Developmental and post-embryonic growth patterns in Longidorus piceicola Lišková, Robbins & Brown, 1997 and L. intermedius Kozlowska & Seinhorst, 1979 (Nematoda: Dorylaimida)

E-mail: laslo[email protected]s Developmental and post-embryonic growth patterns in Longidorus piceicola Lišková, Robbins & Brown, 1997 and L. intermedius Kozlowska & Seinhorst, 1979 (Nematoda: Dorylaimida) Barsi László University of Novi Sad, Faculty of Sciences, Department of Biology and Ecology, Trg Dositeja Obradovića 2, 21000 Novi Sad, Serbia Biologia Serbica, 2025, 47: 0-0 Original paper DOI: 10.5281/zenodo.17872808 Summary. Based solely on molecular markers, D-D3 28R rDNA and ITS1, Longidorus intermedius and L. piceicola share a high level of similarity. However, despite this molecular similarity, L. piceicola and L. intermedius significantly differ in morphometrics and ontogeny. Two key concepts in the development and growth of longidorid nematodes are developmental and growth patterns. Developmental patterns categorize longidorid nematodes according to whether they undergo four or three juvenile developmental stages (JDS) during post-embryonic development. Growth patterns refer to specific morphometric levels, which are more or less similar between species within a genus with the same developmental pattern. There are some differences between different populations of the same species, probably due to intraspecific variability and the environmental conditions in which they live. Longidorus piceicola undergoes four JDS, while L. intermedius undergoes three JDS. In the typical life cycle of longidorid nematodes, there are four or three molts, which occur between the four or three JDS, with the last molt preceding the fully sexually mature adult stage. The similarities and differences between these two developmental patterns are not well known. Growth patterns refer to changes in specific morphometric dimensions during the post-embryonic growth of a nematode, such as: body length, odontostyle and replacement odontostyle size, and body volume. Using these characteristics, a study was conducted to assess whether there are similarities or differences between selected populations of Longidorus piceicola (4 JDS) and L. intermedius (3 JDS). This paper presents results on the developmental and post-embryonic growth patterns of the species Longidorus piceicola and L. intermedius. Keywords: developmental patterns, growth patterns, juvenile stages, Longidoridae, morphometrics, ontogeny. Accepted: 3 December 2025 INTRODUCTION He et al. (2005), in a work dedicated to the phylogeny of the entire family Longidoridae, first noted that based on molecular markers, Longidorus intermedius shows great similarity to L. piceicola. Although this work was based only on the D-D3 28R rDNA marker, the authors did not add any additional comments. Subsequently, Kornobis (2013) and Groza et al. (2017) found that the difference between molecular markers (p-distance) was only 0.3% and 0.3-0.9%, respectively. Additionally, Kornobis (2013) found that the difference between the two species based on the ITS1 marker was 1.4%. Such low differences are usually interpreted as intraspecific variation, rather than as a unique feature of different species. Despite their similarity based on comparison of specific molecular markers, L. piceicola and L. intermedius display significant morphometric and ontogenic differences. Longidorus piceicola undergoes four juvenile developmental stages (JDS) (Liskova et al. 1997, Barsi and Lamberti, 2001; Kornobis and Peneva, 2011; Groza et al. 2017; Barsi 2022), while Published online: 11 December 2025 Developmental and post-embryonic growth patterns in Longidorus piceicola and L. intermedius L. intermedius undergoes three JDS (Peneva et al. 