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Article

The Contribution of Singletons and Doubletons Captured Using Weak Light Heath Traps for the Analysis of the Macroheteroceran Assemblages in Forest Biotopes

by
João Matos da Costa
1,2,* and
Marcin Sielezniew
3
1
Narew National Park, 18-204 Kurowo, Poland
2
Doctoral School of Exact and Natural Sciences, University of Bialystok, ul. K. Ciołkowskiego 1K, 15-245 Białystok, Poland
3
Laboratory of Insect Evolutionary Biology and Ecology, Faculty of Biology, University of Bialystok, ul. K. Ciołkowskiego 1J, 15-245 Białystok, Poland
*
Author to whom correspondence should be addressed.
Diversity 2023, 15(4), 508; https://doi.org/10.3390/d15040508
Submission received: 27 February 2023 / Revised: 29 March 2023 / Accepted: 30 March 2023 / Published: 1 April 2023
(This article belongs to the Special Issue State-of-the-Art Biodiversity Research in Poland)

Abstract

:
In nearly every ecological community, most species are represented by a few individuals, and most individuals come from a few of the most common species. Singletons (one individual sampled) and doubletons (two individuals sampled) are very common in moth community studies. In some reports, these specimens are excluded from the analysis once they are considered a consequence of under-sampling or of contamination with tourist species that are just passing through. Throughout 12 nights in 2018 and 12 nights in 2019, two Heath traps, one with an 8 W ultraviolet lamp and the other with a 15 W actinic lamp, were positioned approximately 50 m apart at nine sites of four different biotopes in a mosaic forest ecosystem in the Narew National Park (NE Poland). We were able to differentiate moth assemblages according to the forest biotopes under study and by the year of research. With our results, it becomes more evident that singletons and doubletons sampled using weak light Heath traps should be included in the ecological analysis of Macroheteroceran moth assemblages, and our research strongly suggests that they are an important and consistent element of such a sampling method. We also demonstrate that weak light Heath traps are suitable for building an inventory scheme of moth assemblages in small forest areas and that singletons and doubletons can be crucial elements in long-term monitoring systems.

1. Introduction

Macrolepidopteran moths (Lepidoptera, Macroheterocera) play an important role in the ecosystem: the adults of many species pollinate flowers while the larvae are herbivores and detritivores [1,2]. Both imagoes and prematures are vital food resources for a variety of animals, such as lizards, birds, mammals, and other insects [3,4,5]. Since they are easy to attract, collect, and identify, they have been used in several ecological studies including those providing evidence of the biodiversity crisis. It has been documented worldwide that ecosystems are losing some species and gaining others, resulting in profound impacts on how these ecosystems function. Species gain is derived from colonization and establishment of new species, processes that are increasing in frequency and intensity [6]. The current global environmental change is driving species to extinction at rates hundreds to thousands of times faster than ever recorded. Monitoring studies confirm that global losses in biodiversity are leading to a mass extinction event and evaluating species diversity becomes increasingly critical. [7,8,9,10,11]. Moth fauna is sensitive to climate and land use changes [12,13] as well as to light pollution [14,15,16]. It is reported that local macro-moth assemblages can reflect even a small variation of vegetation in a small study area [17,18,19,20,21,22].
Sampling methods should be simple, time and cost effective and, above all, efficient to collect as much information as it is possible regarding the diversity of the communities under study. Light traps are one of the best methods to survey moths, as they yield a large number of specimens with a minimum effort. It is known that excessive sampling may not improve the quality of the final data set and that the data obtained are restricted to the assemblages sampled with the light types used [23,24,25,26,27,28,29,30]. Heath traps with lamps emitting short wavelengths, such as ultraviolet light (UV), are commonly used. Other light types, with different spectrums show similar results to the UV light. The spectral sensitivity of the eye receptors of most of the Lepidoptera species have peak absorption wavelengths of 350 (UV), 440 (blue) and 525 nm (green) [31,32,33,34,35,36].
Mark recapture tests have shown that the moth recapture rates decrease with increments in the release distance from the light source [37,38,39,40]. The stronger the intensity of light used in the sampling methods, the higher the number of moths attracted is expected to be. Using more powerful lights can lead to a better prospection of communities, once the number of the attracted specimens increases and more species can be registered [29,41,42]. Leinonen et al. (1998) [43] state that low capture rates and short attraction distances are positive aspects for moth monitoring programs, and Grunsven et al. (2014) [20] demonstrated that weak light Heath traps attract moths from a short range and therefore are suitable for restricted local sampling.
In nearly every ecological community, most species are represented by a few individuals and most individuals come from a few of the most common species [44]. Singletons (one individual sampled) and doubletons (two individuals sampled) are very common in moth assemblages studies. The share of singletons and doubletons captured in light moth studies performed in the tropics as well as in the continental climate varies from 18% to 72% [45,46,47,48,49,50,51,52]. Tikoca et al. (2016) [51] in their studies show that the percentage of singletons is 7.3% (high sampling rate) when the sample is hand made with 125 W mercury vapor lamps and up to 45.5% (low sampling rate) when weak light traps of 15 W are used.
Hilt and Fiedler (2005) [46] mention that if rare species are neglected, whether deliberately by study protocol or because of under-sampling, considerable amounts of information will be lost, rendering faunal comparisons questionable. The use of singletons and doubletons in moth assemblages’ analysis, taking into account the above considerations [46], is not uniform. Several studies do not mention whether singletons and doubletons were used during the analysis [17,50,53]. In some reports, singletons and doubletons are excluded from the analysis, since they are considered a consequence of under-sampling or of contamination with tourist species that are just passing through [30,48,54,55,56,57,58,59]. The use of singletons and doubletons is important for the analysis [17,46,51,53,60]; nevertheless, there seems to be no consensus in their use [57,61].
The goals of this study were to understand:
1. Whether is possible to sample and distinguish macro-moth assemblages, in a mosaic forest ecosystem, using weak light Heath traps.
2. Whether singletons and doubletons sampled using weak light Heath traps should be considered as negligible for macro-moth assemblages studies, or whether they should be included in this kind of analysis.

