populations (LN and XS) had the largest body weight, followed by bivoltine populations (XY and TA), with univoltine populations (e SY and HB) having the smallest body weight. Our results suggest that relationship between voltinism and body size can vary depending on the species. Our study demonstrated that, across all populations, pupae lost significantly more weight during metamorphosis compared to females (Fig. 5 B). This suggests that males either have a higher metabolic rate or are less effective at preventing water loss (Savalli and Fox 1998 ). Consequently, the SSD in pupae is significantly lower than that in adults, indicating that sex-specific weight loss mediated SSD in C. bowringi (Testa et al. 2013 ). Additionally, we observes that female weight loss exhibited a gradually downward latitudinal gradient, while male weight loss followed a U-shaped along the latitudinal gradient, with the mid-latitude populations experiencing the least weight loss (Fig. 5 B). According to Rensch’s rule, SSD tends to increase with body size when males are the larger sex, and decrease with body size when females are the larger sex (Abouheif and Fairbairn 1997 , Fairbairn 1997 ). In our study, for female-biased cabbage beetles reared at 22°C, SSD tended to increase with increasing body weight, and SSD index tended to decrease with increasing latitude, which appears in inconsistent with Rensch’s rule (Teder and Tammaru 2005 ). Similar finding was reported in the male-biased beetle Stator limbatus , where both males and females increased with increasing latitude, but dimorphism decreased with increasing latitude, also not supporting Rensch’s rule (Stillwell et al., 2007 ). Conclusion Our study revealed latitudinal clines in body size, fecundity, development time, growth rate, weight loss and SSD in C. bowringi . Body size and fecundity exhibited converse Bergmann’s cline with a stepwise decline pattern, while development time and growth rate exhibited cogradient variation, with development time increasing stepwise and growth rate decreasing stepwise with latitude. These patterns appear to be related to voltinism, as differences in voltinism across geographical populations collectively construct the stepwise pattern. We also found that female weight loss tended to decrease with increasing latitude, while male weight loss was the lowest in mid-latitude. The SSD index for both pupae and adults tended to decrease with latitude. These joint results indicate that this species comprises regionally adapted populations. These findings broaden our understanding of latitudinal variation in life-history traits in insects. Declarations Supplementary Information The online version contains supplementary material Acknowledgement We thank the National Natural Science Foundation of China for its financial support of this project. Funding The work has been funded by the National Natural Science Foundation of China (32360270) and High-level Scientific Research and Innovation Team of Yuzhang Normal University (YZTD202305). Author contributions HH and XF conceived the ideas and designed the study. HH, HL and XF performed this experiment. XF and HL wrote the original manuscript. HH and TJ analyzed the data. All authors have read and agreed to the published version of the manuscript. Availability Statement of data and materials Data are contained within the article. Code availability Not available Conflicts of Interest: The authors declare no conflict of interest. Ethical approval Not applicable. Consent to participate Not applicable. Consent for publication Not applicable. References Abouheif E, Fairbairn DJ (1997) A comparative analysis of allometry for sexual size dimorphism: assessing Rensch's rule. Am Nat 149:540–562 Atkinson D, Sibly RM (1997) Why are organisms usually bigger in colder environments? Making sense of a life history puzzle. 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Am Nat 170:358–369 Stillwell RC, Fox CW (2009) Geographic variation in body size, sexual size dimorphism and fitness components of a seed beetle: local adaptation versus phenotypic plasticity. Oikos 118:703–712 Tang JJ, He HM, Chen C, Fu S, Xue FS (2017) Latitudinal cogradient variation of development time and growth rate and a negative latitudinal body weight cline in a widely distributed cabbage beetle. PLoS ONE 12(7):0181030 Teder T, Tammaru T (2005) Sexual size dimorphism within species increases with body size in insects. Oikos 108:321–334 Testa ND, Ghosh SM, Shingleton AW (2013) Sex-specific weight loss mediates sexual size dimorphism in Drosophila melanogaster . PLoS ONE 8:e58936 Thompson JN (1988) Evolutionary ecology of the relationship between oviposition preference and performance of offspring in phytophagous insects. Entomol Exp Appl 47:3–14 Tu XY, Chen YS, Xia QW, Chen C, Kuang XJ, Xue FS (2012) Geographic variation in longevity and fecundity of the Asian corn borer, Ostrinia furnacalis Guenée (Lepidoptera: Crambidae). Acta Ecol Sin 32:4160–4165 Xue FS, Li A, Zhu XF, Gui AL, Jiang PL, Liu XF (2002) Diversity in life history of the leaf beetle, Colaphellus bowringi Baly. Acta Entomologica Sinica 45:494–498 Xue FS, Spieth HR, Li AQ, Hua A (2002) The role of photoperiod and temperature in determination of summer and winter diapause in the cabbage beetle, Colaphellus bowringi (Coleoptera: Chrysomelidae). J Insect Physiol 48:279–286 Zeng Y, Zhu DH (2014) Geographical Variation in Body Size, Development Time, and Wing Dimorphism in the Cricket Velarifictorus micado (Orthoptera: Gryllidae). Ann Entomol Soc Am 6:1066–1071 Zheng XL, Yang QS, Hu YW, Lei CL, Wang XP (2015) Latitudinal variation of morphological characteristics in the swallowtail Sericinus montelus Gray, 1798 (Lepidoptera: Papilionidae). Acta Zool–Stockholm 96:242–252 Supplementary Files TableS1Biologyofmatedfemales.doc TableS2Bodyweightandfecundity.doc TableS3Lifehistorydata.doc Cite Share Download PDF Status: Published Journal Publication published 09 Dec, 2025 Read the published version in Oecologia → Version 1 posted Reviewers agreed at journal 30 May, 2025 Reviewers invited by journal 21 Mar, 2025 Editor assigned by journal 28 Feb, 2025 First submitted to journal 26 Feb, 2025 You are reading this latest preprint version Research Square lets you share your work early, gain feedback from the community, and start making changes to your manuscript prior to peer review in a journal. As a division of Research Square Company, we’re committed to making research communication faster, fairer, and more useful. We do this by developing innovative software and high quality services for the global research community. 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Also discoverable on Platform About Our Team In Review Editorial Policies Advisory Board Help Center Resources Author Services Accessibility API Access RSS feed Manage Cookie Preferences © Research Square 2026 | ISSN 2693-5015 (online) Privacy Policy Terms of Service Do Not Sell My Personal Information {"props":{"pageProps":{"initialData":{"identity":"rs-6116757","acceptedTermsAndConditions":true,"allowDirectSubmit":false,"archivedVersions":[],"articleType":"Research Article","associatedPublications":[],"authors":[{"id":431994778,"identity":"41dad7ad-a98d-4530-b81e-00f1b3fe4244","order_by":0,"name":"Lili Huang","email":"data:image/png;base64,iVBORw0KGgoAAAANSUhEUgAAAZAAAAAyAQMAAABI0h/eAAAABlBMVEX///8AAABVwtN+AAAACXBIWXMAAA7EAAAOxAGVKw4bAAAAyUlEQVRIiWNgGAWjYBACNvbGhgMfDGrk+NkbiNTCx3O48eCMimPGkj0HiNQiJ5HefJjnDHPihhsJxDqMIbHhAG8bG+PMmY833mCosYkmQsvBhgOSbTLM/NJpxRYMx9JyGwhqYQR637CNjU1ydo6ZBGPDYSK0MDM2HEhsY+YxuHmGWC1sQC0HzjBLGNzgIVYLD2PDwYaKYwaSPUC/JBDjF/n5zx9//mNQU9/PfnjjjQ81NoS1IAMDiQRSlEO0kKpjFIyCUTAKRgYAAHdvQn1JrctlAAAAAElFTkSuQmCC","orcid":"https://orcid.org/0000-0002-1117-7122","institution":"Yuzhang Normal University","correspondingAuthor":true,"prefix":"","firstName":"Lili","middleName":"","lastName":"Huang","suffix":""},{"id":431994779,"identity":"aff10f3c-8338-4325-ab63-f59015d72a8d","order_by":1,"name":"Fangsen Xue","email":"","orcid":"","institution":"Jiangxi