2001; Barsi and Lamberti 2004; Kumari et al. 2006; Susulovska and Tsaryk 2018). During a typical life-cycle, longidorid nematodes undergo four or three molts, which occur between the four or three juvenile developmental stages (JDS), respectively, with the final molt preceding full sexual maturity (Halbrendt and Brown 1992, 1993; Halbrendt et al. 1997). Although the presence of four or three JDS is a characteristic of the two developmental patterns present in Longidoridae; similarities and differences between these two developmental patterns are not well known. Yeates and Boag (2002) used a volumetric base for comparison of the successive stages of Longidoridae, in order to define post-embryonic growth patterns in longidorid nematodes. They concluded that within the Longidoridae there was generally a larger increase in body volume between the first two juvenile stages. Unfortunately, their results do not give a clear picture of the basic similarities and differences between the two developmental patterns obviously present in Longidoridae. Two important concepts in the development and growth of longidorid nematodes are developmental patterns and growth patterns (growth strategy). In longidorid nematodes, developmental patterns are dictated by the presence of either four or three JDS during post-embryonic development. Growth patterns are defined by specific morphometric levels, which are more or less similar between species within a genus with the same developmental pattern. There are some differences between different populations of the same species, probably due to intraspecific variability and the environmental conditions in which they live. Growth patterns describe the post-embryonic growth of a nematode, and defined as changes in certain measurable morphometric parameters, such as: body length, odontostyle and replacement odontostyle length, and body volume. Using these characteristics, a study was conducted to assess whether there are similarities or differences between selected populations of L. piceicola (4 JDS) and L. intermedius (3 JDS). The present study presents results on the developmental and postembryonic growth patterns of the species L. piceicola and L. intermedius. MATERIAL AND METHODS The present study was based on published data for 6 populations of L. piceicola and 4 populations of L. intermedius (Table 1). For each population, the following data were used: mean body length, odontostyle and replacement odontostyle lengths and body diameter for each developmental stage (all in µm). Body length and body diameter were used to calculate the body volume at every stage, using the empirical equation of Andrássy (1956): V = W2 x L/1.7, where W is the body diameter and L is the body length in µm. These data were tabulated for each population/species, and used for further calculations. A percentage method was used to ensure that the data sets are comparable between each other for every population of both species. Absolute data for females (body and odontostyle length in µm and calculated body volume in µm3) were defined as 100%. The same data set for each juvenile stage was then compared to the female data set and expressed as a percentage. The only exception was the replacement odontostyle length; where the absolute value of the replacement odontostyle length in the pre-adult stage (J4 or JIII, respectively) was defined as 100% for the comparisons. The percent values for each stage of each population/ species were tabulated and the mean value, standard deviation, minimum and maximum values were calculated for Table 1. List and sources of data for linear dimensions of 6 populations of Longidorus piceicola and 4 populations of L. intermedius. Longidorus species Reference L. piceicola1*1Montenegro, Durmitor 