Agricultural University","correspondingAuthor":false,"prefix":"","firstName":"Fangsen","middleName":"","lastName":"Xue","suffix":""},{"id":431994780,"identity":"6bc4fab0-7587-4012-9f07-da349d463508","order_by":2,"name":"Jianjun Tang","email":"","orcid":"","institution":"Jiangxi Agricultural University","correspondingAuthor":false,"prefix":"","firstName":"Jianjun","middleName":"","lastName":"Tang","suffix":""},{"id":431994781,"identity":"dea21d3a-e3da-41e0-ae49-8de88952cdc4","order_by":3,"name":"Haimin He","email":"","orcid":"https://orcid.org/0000-0002-6849-0742","institution":"Jiangxi Agricultural University","correspondingAuthor":false,"prefix":"","firstName":"Haimin","middleName":"","lastName":"He","suffix":""}],"badges":[],"createdAt":"2025-02-27 02:18:20","currentVersionCode":1,"declarations":"","doi":"10.21203/rs.3.rs-6116757/v1","doiUrl":"https://doi.org/10.21203/rs.3.rs-6116757/v1","draftVersion":[],"editorialEvents":[{"content":"https://doi.org/10.1007/s00442-025-05849-3","type":"published","date":"2025-12-09T15:57:27+00:00"}],"editorialNote":"","failedWorkflow":false,"files":[{"id":79587855,"identity":"a34fc976-39be-4003-a275-d64eea4b49c1","added_by":"auto","created_at":"2025-03-31 12:44:27","extension":"png","order_by":1,"title":"Figure 1","display":"","copyAsset":false,"role":"figure","size":102295,"visible":true,"origin":"","legend":"\u003cp\u003eThe collection sites of samples of \u003cem\u003eColaphellus bowringi\u003c/em\u003e. Each dot denotes a collection site.\u003c/p\u003e","description":"","filename":"OnlineFig.1.png","url":"https://assets-eu.researchsquare.com/files/rs-6116757/v1/4743fce2df1b4cea35e18b99.png"},{"id":79588762,"identity":"c6e42dcc-0f53-4520-89eb-2da11a5b50d9","added_by":"auto","created_at":"2025-03-31 12:52:27","extension":"png","order_by":2,"title":"Figure 2","display":"","copyAsset":false,"role":"figure","size":31624,"visible":true,"origin":"","legend":"\u003cp\u003eBody weight and fecundity for post-diapasue females of \u003cem\u003eColaphellus bowringi\u003c/em\u003e in different geographical populations at 22 °C and 15L: 9D photoperiod. Error bars indicate SE. Values with different lowercase among different populations are significantly different at the 0.05 level. The two asterisks indicate a significant negative correlation between fecundity and latitude.\u003c/p\u003e","description":"","filename":"OnlineFig.2.png","url":"https://assets-eu.researchsquare.com/files/rs-6116757/v1/5b79adf50cc27d9631ddb6de.png"},{"id":79587858,"identity":"86a75a58-6201-497c-8013-3584713c1a7d","added_by":"auto","created_at":"2025-03-31 12:44:27","extension":"png","order_by":3,"title":"Figure 3","display":"","copyAsset":false,"role":"figure","size":26480,"visible":true,"origin":"","legend":"\u003cp\u003eLarval and pupal development times of \u003cem\u003eColaphellus bowringi \u003c/em\u003efemales and males in different geographical populations at 22 °C and 15L: 9D photoperiod. Error bars indicate SE. Values with different lowercase among different populations are significantly different at the 0.05 level.\u003c/p\u003e","description":"","filename":"OnlineFig.3.png","url":"https://assets-eu.researchsquare.com/files/rs-6116757/v1/eeb6b8ce785950fab66c71b9.png"},{"id":79590008,"identity":"90b05693-3916-407d-9a75-a2ab3f5c477b","added_by":"auto","created_at":"2025-03-31 13:00:27","extension":"png","order_by":4,"title":"Figure 4","display":"","copyAsset":false,"role":"figure","size":26665,"visible":true,"origin":"","legend":"\u003cp\u003ePupal weight and larval growth rate of \u003cem\u003eColaphellus bowringi\u003c/em\u003e females and males in different geographical populations at 22 °C and 15L: 9D photoperiod. Error bars indicate SE. Values with different lowercase among different populations are significantly different at the 0.05 level. The asterisk means significant difference between sexes (\u003cem\u003eT \u003c/em\u003etest, \u003cem\u003eP \u003c/em\u003e\u0026lt; 0.05).\u003c/p\u003e","description":"","filename":"OnlineFig.4.png","url":"https://assets-eu.researchsquare.com/files/rs-6116757/v1/a99c8c9fb8fa64ae594c94d6.png"},{"id":79587860,"identity":"d85cbfa3-b805-41f4-bd5d-21ae5368e3ac","added_by":"auto","created_at":"2025-03-31 12:44:27","extension":"png","order_by":5,"title":"Figure 5","display":"","copyAsset":false,"role":"figure","size":25478,"visible":true,"origin":"","legend":"\u003cp\u003eAdult weight and proportionate weight loss of \u003cem\u003eColaphellus bowringi\u003c/em\u003e females and males in different geographical populations at 22 °C and 15L: 9D photoperiod. Error bars indicate SE. Values with different lowercase among different populations are significantly different at the 0.05 level. The asterisk means significant difference between sexes (\u003cem\u003eT \u003c/em\u003etest, \u003cem\u003eP \u003c/em\u003e\u0026lt; 0.05).\u003c/p\u003e","description":"","filename":"OnlineFig.5.png","url":"https://assets-eu.researchsquare.com/files/rs-6116757/v1/e83415ec6d83fc578d2183d4.png"},{"id":79588763,"identity":"688d2ed3-b030-41e4-ad93-8094d3d526ab","added_by":"auto","created_at":"2025-03-31 12:52:27","extension":"png","order_by":6,"title":"Figure 6","display":"","copyAsset":false,"role":"figure","size":29983,"visible":true,"origin":"","legend":"\u003cp\u003eSexual size dimorphism of \u003cem\u003eColaphellus bowringi\u003c/em\u003e pupa and adult in different geographical populations at 22 °C and 15L: 9D photoperiod. Error bars indicate SE. Values with different lowercase among different populations are significantly different at the 0.05 level. The asterisk means significant difference between sexes (\u003cem\u003eT \u003c/em\u003etest, \u003cem\u003eP \u003c/em\u003e\u0026lt; 0.05).\u003c/p\u003e","description":"","filename":"OnlineFig.6.png","url":"https://assets-eu.researchsquare.com/files/rs-6116757/v1/c0df6cba89d7eb7bf556329f.png"},{"id":98243989,"identity":"cc817412-a538-4652-8e28-bda722b39849","added_by":"auto","created_at":"2025-12-15 16:12:08","extension":"pdf","order_by":0,"title":"","display":"","copyAsset":false,"role":"manuscript-pdf","size":1161529,"visible":true,"origin":"","legend":"","description":"","filename":"manuscript.pdf","url":"https://assets-eu.researchsquare.com/files/rs-6116757/v1/4cfd3813-b157-4b3f-ac6d-f73f0a854a72.pdf"},{"id":79587872,"identity":"506c666d-8346-4015-a26e-46e2a02ccb23","added_by":"auto","created_at":"2025-03-31 12:44:27","extension":"doc","order_by":11,"title":"","display":"","copyAsset":false,"role":"supplement","size":38400,"visible":true,"origin":"","legend":"","description":"","filename":"TableS1Biologyofmatedfemales.doc","url":"https://assets-eu.researchsquare.com/files/rs-6116757/v1/eec2467ece1d89ecae7002e1.doc"},{"id":79588765,"identity":"4a61a9e5-08ca-4649-a661-256283a4e2ee","added_by":"auto","created_at":"2025-03-31 12:52:27","extension":"doc","order_by":12,"title":"","display":"","copyAsset":false,"role":"supplement","size":34304,"visible":true,"origin":"","legend":"","description":"","filename":"TableS2Bodyweightandfecundity.doc","url":"https://assets-eu.researchsquare.com/files/rs-6116757/v1/9154eafae02a1bb324e01a97.doc"},{"id":79587868,"identity":"a8543088-8b09-4bad-8043-92a7a635336e","added_by":"auto","created_at":"2025-03-31 12:44:27","extension":"doc","order_by":13,"title":"","display":"","copyAsset":false,"role":"supplement","size":50688,"visible":true,"origin":"","legend":"","description":"","filename":"TableS3Lifehistorydata.doc","url":"https://assets-eu.researchsquare.com/files/rs-6116757/v1/fdc1d3eafcde04d3d8e650ab.doc"}],"financialInterests":"","formattedTitle":"Latitudinal clines in life-history traits of the cabbage beetle, Colaphellus bowringi: showing a stepwise pattern","fulltext":[{"header":"Introduction","content":"\u003cp\u003eDevelopment time, body size (or body weight), growth rate and fecundity are four critical life-history traits in insects that significantly influence various fitness components of individuals (Nylin and Gotthard \u003cspan citationid=\"CR32\" class=\"CitationRef\"\u003e1998\u003c/span\u003e; Rhainds and