1Barsi and Lamberti 2011 L. piceicola2 2Kopaonik 2Barsi 2022 L. piceicola3 3Slovakia 3Lišková et al. 1997 L. piceicola4 4Romania, Bran population 4Groza et al. 2017 L. piceicola5 5Poland 5Kornobis and Peneva 2011 L. piceicola6 6Romania, Cernica 6Groza et al. 2017 L. intermedius7** 7Serbia 7Barsi and Lamberti 2004 L. intermedius8 8Bulgaria, Dedevo-Rhodopi 8Peneva et al. 2001 L. intermedius9 9Czech Republic 9Kumari et al. 2006 L. intermedius10 10Ukraine, Ustia, 10Susulovska and Tsaryk 2018 *Longidorus piceicola has 4 juvenile developmental stages (JDS); **L. interemdius has 3 JDS. L. Barsi each character studied. Also, using percent values for each stage, the relative increase in the analyzed morphometric characters between successive stages were calculated for each population/species. Analyses were conducted at the species level (intrageneric), separately for the populations of L. piceicola and L. intermedius, respectively. RESULTS AND DISCUSSION A comparison of selected characteristics of 6 populations of L. piceicola with 4 JDS, and 4 populations of L. intermedius with 3 JDS, revealed the presence of specific basic similarities and differences between these two species. The final data sets used in the present study are presented in Tables 2 and 3. Analyses of these data sets were conducted at the species (intra-generic) level. However, it must be strongly emphasized that these average values provide only an average picture of the relationships present between these species. Body length (Fig. 1A, B) Based on results for the analyzed data from the studied populations, it appears that in L. intermedius, body lengths for the JI, JII, and JIII stages are greater than the body lengths in the corresponding J1, J2, and J3 stages in L. piceicola. In general, in L. intermedius, the body length during JI is 1.2 times that of the body length of L. piceicola during J1, JII is 1.29 times that of J2, and JIII is 1.31 times that of J3, for L. intermedius vs. L. piceicola. Although inclusion of additional populations of both species would certainly affect the numerical values obtained, the trends would be expected to remain the same. Using the percentage body lengths for each stage, the relative increase from stage to stage was calculated for each species (Tables 2 and 3). The sum of the relative increases between successive stages is lower in L. intermedius (4.42) than in L. piceicola (5.62). This also supports the observation that for L. intermedius, specimens in stages JI, JII and JIII generally have a longer body length than L. piceicola specimens in stages J1, J2 and J3. Therefore, L. intermedius nema - todes can probably reach maturity more quickly, through only three molts. Body volume (Fig. 2A, B) It is known that L. intermedius reaches maturity through 3 JDS and L. piceicola through 4 JDS, which represent two different developmental patterns. Based on the results of the analyzed data from the studied populations, it appears that the body volumes during stages JI, JII and JIII in L. intermedius are larger than the body volumes in the corresponding stages J1, J2 and J3 in L. piceicola. In general, the body volume during JI for L. intermedius is 1.17 times the body volume during J1 for L. piceicola, while JII is 1.45 times the volume of J2, and JIII is 1.51 times the volume of J3 for L. intermedius vs. L. piceicola, respectively. Body volume is a useful metric for describing processes during postembryonic development in nematodes, but it should be used with caution, especially when comparing different populations of the same species or other species, and for drawing conclusions. Body volume depends on body length and width. Unlike body length, body width may be more affected by flattening during the mounting