Fagan \u003cspan citationid=\"CR36\" class=\"CitationRef\"\u003e2010\u003c/span\u003e). During the non-breeding season, the shorter development time typically reduces the risk of mortality before breeding (Sibly and Calow \u003cspan citationid=\"CR43\" class=\"CitationRef\"\u003e1986\u003c/span\u003e). In contrast, during the breeding season, shortened development times can increase the number of generations per year, thereby enhancing population intrinsic growth rates (Lewontin \u003cspan citationid=\"CR28\" class=\"CitationRef\"\u003e1965\u003c/span\u003e). In most cases, high growth rates are advantageous, as herbivorous insect females often select host plants that support rapid larval development (Thompson 1980) or orient towards sites with optimal sunlight exposure for oviposition (Kadej et al. \u003cspan citationid=\"CR23\" class=\"CitationRef\"\u003e2018\u003c/span\u003e). Female body size is usually positively correlated with fecundity, where larger females tend to produce more eggs (Honěk 1993, Nylin and Gotthard \u003cspan citationid=\"CR32\" class=\"CitationRef\"\u003e1998\u003c/span\u003e). Male body size is positively associated with mating success and survival ability (Blanckenhorn \u003cspan citationid=\"CR4\" class=\"CitationRef\"\u003e2000\u003c/span\u003e). Fecundity serves as a species-specific indicator of reproductive potential and can be used to predict future population dynamics (Figueiredo et al. \u003cspan citationid=\"CR11\" class=\"CitationRef\"\u003e2008\u003c/span\u003e; Stillwell and Fox \u003cspan citationid=\"CR47\" class=\"CitationRef\"\u003e2009\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eFor widely distributed insect species, clinal variations in life-history traits have been documented in a variety of insect species. The term \u0026ldquo;cline\u0026rdquo; was introduced by Huxley (\u003cspan citationid=\"CR21\" class=\"CitationRef\"\u003e1938\u003c/span\u003e), refers to an observable gradient reflecting continuous changes in the biological characteristics of a species over extensive geographical areas (Koch \u003cspan citationid=\"CR25\" class=\"CitationRef\"\u003e1986\u003c/span\u003e). Clines serve as valuable tools for tracking interactions between climate variables and organisms (Mayekar et al. \u003cspan citationid=\"CR30\" class=\"CitationRef\"\u003e2022\u003c/span\u003e). Two primary patterns describe clinal variation in development time and growth rate: cogradient and countergradient variation (Conover et al. \u003cspan citationid=\"CR8\" class=\"CitationRef\"\u003e2009\u003c/span\u003e). Cogradient variation characterizes geographic patterns where genetic and environmental influences on phenotypic expression act synergistically in the same direction (Falconer \u003cspan citationid=\"CR10\" class=\"CitationRef\"\u003e1990\u003c/span\u003e). For instance, Barton et al. (\u003cspan citationid=\"CR3\" class=\"CitationRef\"\u003e2014\u003c/span\u003e) observed that low latitude populations of the common brown butterfly, \u003cem\u003eHeteronympha merope\u003c/em\u003e exhibited faster growth and development rates compared to higher latitude populations. Countergradient variation describes geographic patterns where genetic and environmental influences on phenotypes oppose each another (Falconer \u003cspan citationid=\"CR10\" class=\"CitationRef\"\u003e1990\u003c/span\u003e). Kivel\u0026auml; et al. (\u003cspan citationid=\"CR24\" class=\"CitationRef\"\u003e2011\u003c/span\u003e) reported that larval development time decreased with increasing latitudinal gradient in four species of geometrid moths: \u003cem\u003eCabera exanthemata\u003c/em\u003e, \u003cem\u003eCabera pusaria, Chiasmia clathrata\u003c/em\u003e and \u003cem\u003eLomaspilis marginata\u003c/em\u003e. Zeng and Zhu (\u003cspan citationid=\"CR55\" class=\"CitationRef\"\u003e2014\u003c/span\u003e) found that nymphal development in the Cricket \u003cem\u003eVelarifictorus micado\u003c/em\u003e decreased with increasing latitude. Blankenhorn et al. (2018) noted that high-latitude population of the yellow dung fly \u003cem\u003eScathophaga stercoraria\u003c/em\u003e had faster growth rates and short development time than low-latitude populations. Kojima et al. (\u003cspan citationid=\"CR26\" class=\"CitationRef\"\u003e2020\u003c/span\u003e) observed a strong positive correlation between growth rate and latitude in the univoltine Japanese rhinoceros beetle \u003cem\u003eTrypoxylus dichotomu\u003c/em\u003es. Fu et al. (\u003cspan citationid=\"CR12\" class=\"CitationRef\"\u003e2022\u003c/span\u003e) reported that high-latitude populations of the Asian corn borer, \u003cem\u003eOstrinia furnacalis\u003c/em\u003e, showed significantly shorter larval developmental times compared to low-latitude populations. According to Conover et al. (\u003cspan citationid=\"CR8\" class=\"CitationRef\"\u003e2009\u003c/span\u003e), among 15 insect species studied, 13 exhibited countergradient variation, while only two species\u0026mdash;\u003cem\u003eDrosophila melanogaster\u003c/em\u003e (James et al. \u003cspan citationid=\"CR22\" class=\"CitationRef\"\u003e1995\u003c/span\u003e) and \u003cem\u003eDrosophila subobscura\u003c/em\u003e (Gilchrist et al. \u003cspan citationid=\"CR13\" class=\"CitationRef\"\u003e2004\u003c/span\u003e) showed cogradient variation. This suggests that countergradient variation is more prevalent than cogradient variation in insects. However, the underlying mechanisms driving these patterns remain unclear.\u003c/p\u003e \u003cp\u003eBoth Bergmann\u0026rsquo;s cline and converse Bergmann\u0026rsquo;s cline have primarily been utilized to characterize clinal variation in body size (Blanckenhorn and Demont \u003cspan citationid=\"CR5\" class=\"CitationRef\"\u003e2004\u003c/span\u003e; Shelomi \u003cspan citationid=\"CR42\" class=\"CitationRef\"\u003e2012\u003c/span\u003e). According to Bergmann\u0026rsquo;s rule, body size exhibits a positive correlation with latitude (Atkinson and Sibly \u003cspan citationid=\"CR2\" class=\"CitationRef\"\u003e1997\u003c/span\u003e), whereas the converse Bergmann rule posits a negative correlation between body size and latitude (Mousseau \u003cspan citationid=\"CR31\" class=\"CitationRef\"\u003e1997\u003c/span\u003e). These two clines have been observed in 234 insect species, with 123 studies documenting Bergmann clines and 111 studies reporting converse Bergmann clines (Blanckenhorn and Demont \u003cspan citationid=\"CR5\" class=\"CitationRef\"\u003e2004\u003c/span\u003e). Subsequently, converse Bergmann clines have been identified in various species, including the beetle \u003cem\u003eParopsis atomaria\u003c/em\u003e (Schutze and Clarke \u003cspan citationid=\"CR40\" class=\"CitationRef\"\u003e2008\u003c/span\u003e), the generalist grasshopper, \u003cem\u003eMelanoplus femurrubrum\u003c/em\u003e (Parsons and Joern \u003cspan citationid=\"CR35\" class=\"CitationRef\"\u003e2014\u003c/span\u003e), the swallowtail \u003cem\u003eSericinus montelus\u003c/em\u003e (Zheng et al. \u003cspan citationid=\"CR56\" class=\"CitationRef\"\u003e2015\u003c/span\u003e), the univoltine damselfly \u003cem\u003eLestes spons\u003c/em\u003e (Sniegula, et al. \u003cspan citationid=\"CR44\" class=\"CitationRef\"\u003e2016\u003c/span\u003e), the cabbage beetle \u003cem\u003eColaphellus bowringi\u003c/em\u003e (Tang et al. \u003cspan citationid=\"CR48\" class=\"CitationRef\"\u003e2017\u003c/span\u003e) and the dung fly \u003cem\u003eSepsis fulgens\u003c/em\u003e (Roy et al. \u003cspan citationid=\"CR38\" class=\"CitationRef\"\u003e2018\u003c/span\u003e). In the Asian corn borer, \u003cem\u003eO. furnacalis\u003c/em\u003e, populations at high-latitudes exhibited significantly greater body weight compared to those at lower latitudes (Fu et al. \u003cspan citationid=\"CR12\" class=\"CitationRef\"\u003e2022\u003c/span\u003e), consistent with Bergmann\u0026rsquo;s rule. Given that fecundity generally increases with female body size (Honek 1993), it is reasonable to expect that species exhibiting body size clines would also display clinal variations in fecundity. Fecundity has been shown to increase with latitude in \u003cem\u003eD. melanogaster\u003c/em\u003e (Schmidt et al. \u003cspan citationid=\"CR41\" class=\"CitationRef\"\u003e2005\u003c/span\u003e)d \u003cem\u003efurnacalis\u003c/em\u003e (Tu et al. \u003cspan citationid=\"CR52\" class=\"CitationRef\"\u003e2012\u003c/span\u003e), aligning the Bergmann\u0026rsquo;s rule, while it decreased with increasing latitude in the spruce budworm \u003cem\u003eChoristoneura fumiferana\u003c/em\u003e (Harvey \u003cspan citationid=\"CR17\" class=\"CitationRef\"\u003e1983\u003c/span\u003e), in the forest tent caterpillar \u003cem\u003eMalacosoma disstria\u003c/em\u003e (Parry et al. \u003cspan citationid=\"CR34\" class=\"CitationRef\"\u003e2001\u003c/span\u003e), in the evergreen bagworm, \u003cem\u003eThyridopteryx ephemeraeformis\u003c/em\u003e (Rhainds and Fagan \u003cspan citationid=\"CR36\" class=\"CitationRef\"\u003e2010\u003c/span\u003e) and in green-veined white butterfly \u003cem\u003ePieris napi\u003c/em\u003e (G\u0026uuml;nter et al. \u003cspan citationid=\"CR15\" class=\"CitationRef\"\u003e2020\u003c/span\u003e), following the converse Bergmann\u0026rsquo;s rule.\u003c/p\u003e \u003cp\u003eTo date, only a limited number of studies have simultaneously examined all four traits when investigating life history variation along a latitudinal gradient. Building upon previous studies (Tang et al. \u003cspan citationid=\"CR48\" class=\"CitationRef\"\u003e2017\u003c/span\u003e), this study extends the investigation into the latitudinal clines in developmental time, body weight, growth rate and fecundity of the cabbage beetle \u003cem\u003eC. bowringi\u003c/em\u003e. The specimens were collected from six sites spanning a 21\u0026ordm; latitudinal range, with an expanded scope of study content and increased sample size. Examining the latitudinal clines in life-history traits is crucial for understanding how organisms adapt to seasonal environments and for predicting their potential responses to climate change.\u003c/p\u003e"},{"header":"Materials and methods","content":"\u003cp\u003e \u003cb\u003eLife history of\u003c/b\u003e \u003cb\u003eC. bowringi\u003c/b\u003e\u003c/p\u003e \u003cp\u003eThe cabbage beetle \u003cem\u003eC. bowringi\u003c/em\u003e, is a leaf-feeding pest primarily affecting cruciferous plants and is widely distributed across China. This study examines six populations that exhibit diverse life histories. Populations from lower latitudes (LN and XS) demonstrate multivoltine annual life history (Xue et al. \u003cspan citationid=\"CR53\" class=\"CitationRef\"\u003e2002\u003c/span\u003ea; Lai et al. \u003cspan citationid=\"CR27\" class=\"CitationRef\"\u003e2008\u003c/span\u003e). In the field, these populations display two distinct infestation peaks, one in spring with a single generation and another in autumn with three generations. Both populations undergo an aestivating and hibernating imaginal diapause in the soil. Temperature-dependent short-day response are observed, where shorter day lengths coupled with higher temperature lead to non-diapause development, while longer daylengths coupled with higher temperature induce diapause (Xue et al. \u003cspan citationid=\"CR53\" class=\"CitationRef\"\u003e2002\u003c/span\u003eb; Lai et al. \u003cspan citationid=\"CR27\" class=\"CitationRef\"\u003e2008\u003c/span\u003e). Regardless of photoperiod, diapause is triggered when mean daily temperature drop to \u0026le;\u0026thinsp;18\u0026deg;C for LN population or \u0026le;\u0026thinsp;20\u0026deg;C for XS population. Populations from mid-latitudes (XY and TA) exhibit bivoltine annual life history (Lai et al. \u003cspan citationid=\"CR27\" class=\"CitationRef\"\u003e2008\u003c/span\u003e; Tang et al. \u003cspan citationid=\"CR48\" class=\"CitationRef\"\u003e2017\u003c/span\u003e). These populations also show two distinct infestation peaks, one in spring and another in autumn, each with a single generation. They undergo imaginal summer and winter diapause in the soil. Unlike lower latitude populations, these populations lack photoperiodic response, and all individuals are induced into diapause at temperatures below 25\u0026deg;C (Tang et al. \u003cspan citationid=\"CR48\" class=\"CitationRef\"\u003e2017\u003c/span\u003e; Lai et al. \u003cspan citationid=\"CR27\" class=\"CitationRef\"\u003e2008\u003c/span\u003e). Higher latitude populations (SY and HB) exhibit univoltine annual life history. Only one generation is produced in summer, and adults overwinter in the soil as diapausing adults. All almost individuals enter diapause at \u0026le;\u0026thinsp;28\u0026deg;C regardless of photoperiod (Lai et al. \u003cspan citationid=\"CR27\" class=\"CitationRef\"\u003e2008\u003c/span\u003e; Chen et al. \u003cspan citationid=\"CR7\" class=\"CitationRef\"\u003e2014\u003c/span\u003e; He et al. \u003cspan citationid=\"CR18\" class=\"CitationRef\"\u003e2021\u003c/span\u003e ). Lai et al. (\u003cspan citationid=\"CR27\" class=\"CitationRef\"\u003e2008\u003c/span\u003e) reported that under controlled conditions of 25\u0026deg;C and L12:D12 photoperiod, the incidence of diapause increased with increasing latitude, indicating a latitudinal cline in diapause incidence.\u003c/p\u003e \u003cdiv id=\"Sec3\" class=\"Section2\"\u003e \u003ch2\u003eCollection sites and insect culture\u003c/h2\u003e \u003cp\u003eIn spring of 2022, naturally diapausing adults of \u003cem\u003eC. bowringi\u003c/em\u003e were collected from six locations: Longnan County (LN, 24\u0026deg;9' N, 114\u0026deg;8' E, as LN population), Xiushui County (XS, 29\u0026deg;1' N, 114\u0026deg;4' E, as XS population), Xinyang County (XY, 31\u0026deg;48' N, 114\u0026deg;03' E, as XY population), Taian City (TA, 36\u0026deg;2' N, 117\u0026deg;1' E, as TA population), Shenyang City (SY, 41\u0026deg;48' N, 123\u0026deg;23' E) and Harbin City (HB, 45\u0026deg;8' N, 126\u0026deg;6' E) (see Fig.\u0026nbsp;\u003cspan refid=\"Fig1\" class=\"InternalRef\"\u003e1\u003c/span\u003e). Diapause adults from each population were transferred into glass bottles (diameter 18 cm; height 32 cm) containing soil, allowing them to burrow for dormancy The bottles were maintained under semi-natural conditions (a covered, open-air room with natural temperature and photoperiod fluctuations) at Jiangxi Agricultural University in Nanchang, Jiangxi Province (28\u0026deg;46' N, 115\u0026deg;59' E). In spring 2023, post-diapausing adults emerged from the soil, and males and females from each population were weighed using an electric balance (AUY120, SHIMADZU Corporation, Japan). Females and males were paired randomly in petri dishes (9.0 cm diameter, 2.0 cm height) lined with filter paper and fresh leaves of the potherb mustard (\u003cem\u003eBrassica juncea\u003c/em\u003e (L.) Czern. et Coss. var. multiceps Tsen et Lee) for mating and oviposition. The lifetime number of eggs laid by each pair was recorded (see Table \u003cspan refid=\"MOESM1\" class=\"InternalRef\"\u003eS1\u003c/span\u003e). Eggs laid with the first 3 days were collected for subsequent experiments and checked daily until hatching.\u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003c/div\u003e\n\u003ch3\u003eExperimental design\u003c/h3\u003e\n\u003cp\u003eUp hatching, larvae for each population were transferred to rearing boxes (16 \u0026times; 11 \u0026times; 5.5 cm) containing fresh potherb mustard leaves. Each box housed 80 larvae, and the boxes were cleaned daily with fresh leaves provided as needed. Each population had three to four groups, with each group consisting of 1\u0026ndash;2 boxes. After reaching maturity, larvae were individually placed in cell culture plates with 12 holes (hole: diameter: 2.4 cm, height: 2 cm) for pupation and emergence. Pupation time and adult emergence times were observed every morning.