process. Therefore, uncritical use of published data may lead to erroneous conclusions. Including additional populations of both species would certainly affect the numerical values obtained, but the trends would be expected to remain the same. Using percent body volumes for each stage, the relative increase from stage to stage was calculated for each species (Tables 2 and 3). The sum of the relative increases between successive stages was lower in L. intermedius (7.99) than in L. piceicola (8.84). This also supports the results that L. intermedius specimens in stages JI, JII and JIII generally have a larger body volume than corresponding L. piceicola specimens in stages J1, J2 and J3. Odontostyle length (Fig. 3A, B) In L. intermedus, specimens in stages JI, JII and JIII generally have a 1.14, 1.12 and 1.11 fold longer odontostyle than corresponding L. piceicola specimens in stages J1, J2 and J3 (Fig. 3A). These JDSs are not directly comparable to each other (Tables 2 and 3). However, comparison of specimens in stages JI, JII, and JIII with specimens in stages J2, J3, and J4, respectively, reveals a very similar and comparable growth pattern (Fig. 3B). The odontostyle in JI is 1.67% longer than in J2, while in JII it is 4.02% shorter than in J3 and in JIII it is 2.06% shorter than in J4. Due to its longer odontostyle in the JI stage, L. intermedus can probably reach its final odontostyle length more quickly after only three molts. Including additional populations of both species would certainly affect the numerical values obtained, but the trends would be expected to remain the same. Using percent odontostyle lengths for each stage, the relative increase from stage to stage was calculated for both species (Tables 2 and 3). The sum of the relative increases between successive stages was smaller for L. intermedius (3.43) than for L. piceicola (4.57). This also supports the observation that L. intermedius Developmental and post-embryonic growth patterns in Longidorus piceicola and L. intermedius Fig. 1. Comparison of body length growth patterns in 6 populations of Longidorus piceicola and 4 populations of L. intermedius. L. Barsi Fig. 2. Comparison of body volume growth patterns in 6 populations of Longidorus piceicola and 4 populations of L. intermedius. Developmental and post-embryonic growth patterns in Longidorus piceicola and L. intermedius Table 2. Growth patterns of the body length, body volume, odontostyle length and replacement odontostyle length and relative increase in the body length, body volume, odontostyle length and replacement odontostyle length between successive stages in six populations of Longidorus piceicola*. Growing patterns Relative increase L% J1 J2 J3 J4 F L µm J2/J1 J3/J2 J4/J3 F/J4 Sum L. piceicola126.2 38.7 55.7 79.0 100.0 5953 1.48 1.44 1.42 1.27 5.60 L. piceicola229.2 34.6 54.6 72.2 100.0 6332 1.18 1.58 1.32 1.39 5.47 L. piceicola328.9 45.5 57.8 72.3 100.0 5190 1.57 1.27 1.25 1.38 5.48 L. piceicola426.9 37.3 53.5 70.3 100.0 4900 1.39 1.43 1.31 1.42 5.56 L. piceicola522.3 32.7 44.5 65.6 100.0 6477 1.47 1.36 1.47 1.52 5.83 L. piceicola623.0 33.6 51.8 67.2 100.0 5880 1.46 1.54 1.30 1.49 5.79 Avg 26.1 37.1 53.0 71.1 100.0 5789 1.42 1.44 1.35 1.41 5.62 Min 22.3 32.7 44.5 65.6 100.0 4900 1.18 1.27 1.25 1.27 5.47 Max 29.2 45.5 57.8 79.0 100.0 6477 1.57 1.58 1.47 1.52 5.83 Stdev 2.9 4.7 4.6 4.7 0 625 0.13 0.11 0.08 0.09 0.15 V% J1 J2 J3 J4 F V mm3J2/J1 J3/J2 J4/J3 F/J4 Sum L. piceicola14.4 12.0 28.7 63.6 100.0 0.0115 2.73 2.39 2.22 1.57 8.91 L. piceicola27.4 10.1 29.6 50.2 100.0 0.0117 1.36 2.93 1.70 1.99 7.98 L. piceicola35.8 18.8 31.0 50.9 100.0 0.0096 3.26 1.65 1.64 1.97 8.51 L. piceicola44.4 10.3 23.5 47.6 100.0 0.0099 2.34 2.28 2.03 2.10 8.75 L. piceicola53.8 8.7 17.9 40.2 100.0 0.0129 2.29 2.06 2.25 