\u003c/p\u003e \u003cp\u003eFor each individual, we recorded the development time from hatching to pupation and to adult emergence and weighed the pupa and adult. Growth rate, proportional weight loss during metamorphosis, and sexual size dimorphism (SSD) were calculated. Pupae were weighed the next day after pupation, and adults were weighed on the day of emergence after their elytra hardened. Growth rate was computed as ln pupal weight / larval development times (Gotthard et al. \u003cspan citationid=\"CR14\" class=\"CitationRef\"\u003e1994\u003c/span\u003e). Weight loss ratio was calculated as l-(adult weight/pupal weight). The SSD was estimated using a sexual dimorphism index (Lovich and Gibbons \u003cspan citationid=\"CR29\" class=\"CitationRef\"\u003e1992\u003c/span\u003e), defined as SSD = (size of the larger sex/size of the smaller sex)-1. Overall, 1811 offspring (903 females and 908 males) from six populations were raised to adulthood and weighed.\u003c/p\u003e \u003cp\u003eAll experiments were conducted in illuminated incubators (LRH-250-GS, Guangdong Medical Appliances Plant, Guangdong,China). The temperature was maintained at 22\u0026deg;C with a photoperiod of L15:D9 h. Under these conditions, all individuals were induced into adult diapause.\u003c/p\u003e\n\u003ch3\u003eStatistical analyses\u003c/h3\u003e\n\u003cp\u003eStatistical analyses were conducted using IBM SPSS Statistics 22.0. A linear mixed model (LMM) was utilized to assess the effects of population, sex, and their interaction on life-history traits. In this model, population and sex were treated as fixed factors, while rearing group was considered a random factor. The model was specified as: Trait\u0026thinsp;~\u0026thinsp;population * sex + (1|rearing group). Residual diagnostics confirmed that the assumptions of normality and homoscedasticity were met. To further investigate population-level differences identified by the LMM, one-way ANOVA followed by Tukey\u0026rsquo;s HSD post hoc tests were employed. Independent samples t-tests, with Welch\u0026rsquo;s correction for unequal variances, were used to compare traits between sexes. Linear regression analyses were conducted to examine the relationships between latitude and both fecundity and sexual size dimorphism (SSD).\u003c/p\u003e"},{"header":"Results","content":"\u003cdiv id=\"Sec7\" class=\"Section2\"\u003e \u003ch2\u003eBody weight and fecundity in post-diapause mated females\u003c/h2\u003e \u003cp\u003e \u003cb\u003eS\u003c/b\u003eignificant differences were observed in body weight (F\u003csub\u003e5, 116\u003c/sub\u003e = 109.116, \u003cem\u003eP\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.001) and fecundity (F\u003csub\u003e5, 116\u003c/sub\u003e = 8.557, \u003cem\u003eP\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.001) among population. Both body weight and fecundity showed a stepwise decline with increasing latitude (Fig.\u0026nbsp;\u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e2\u003c/span\u003eA), forming a distinct latitudinal gradient. Low-latitude populations maintained consistent body weight and fecundity levels, while mid- and high-latitude latitude populations showed decreasing trends. This pattern aligns with the converse Bergmann\u0026rsquo;s rule. A significant positive correlation was found between female body weight and fecundity (Fig.\u0026nbsp;\u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e2\u003c/span\u003eB, see also Table \u003cspan refid=\"MOESM2\" class=\"InternalRef\"\u003eS2\u003c/span\u003e).\u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec8\" class=\"Section2\"\u003e \u003ch2\u003eProgeny development time\u003c/h2\u003e \u003cp\u003eLarval development time was significantly influenced by population (Table\u0026nbsp;\u003cspan refid=\"Tab1\" class=\"InternalRef\"\u003e1\u003c/span\u003e), but not by sex or the interaction population between population and sex. Larval development time increased stepwise with increasing latitude (Fig.\u0026nbsp;\u003cspan refid=\"Fig3\" class=\"InternalRef\"\u003e3\u003c/span\u003eA), exemplifying cogradient variation. In contrast, pupal development time was unaffected by population, sex, or their interactions (Table\u0026nbsp;\u003cspan refid=\"Tab1\" class=\"InternalRef\"\u003e1\u003c/span\u003e, Fig.\u0026nbsp;\u003cspan refid=\"Fig3\" class=\"InternalRef\"\u003e3\u003c/span\u003eB).\u003c/p\u003e \u003cp\u003e \u003cdiv class=\"gridtable\"\u003e\u003ctable float=\"Yes\" id=\"Tab1\" border=\"1\"\u003e \u003ccaption language=\"En\"\u003e \u003cdiv class=\"CaptionNumber\"\u003eTable 1\u003c/div\u003e \u003cdiv class=\"CaptionContent\"\u003e \u003cp\u003eResults from a linear mixed-model analysis of fixed effects on larval time, pupal time, pupal weight, growth rate, adult weight and proportionate weight loss in \u003cem\u003eColaphellus bowringi\u003c/em\u003e in relation to population and sex\u003c/p\u003e \u003c/div\u003e \u003c/caption\u003e \u003ccolgroup cols=\"5\"\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c1\" colnum=\"1\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c2\" colnum=\"2\"\u003e\u003c/div\u003e \u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c3\" colnum=\"3\"\u003e\u003c/div\u003e \u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c4\" colnum=\"4\"\u003e\u003c/div\u003e \u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c5\" colnum=\"5\"\u003e\u003c/div\u003e \u003cthead\u003e \u003ctr\u003e \u003cth align=\"left\" colname=\"c1\"\u003e \u003cp\u003eLife-history traits\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c2\"\u003e \u003cp\u003eFixed factors\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c3\"\u003e \u003cp\u003e\u003cem\u003edf\u003c/em\u003e\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c4\"\u003e \u003cp\u003e\u003cem\u003eF\u003c/em\u003e -value\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c5\"\u003e \u003cp\u003e\u003cem\u003eP\u003c/em\u003e -value\u003c/p\u003e \u003c/th\u003e \u003c/tr\u003e \u003c/thead\u003e \u003ctbody\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\" morerows=\"2\" rowspan=\"3\"\u003e \u003cp\u003eLarval time\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003ePopulation\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e5\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e776.439\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e\u0026lt;\u0026thinsp;0.001\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eSex\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e1\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e0.305\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e0.581\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003ePopulation \u0026times; sex\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e5\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e0.416\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e0.838\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\" morerows=\"2\" rowspan=\"3\"\u003e \u003cp\u003ePupal time\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003ePopulation\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e5\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e1.848\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e0.100\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eSex\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e1\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e0.031\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e0.861\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003ePopulation \u0026times; sex\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e5\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e0.175\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e0.972\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\" morerows=\"2\" rowspan=\"3\"\u003e \u003cp\u003ePupal weight\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003ePopulation\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e5\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e905.185\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e\u0026lt;\u0026thinsp;0.001\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eSex\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e1\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e2969.607\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e\u0026lt;\u0026thinsp;0.001\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003ePopulation \u0026times; sex\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e5\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e24.300\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e\u0026lt;\u0026thinsp;0.001\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\" morerows=\"2\" rowspan=\"3\"\u003e \u003cp\u003eGrowth rate\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003ePopulation\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e5\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e1521.485\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e\u0026lt;\u0026thinsp;0.001\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eSex\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e1\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e1129.139\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e\u0026lt;\u0026thinsp;0.001\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003ePopulation \u0026times; sex\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e5\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e3.552\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e0.003\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\" morerows=\"2\" rowspan=\"3\"\u003e \u003cp\u003eAdult weight\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003ePopulation\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e5\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e760.767\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e\u0026lt;\u0026thinsp;0.001\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eSex\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e1\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e3724.932\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e\u0026lt;\u0026thinsp;0.001\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003ePopulation \u0026times; sex\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e5\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e30.422\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e\u0026lt;\u0026thinsp;0.001\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\" morerows=\"2\" rowspan=\"3\"\u003e \u003cp\u003eProportionate weight loss\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003ePopulation\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e5\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e43.497\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e\u0026lt;\u0026thinsp;0.001\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eSex\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e1\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e459.878\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e\u0026lt;\u0026thinsp;0.001\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003ePopulation \u0026times; sex\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e5\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e10.941\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e\u0026lt;\u0026thinsp;0.001\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003c/tbody\u003e \u003c/colgroup\u003e \u003c/table\u003e\u003c/div\u003e \u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003c/div\u003e\n\u003ch3\u003eProgeny pupal weight and growth rate\u003c/h3\u003e\n\u003cp\u003ePupal weight and growth rate were significantly affected by population, sex and their interactions (Table\u0026nbsp;\u003cspan refid=\"Tab1\" class=\"InternalRef\"\u003e1\u003c/span\u003e). Both Pupal weight and growth rate decreased stepwise with increasing latitude, adhering to the converse Bergmann\u0026rsquo;s rule or cogradient variation (Fig.\u0026nbsp;\u003cspan refid=\"Fig4\" class=\"InternalRef\"\u003e4\u003c/span\u003eA and \u003cspan refid=\"Fig4\" class=\"InternalRef\"\u003e4\u003c/span\u003eB). Across all populations, female pupae exhibited significantly higher body weight and growth rate compared to male pupae (see Table \u003cspan refid=\"MOESM3\" class=\"InternalRef\"\u003eS3\u003c/span\u003e, \u003cem\u003eP\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.05).\u003c/p\u003e \u003cp\u003e \u003c/p\u003e\n\u003ch3\u003eProgeny adult weight and weight loss\u003c/h3\u003e\n\u003cp\u003eAdult weight and weight loss were significantly influenced by population, sex and their interactions (Table\u0026nbsp;\u003cspan refid=\"Tab1\" class=\"InternalRef\"\u003e1\u003c/span\u003e). Both female and male adults showed a stepwise decline in weight with increasing latitude (Fig.\u0026nbsp;\u003cspan refid=\"Fig5\" class=\"InternalRef\"\u003e5\u003c/span\u003eA), confirming a converse Bergmann\u0026rsquo;s cline. Male adults experienced significantly greater weight loss than females (Fig.\u0026nbsp;\u003cspan refid=\"Fig5\" class=\"InternalRef\"\u003e5\u003c/span\u003eB, see Table \u003cspan refid=\"MOESM3\" class=\"InternalRef\"\u003eS3\u003c/span\u003e). For males, the highest weight loss occurred at low latitudes, followed by high and middle latitudes. For females, the weight loss gradually decreased with increasing latitude, indicating a latitudinal cline. The higher weight loss in low-latitudinal populations suggests that larger individuals lose more weight during metamorphosis compared to smaller ones.\u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003cdiv id=\"Sec11\" class=\"Section2\"\u003e \u003ch2\u003eSexual size dimorphism (SSD)\u003c/h2\u003e \u003cp\u003eSSD for both pupae and adults varied substantially with latitude (Fig.\u0026nbsp;\u003cspan refid=\"Fig6\" class=\"InternalRef\"\u003e6\u003c/span\u003eA). The SSD index for pupae was lower than that for adults, particularly in low-latitude populations. Higher latitudes exhibited lower SSD indices compared to middle and low latitudes. Linear regression analysis revealed a negative correlation between SSD and Latitude (Fig.\u0026nbsp;\u003cspan refid=\"Fig6\" class=\"InternalRef\"\u003e6\u003c/span\u003eB).\u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003c/div\u003e"},{"header":"Discussion","content":"\u003cp\u003eOur study revealed for the first time that adult body weigh and fecundity \u003cem\u003ein C. bowringi\u003c/em\u003e decreased with increasing latitude after diapause, and female body weight and fecundity were significantly positively correlated (Fig.\u0026nbsp;\u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e2\u003c/span\u003e). This latitudinal cline in body weight and fecundity is converse to Bergmann\u0026rsquo;s cline. Notably, the latitudinal clines in body weight and fecundity exhibited a stepwise decline pattern. Specifically, populations with multivoltine annual life history at low latitudes (LN and XS) showed similar body weight and fecundity. Similarly, mid-latitude populations (XY and TA) with bivoltine annual life history exhibited comparable body weight and fecundity, as did high-latitude populations (SY and HB) with univoltine annual life history. This stepwise pattern has only been reported in two other species (Shelomi \u003cspan citationid=\"CR42\" class=\"CitationRef\"\u003e2012\u003c/span\u003e), suggesting that voltinism plays a crucial role in determining life-history patterns. Furthermore, the weight of the pupae and adults produced by post-diapause individuals also decreased in a stepwise manner with increasing latitude. (Fig.\u0026nbsp;\u003cspan refid=\"Fig5\" class=\"InternalRef\"\u003e5\u003c/span\u003eA and Fig.\u0026nbsp;\u003cspan refid=\"Fig6\" class=\"InternalRef\"\u003e6\u003c/span\u003eA).\u003c/p\u003e \u003cp\u003eAdditionally, we observed a latitudinal cline in larval development time and growth rate. Larval development time increased significantly in a stepwise manner with increasing latitude (Fig.\u0026nbsp;\u003cspan refid=\"Fig3\" class=\"InternalRef\"\u003e3\u003c/span\u003eA), while growth rate decreased significantly in a stepwise manner (Fig.