2.49 9.08 L. piceicola63.1 7.6 25.9 47.6 100.0 0.0138 2.45 3.41 1.84 2.10 9.80 Avg 4.8 11.2 26.1 50.0 100.0 0.0111 2.41 2.45 1.94 2.04 8.84 Min 3.1 7.6 17.9 40.2 100.0 0.0096 1.36 1.65 1.64 1.57 7.98 Max 7.4 18.8 31.0 63.6 100.0 0.0129 3.26 3.41 2.25 2.49 9.80 Stdev 1.5 4.0 5.4 7.7 0.0 0.0016 0.69 0.47 0.28 0.33 0.43 O% J1 J2 J3 J4 F O µm J2/J1 J3/J2 J4/J3 F/J4 Sum L. piceicola161.1 68.0 79.0 89.6 100.0 178.2 1.11 1.16 1.13 1.12 4.52 L. piceicola257.3 65.5 79.2 88.2 100.0 175.6 1.14 1.21 1.11 1.13 4.60 L. piceicola357.5 61.9 74.4 83.1 100.0 160.0 1.08 1.20 1.12 1.20 4.60 L. piceicola461.6 68.8 76.1 88.9 100.0 155.5 1.12 1.11 1.17 1.12 4.52 L. piceicola559.0 66.5 78.3 85.8 100.0 153.9 1.13 1.18 1.10 1.17 4.57 L. piceicola655.9 64.0 75.9 88.0 100.0 155.4 1.14 1.19 1.16 1.14 4.63 Avg 58.7 65.8 77.1 87.3 100.0 163.1 1.12 1.17 1.13 1.15 4.57 Min 55.9 61.9 74.4 83.1 100.0 153.9 1.08 1.11 1.10 1.12 4.52 Max 61.6 68.8 79.2 89.6 100.0 178.2 1.14 1.21 1.17 1.20 4.63 Stdev 2.3 2.6 2.0 2.4 0.0 10.9 0.03 0.04 0.03 0.04 0.04 R% J1 J2 J3 J4 R µm – J2/J1 J3/J2 J4/J3 Sum – L. piceicola166.4 77.4 88.9 100.0 177.6 – 1.17 1.15 1.12 3.44 – L. piceicola266.4 76.0 88.6 100.0 176.7 – 1.14 1.17 1.13 3.44 – L. piceicola362.7 77.2 77.3 100.0 158.0 – 1.23 1.13 1.15 3.51 – L. piceicola467.9 75.6 90.2 100.0 152.7 – 1.11 1.19 1.11 3.42 – L. piceicola565.2 76.8 86.7 100.0 151.5 – 1.18 1.13 1.15 3.46 – L. piceicola660.8 69.9 86.9 100.0 157.3 – 1.15 1.24 1.15 3.54 Avg 64.9 75.5 88.1 100.0 162.3 – 1.16 1.17 1.14 3.47 – Min 60.8 69.9 86.7 100.0 151.5 – 1.11 1.13 1.11 3.42 – Max 67.9 77.4 90.2 100.0 177.6 – 1.23 1.24 1.15 3.54 – Stdev 2.7 2.8 1.4 0.0 11.8 – 0.04 0.03 0.02 0.04 – *Longidorus piceicola, a species with 4 juvenile developmental stages (JDS). Sources: 1Barsi and Lamberti 2011; 2Barsi 2022; 3Lišková et al. 1997; 4,6Groza et al. 2017; 5Kornobis and Peneva 2011. L = body length; V = body volume; O = odontostyle length; R = replacement odontostyle length; F = female. L. Barsi Table 3. Growth patterns of the body length, body volume, odontostyle length and replacement odontostyle length and relative increase in the body length, body volume, odontostyle length and replacement odontostyle length between successive stages in four populations of Longidorus intermedius*. Growing patterns Relative increase L% JI JII JIII F L µm JII/JI JIII/JII F/JIII Sum L. intermedius732.1 44.6 68.2 100.0 4352 1.39 1.53 1.47 4.38 L. intermedius832.7 52.0 70.9 100.0 3759 1.59 1.36 1.41 4.36 L. intermedius931.1 48.6 69.9 100.0 3993 1.56 1.44 1.43 4.43 L. intermedius10 29.4 45.5 68.5 100.0 4220 1.55 1.51 1.46 4.51 Avg 31.3 47.7 69.4 100.0 4081 1.52 1.46 1.44 4.42 Min 29.4 44.6 68.2 100.0 3759 1.39 1.36 1.41 4.36 Max 32.7 52.0 70.9 100.0 4352 1.59 1.53 1.47 4.51 Stdev 1.4 3.4 1.3 0.0 261 0.09 0.07 0.03 0.07 V% JI JII JIII F V mm3JII/JI JIII/JII F/JIII Sum L. intermedius77.0 14.8 37.7 100.0 0.0065 2.11 2.55 2.65 7.31 L. intermedius85.1 20.8 42.4 100.0 0.0062 4.08 2.04 2.36 8.48 L. intermedius94.9 13.9 38.4 100.0 0.0079 2.84 2.76 2.60 8.20 L. intermedius10 5.4 15.4 39.5 100.0 0.0071 2.85 2.56 2.53 7.95 Avg 5.6 16.2 39.5 100.0 0.0069 2.97 2.48 2.54 7.99 Min 4.9 13.9 37.7 100.0 0.0062 2.11 2.04 2.36 7.31 Max 7.0 20.8 42.4 100.0 0.0079 4.08 2.76 2.65 8.48 Stdev 1.0 3.1 2.1 0.0 0.0008 0.81 0.31 0.13 0.50 O% JI JII JIII F O µm JII/JI JIII/JII F/JIII Sum L. intermedius765.4 73.9 86.4 100.0 112.9 1.13 1.17 1.16 3.46 L. intermedius867.6 75.2 86.5 100.0 111.0 1.11 1.15 1.16 3.42 L. intermedius968.2 73.6 84.5 100.0 110.0 10.8 1.18 1.18 3.41 L. intermedius10 66.5 73.4 84.4 100.0 111.2 1.10 1.15 1.18 3.44 Avg 66.9 74.0 85.5 100.0 111.3 1.11 1.15 1.17 3.43 Min 65.4 73.4 84.4 100.0 110.0 1.08 1.15 1.16 3.41 Max 68.2 75.2 86.5 100.0 112.9 1.13 1.17 1.18 3.46 Stdev 1.2 0.8 1.2 0.0 1.2 