\u0026nbsp;\u003cspan refid=\"Fig4\" class=\"InternalRef\"\u003e4\u003c/span\u003eB). Both variables exhibited cogradient variation, consistent with findings in \u003cem\u003eH. merope\u003c/em\u003e, where low-latitude populations had faster growth and shorter development time compared to high-latitude populations (Barton et al. \u003cspan citationid=\"CR3\" class=\"CitationRef\"\u003e2014\u003c/span\u003e). Unlike the \u003cem\u003eC. bowringi\u003c/em\u003e, \u003cem\u003eH. merope\u003c/em\u003e did not show latitudinal cline in body size, indicating that selection pressures on development time and body weight may follow different evolutionary pathways.\u003c/p\u003e \u003cp\u003eWe also found significant variation across populations in the relationship between larval development and body weight. Low-latitude populations (LN and XS) had the shortest development time and largest body weight while mid-latitude populations (XY and TA) exhibited intermediate values, and high-latitude populations (SY and HB) had the longest development time and smallest body weight. These findings align with previous studies (Tang et al. \u003cspan citationid=\"CR48\" class=\"CitationRef\"\u003e2017\u003c/span\u003e; He et al. \u003cspan citationid=\"CR18\" class=\"CitationRef\"\u003e2021\u003c/span\u003e). Generally, larval development time is positively correlated with body size, that is, animals with longer development time are larger at maturity (Nijhout et al. \u003cspan citationid=\"CR33\" class=\"CitationRef\"\u003e2010\u003c/span\u003e) or take longer time to grow larger (Roff \u003cspan citationid=\"CR37\" class=\"CitationRef\"\u003e2000\u003c/span\u003e; Stearns \u003cspan citationid=\"CR45\" class=\"CitationRef\"\u003e1992\u003c/span\u003e), but this assumption does not hold when comparing populations from different regions. For instance, in \u003cem\u003eO. furnacalis\u003c/em\u003e for the diapausing generation, the high-latitude populations exhibited significantly longer larval development time and lower body weight compared to low-latitude populations (Fu et al. \u003cspan citationid=\"CR12\" class=\"CitationRef\"\u003e2022\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eThe converse Bergmann\u0026rsquo;s cline (decreasing body size with increasing latitude) is typically attributed to season length effect (Blanckenhorn and Demont \u003cspan citationid=\"CR5\" class=\"CitationRef\"\u003e2004\u003c/span\u003e). However, in our study, the converse Bergmann\u0026rsquo;s cline in \u003cem\u003eC. bowringi\u003c/em\u003e is unlikely to be associated with gradual changes in seasonality. Reproductive activity in high-latitude SY and HB populations occurs during summer, providing sufficient time for larvae to develop without seasonal constraints. The significantly longer development times and small body weights in northern populations may instead be related to local climatic conditions rather than season length. The stepwise clines in development time and body weight in \u003cem\u003eC. bowringi\u003c/em\u003e suggest genetic regional differences in life-history traits.\u003c/p\u003e \u003cp\u003eBased on Horne et al. (\u003cspan citationid=\"CR20\" class=\"CitationRef\"\u003e2015\u003c/span\u003e), voltinism in arthropods is strongly correlated with body weight, with univoltine species often being larger than multivoltine species. However, this rule does not apply to \u003cem\u003eC. bowringi\u003c/em\u003e. In this study, the multivoltine populations (LN and XS) had the largest body weight, followed by bivoltine populations (XY and TA), with univoltine populations (e SY and HB) having the smallest body weight. Our results suggest that relationship between voltinism and body size can vary depending on the species.\u003c/p\u003e \u003cp\u003eOur study demonstrated that, across all populations, pupae lost significantly more weight during metamorphosis compared to females (Fig.\u0026nbsp;\u003cspan refid=\"Fig5\" class=\"InternalRef\"\u003e5\u003c/span\u003eB). This suggests that males either have a higher metabolic rate or are less effective at preventing water loss (Savalli and Fox \u003cspan citationid=\"CR39\" class=\"CitationRef\"\u003e1998\u003c/span\u003e). Consequently, the SSD in pupae is significantly lower than that in adults, indicating that sex-specific weight loss mediated SSD \u003cem\u003ein C. bowringi\u003c/em\u003e (Testa et al. \u003cspan citationid=\"CR50\" class=\"CitationRef\"\u003e2013\u003c/span\u003e). Additionally, we observes that female weight loss exhibited a gradually downward latitudinal gradient, while male weight loss followed a U-shaped along the latitudinal gradient, with the mid-latitude populations experiencing the least weight loss (Fig.\u0026nbsp;\u003cspan refid=\"Fig5\" class=\"InternalRef\"\u003e5\u003c/span\u003eB).\u003c/p\u003e \u003cp\u003eAccording to Rensch\u0026rsquo;s rule, SSD tends to increase with body size when males are the larger sex, and decrease with body size when females are the larger sex (Abouheif and Fairbairn \u003cspan citationid=\"CR1\" class=\"CitationRef\"\u003e1997\u003c/span\u003e, Fairbairn \u003cspan citationid=\"CR9\" class=\"CitationRef\"\u003e1997\u003c/span\u003e). In our study, for female-biased cabbage beetles reared at 22\u0026deg;C, SSD tended to increase with increasing body weight, and SSD index tended to decrease with increasing latitude, which appears in inconsistent with Rensch\u0026rsquo;s rule (Teder and Tammaru \u003cspan citationid=\"CR49\" class=\"CitationRef\"\u003e2005\u003c/span\u003e). Similar finding was reported in the male-biased beetle \u003cem\u003eStator limbatus\u003c/em\u003e, where both males and females increased with increasing latitude, but dimorphism decreased with increasing latitude, also not supporting Rensch\u0026rsquo;s rule (Stillwell et al., \u003cspan citationid=\"CR46\" class=\"CitationRef\"\u003e2007\u003c/span\u003e).\u003c/p\u003e"},{"header":"Conclusion","content":"\u003cp\u003eOur study revealed latitudinal clines in body size, fecundity, development time, growth rate, weight loss and SSD \u003cem\u003ein C. bowringi\u003c/em\u003e. Body size and fecundity exhibited converse Bergmann\u0026rsquo;s cline with a stepwise decline pattern, while development time and growth rate exhibited cogradient variation, with development time increasing stepwise and growth rate decreasing stepwise with latitude. These patterns appear to be related to voltinism, as differences in voltinism across geographical populations collectively construct the stepwise pattern. We also found that female weight loss tended to decrease with increasing latitude, while male weight loss was the lowest in mid-latitude. The SSD index for both pupae and adults tended to decrease with latitude. These joint results indicate that this species comprises regionally adapted populations. These findings broaden our understanding of latitudinal variation in life-history traits in insects.\u003c/p\u003e"},{"header":"Declarations","content":"\u003cp\u003e\u003cstrong\u003eSupplementary Information\u003c/strong\u003e The online version contains supplementary material\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eAcknowledgement\u0026nbsp;\u003c/strong\u003eWe thank\u003cstrong\u003e\u0026nbsp;\u003c/strong\u003ethe National Natural Science Foundation of China for its financial support of this project.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eFunding\u0026nbsp;\u003c/strong\u003eThe work has been funded by the\u003cstrong\u003e\u0026nbsp;\u003c/strong\u003eNational Natural Science Foundation of China (32360270) and High-level Scientific Research and Innovation Team of Yuzhang Normal University (YZTD202305).\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eAuthor contributions\u003c/strong\u003e HH and XF conceived the ideas and designed the study. HH, HL and XF performed this experiment. XF and HL wrote the original manuscript. HH and TJ analyzed the data. All authors have read and agreed to the published version of the manuscript.\u0026nbsp;\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eAvailability Statement of data and materials\u0026nbsp;\u003c/strong\u003eData are contained within the article.