0.02 0.01 0.02 0.02 R% JI JII JIII R µm – JII/JI JIII/JII Sum – L. intermedius774.4 86.3 100.0 111.2 – 1.16 1.16 2.32 – L. intermedius877.6 89.3 100.0 107.0 – 1.15 1.12 2.27 – L. intermedius974.3 83.5 100.0 109.0 – 1.12 1.20 2.31 – L. intermedius10 72.1 86.1 100.0 112.6 – 1.19 1.16 2.36 – Avg 74.6 86.3 100.0 110.0 – 1.16 1.16 2.32 – Min 72.1 83.5 100.0 107.0 – 1.12 1.12 2.27 – Max 77.6 89.3 100.0 112.6 – 1.19 1.20 2.36 – Stdev 2.3 2.4 0.0 2.5 – 0.03 0.03 0.03 – *Longidorus intermedius, a species with 3 juvenile developmental stages (JDS). Sources: 7Barsi and Lamberti 2004; 8Peneva et al. 2001; 9Kumari et al. 2006; 10Susulovska and Tsaryk 2018. L = body length; V = body volume; O = odontostyle length; R = replacement odontostyle length; F = female. Developmental and post-embryonic growth patterns in Longidorus piceicola and L. intermedius Fig. 3. Comparison of odontostyle length growth patterns in 6 populations of Longidorus piceicola and 4 populations of L. intermedius. L. Barsi specimens in the JI stage generally have a longer odontostyle length compared to J1 stage specimens of L. piceicola. Therefore, they may likely reach maturity more quickly after only three molts. Replacement odontostyle length (Fig. 4A, B) Reminder: in the case of replacement odontostyle length, the absolute value of the replacement odontostyle Fig. 4. Comparison of growth patterns of replacement odontostyle length in 6 populations of Longidorus piceicola and 4 populations of L. intermedius. Developmental and post-embryonic growth patterns in Longidorus piceicola and L. intermedius Table 6. Odontostyle and replacement odontostyle growth patterns in Longidorus intermedius (Obedska bara, Serbia). JI L O R O_R% I% JII L O R O_R% I% 1 1426 68.7 80.0 85.9 14.1 1 1915 78.7 95.0 82.8 17.2 2 1276 68.7 82.5 83.3 16.7 2 2164 78.7 98.1 80.2 19.8 3 1338 71.9 80.6 89.2 10.8 3 2287 80.0 103.7 77.1 22.9 4 1387 72.5 78.7 92.1 7.9 4 1793 82.5 95.0 86.8 13.2 5 1276 72.5 81.9 88.5 11.5 5 2065 82.5 101.2 81.5 18.5 6 1437 72.5 83.7 86.6 13.4 6 1981 82.5 95.0 86.8 13.2 7 1415 73.1 83.7 87.3 12.7 7 1776 82.5 92.5 89.2 10.8 8 1304 73.7 81.2 90.8 9.2 8 2065 82.5 95.0 86.8 13.2 9 1382 75.0 82.5 90.9 9.1 9 2053 83.7 98.7 84.8 15.2 10 1276 75.0 82.5 90.9 9.1 10 1759 83.7 93.7 89.3 10.7 11 1537 75.6 85.0 88.9 11.1 11 1942 83.7 93.7 89.3 10.7 12 1498 77.5 85.0 91.2 8.8 12 1654 83.7 92.5 90.5 9.5 13 1510 78.1 85.0 91.9 8.1 13 1937 83.7 96.2 87 13.0 14 1471 78.7 85.0 92.6 7.4 14 1987 85.0 96.2 88.4 11.6 15 1832 85.0 93.7 90.7 9.3 16 1915 85.0 98.7 86.1 13.9 17 1887 85.6 100 85.6 14.4 18 1831 86.2 92.5 93.2 6.8 19 1942 86.2 91.2 94.5 5.5 20 2037 86.2 97.5 88.4 11.6 JIII L O R O_R% I% 1 3280 91.9 116.2 79.1 20.9 2 2509 93.7 103.7 90.4 9.6 3 2758 93.7 106.2 88.2 11.8 4 2775 93.7 106.2 88.2 11.8 5 2775 93.7 111.2 84.3 15.7 6 3252 93.7 111.2 84.3 15.7 7 2886 95.0 106.2 89.5 10.5 8 3324 95.0 108.7 87.4 12.6 9 2664 95.6 107.5 88.9 11.1 10 3052 96.9 111.2 87.1 12.9 11 2581 96.9 112.5 86.1 13.9 12 3225 97.5 108.7 89.7 10.3 13 3008 97.5 108.7 89.7 10.3 14 2775 97.5 111.9 87.1 12.9 15 2969 97.5 112.5 86.7 13.3 16 3036 97.5 112.5 86.7 13.3 17 2653 98.7 111.2 88.8 11.2 18 2775 98.7 112.5 87.7 12.3 19 2997 99.4 112.5 88.4 11.6 20 2442 99.4 116.2 85.5 14.5 21 3341 100.0 112.5 88.9 11.1 22 3607 100.0 115.0 87.0 13.0 23 3225 100.6 111.2 90.5 9.5 24 3274 100.6 112.5 89.4 10.6 25 2775 100.6 120.6 83.4 16.6 26 2703 101.9 106.2 96.0 4.0 27 3008 102.5 111.2 92.2 7.8 28 3430 103.7 117.5 88.3 11.7 Note. Data sorted by odontostyle length. JI-JIII = juvenile stages; L = body length (µm); O = odontostyle (µm); R = replacement odontostyle (µm); O_R% = odontostyle length in relation to replacement odontostyle length (%); I = increase of replacement odontostyle in relation to odontostyle length (%).