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eCode availability \u0026nbsp;Not available\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eConflicts of Interest:\u0026nbsp;\u003c/strong\u003eThe authors declare no conflict of interest.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eEthical approval\u0026nbsp;\u003c/strong\u003e\u003cstrong\u003eNot applicable.\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eConsent to participate\u0026nbsp;\u003c/strong\u003e\u003cstrong\u003eNot applicable.\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eConsent for publication\u0026nbsp;\u003c/strong\u003e\u003cstrong\u003eNot applicable.\u003c/strong\u003e\u003c/p\u003e"},{"header":"References","content":"\u003col\u003e\u003cli\u003e\u003cspan\u003eAbouheif E, Fairbairn DJ (1997) A comparative analysis of allometry for sexual size dimorphism: assessing Rensch's rule. Am Nat 149:540\u0026ndash;562\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eAtkinson D, Sibly RM (1997) Why are organisms usually bigger in colder environments? Making sense of a life history puzzle. Trends Ecol Evol 12:235\u0026ndash;239\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eBarton M, Sunnucks P, Norgate M, Murray N, Kearney M (2014) Co-Gradient variation in growth rate and development time of a broadly distributed butterfly. PLoS ONE 9:1\u0026ndash;8\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eBlanckenhorn WU (2000) The evolution of body size: what keeps organisms small? Q Rev Biol 75:385\u0026ndash;407\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eBlanckenhorn WU, Demont M (2004) Bergmann and converse Bergmann latitudinal clines in arthropods: two ends of a continuum? Integr Comp Biol 44:413\u0026ndash;424\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eBlanckenhorn WU, Bauerfeind SS, Berger D, Davidowitz G, Fox CW, Guillaume F et al (2018) Life history traits, but not body size, vary systematically along latitudinal gradients on three continents in the widespread yellow dung fly. Ecography 41:2080\u0026ndash;2091\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eChen C, Xiao L, He HM, Xu J, Xue FS (2014) A genetic analysis of diapause in crosses of a southern and a northern strain of the cabbage Colaphellus bowringi (Coleoptera: Chrysomelidae). B Entomol Res 104:586\u0026ndash;591\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eConover DO, Duffy TA, Hice LA (2009) The covariance between genetic and environmental influences across ecological gradients: reassessing the evolutionary significance of countergradient and cogradient variation. 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Acta Zool\u0026ndash;Stockholm 96:242\u0026ndash;252\u003c/span\u003e\u003c/li\u003e\u003c/ol\u003e"}],"fulltextSource":"","fullText":"","funders":[],"hasAdminPriorityOnWorkflow":false,"hasManuscriptDocX":true,"hasOptedInToPreprint":true,"hasPassedJournalQc":"","hasAnyPriority":false,"hideJournal":false,"highlight":"","institution":"","isAcceptedByJournal":true,"isAuthorSuppliedPdf":false,"isDeskRejected":"","isHiddenFromSearch":false,"isInQc":false,"isInWorkflow":true,"isPdf":false,"isPdfUpToDate":true,"isWithdrawnOrRetracted":false,"journal":{"display":true,"email":"[email protected]","identity":"oecologia","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":false,"externalIdentity":"oeco","sideBox":"Learn more about [Oecologia](https://www.springer.com/journal/442)","snPcode":"442","submissionUrl":"https://submission.nature.com/new-submission/442/3","title":"Oecologia","twitterHandle":"","acdcEnabled":true,"dfaEnabled":true,"editorialSystem":"em","reportingPortfolio":"Springer Hybrid","inReviewEnabled":true,"inReviewRevisionsEnabled":false},"keywords":"Latitudinal cline, Body weight, Development time, Growth rate, Fecundity","lastPublishedDoi":"10.21203/rs.3.rs-6116757/v1","lastPublishedDoiUrl":"https://doi.org/10.21203/rs.3.rs-6116757/v1","license":{"name":"CC BY 4.0","url":"https://creativecommons.org/licenses/by/4.0/"},"manuscriptAbstract":"\u003cp\u003eStudying the latitudinal cline in life-history traits is crucial for understanding how organisms adapt to seasonal environments and for predicting their potential responses to climate change. In this study, we systematically examined the life-history traits of the cabbage beetle \u003cem\u003eColaphellus bowringi\u003c/em\u003e collected from six sites spanning a 21\u0026ordm; latitudinal range. Our results demonstrated that post-diapause female body weight and fecundity decreased in a stepwise manner with increasing latitude, consistent with the converse Bergmann\u0026rsquo;s rule. This pattern was also found in pupal and adult weight of their offspring. Larval development time increased while growth rate decreased in a stepwise manner with increasing latitude, indicating cogradient variation. We further found that these stepwise changes are associated with voltinism. Specifically, multivoltine populations exhibited one set of life-history trait pattern, bivoltine populations another, and univoltine populations yet another, collectively forming a stepwise pattern. Additionally, male pupae experienced significantly greater weight loss during metamorphosis compared to female pupae, resulting in lower sexual size dimorphism (SSD) in pupae than in adults. This suggests that sex-specific weight loss during metamorphosis mediates SSD. In summary, our study provides a comprehensive example of insect life-history evolution, particularly in the empirical study of stepped variation patterns. These findings enhance our understanding of latitudinal variation in life-history traits.\u003c/p\u003e","manuscriptTitle":"Latitudinal clines in life-history traits of the cabbage beetle, Colaphellus bowringi: showing a stepwise pattern","msid":"","msnumber":"","nonDraftVersions":[{"code":1,"date":"2025-03-31 12:44:22","doi":"10.21203/rs.3.rs-6116757/v1","editorialEvents":[{"type":"communityComments","content":0},{"type":"reviewerAgreed","content":"","date":"2025-05-30T07:44:31+00:00","index":0,"fulltext":""},{"type":"reviewersInvited","content":"","date":"2025-03-21T09:07:54+00:00","index":"","fulltext":""},{"type":"editorAssigned","content":"","date":"2025-02-28T11:47:47+00:00","index":"","fulltext":""},{"type":"submitted","content":"Oecologia","date":"2025-02-26T21:16:56+00:00","index":"","fulltext":""}],"status":"published","journal":{"display":true,"email":"[email protected]","identity":"oecologia","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":false,"externalIdentity":"oeco","sideBox":"Learn more about [Oecologia](https://www.springer.com/journal/442)","snPcode":"442","submissionUrl":"https://submission.nature.com/new-submission/442/3","title":"Oecologia","twitterHandle":"","acdcEnabled":true,"dfaEnabled":true,"editorialSystem":"em","reportingPortfolio":"Springer Hybrid","inReviewEnabled":true,"inReviewRevisionsEnabled":false}}],"origin":"","ownerIdentity":"f981a51a-ee12-4819-9620-56a1f164c04c","owner":[],"postedDate":"March 31st, 2025","published":true,"recentEditorialEvents":[],"rejectedJournal":[],"revision":"","amendment":"","status":"published-in-journal","subjectAreas":[],"tags":[],"updatedAt":"2025-12-15T16:04:29+00:00","versionOfRecord":{"articleIdentity":"rs-6116757","link":"https://doi.org/10.1007/s00442-025-05849-3","journal":{"identity":"oecologia","isVorOnly":false,"title":"Oecologia"},"publishedOn":"2025-12-09 15:57:27","publishedOnDateReadable":"December 9th, 2025"},"versionCreatedAt":"2025-03-31 12:44:22","video":"","vorDoi":"10.1007/s00442-025-05849-3","vorDoiUrl":"https://doi.org/10.1007/s00442-025-05849-3","workflowStages":[]},"version":"v1","identity":"rs-6116757","journalConfig":"researchsquare"},"__N_SSP":true},"page":"/article/[identity]/[[...version]]","query":{"redirect":"/article/rs-6116757","identity":"rs-6116757","version":["v1"]},"buildId":"8U1c8b4HqxoKbykW_rLl7","isFallback":false,"isExperimentalCompile":false,"dynamicIds":[84888],"gssp":true,"scriptLoader":[]}