2. Materials and Methods

This study was conducted in the Narew National Park (NNP), North-East Poland, in the Podlaskie Voivodeship. The Park occupies the marshy Narew valley; marshlands and wasteland are the dominating ecosystems and cover about 90% of the Park area. In 2013, forests occupied 10% of the area of the Narew National Park (665 ha) and occurred mainly on swampy habitats (83%). In the NNP, alder is the predominant type, occupying over 84% of the forest area. In addition, there are also pine stands (8.5%) and small areas of birch, aspen, oak, spruce, and maple [62].
A project to conduct an inventory of the Macroheterocera fauna of the NNP was started in 2017 [44]. Until 2019, moths were sampled in 15 sites with different forest biotopes, such as Querco-Pinetum (QP), Ribeso nigri-Alnetum (RN), and in sites that were built up under the high pressure of human activity, designated as Substitute Communities (SC). Such biotopes are not uniformly developed and form very different species combinations [63]. According to the Forest Management Manual 2012 [64], used by the Polish State Forests, SC biotopes have more than 60% of ecological and more than 30% of geographically alien species. In 2018 and 2019, the inventory was conducted in nine sites (QP1, QP2, QP3, RN1, RN2, RN3, RN4, SQP, and SRN). The localization of the sites in the NNP (Figure 1) and the distances between them are presented in Table 1. The QP and SQP sites were dominated by Pinus sylvestris L. and RN and SRN mainly by Alnus glutinosa L. or Betula spp. had the largest share in the forest stand. The studied sites were also somewhat different with respect to the shrub cover as well as the surroundings (Table 2, Appendix A). The site SQP was chosen as the “control site” due to the highest number of species observed in 2017, and macro-moths were collected in 2018 (SQPa) and in 2019 (SQPb).
Between the last and the first quarter of the moon phases, from May to October, throughout 12 nights in 2018 and 12 nights in 2019, two Heath traps, one with an 8 W ultraviolet lamp (Philips TL 8 W BLB) and other with a 15 W actinic lamp (Philips Actinic BL TLD 15 W), were positioned approximately 50 m apart in each site. Both traps were powered by 12 V-14 Ah batteries, positioned at ground level, and operated from dusk till dawn. The actinic lamps used in this study emit wavelengths between 320-400 nm plus a peak at 405 nm and another at 440 nm [65]. The ultraviolet lamps emit similar ultraviolet wavelengths, between 320 and 400 nm, and a small peak at 405 nm [66], Figure 2. The specimens collected from both traps during the entire season were pooled for each site. We did not aim to compare the effectiveness of different lamp types during this study. The collected fauna was euthanized with ethyl acetate inside the traps, packed in plastic bags and frozen. Macrolepidoptera specimens were identified according to their wing pattern [67,68,69,70,71,72] and then stored in the NNP entomological collection.
Table 2. Specification of the sites where the inventory of the Macroheterocera fauna was carried out in the Narew National Park (NNP), Podlaskie Voivodeship, North-East Poland [73].
Table 2. Specification of the sites where the inventory of the Macroheterocera fauna was carried out in the Narew National Park (NNP), Podlaskie Voivodeship, North-East Poland [73].
NameYearArea (ha)Forest BiotopeTree spp. CoverShrubs spp. CoverSurrounded by:
QP120183.01Querco-Pinetum35-year-old P. sylvestris (80%)
35-to 65-year-old Betula spp.
45-year-old Picea abies L.
30-year-old A. glutinosa
Frangula alnus L.
Sorbus aucuparia L.
Juniperus spp.
RN forests
QP220190.4530-year-old P. sylvestris (70%)
30-year-old Betula spp.
25-year-old A. glutinosa
RN forests and
crop fields
QP320197.0755- to 80-year-old P. sylvestris (80%)
40-year-old Betula spp.
RN120181.09Ribeso
nigri-Alnetum
30- to 50-year-old A. glutinosa (70%)
50-year-old Betula spp.
Salix spp.
F. alnusP. sylvestris and
Betula spp. forests
RN220180.8325-year-old Betula spp. (60%)
25-year-old A. glutinosa
Salix spp.
F. alnus
RN320182.0430- to 65-year-old A. glutinosa (90%)
45-year-old Betula spp.
Salix spp.
F. alnus
Padus avium L.
RN forests and small patches of P. sylvestris
RN420182.7425- to 55-year-old A. glutinosa (90%)
50-year-old Salix spp.
F. alnus
P. avium
SQPa20180.61Substitute Community of
Querco-Pinetum
33- to 45-year-old P. sylvestris (80%)
45-year-old Betula spp.
Salix spp.
F. alnus
S. aucuparia
P. avium
RN forests and
crop fields
SQPb2019
SRN20190.31Substitute Community of
Ribeso nigri-Alnetum
33-year-old A. glutinosa (80%)
55- to 70-year-old Betula spp.
Salix spp.
P. sylvestris forests and marshlands
The species were segregated according to their larval food necessities: to their host plant specificity: (i) m1-1st degree monophagous (on one plant species), m2-2nd degree monophagous (on one plant genus), o1-1st degree oligophagous (on one plant family), o2-2nd degree oligophagous (on two to four plant family), po-polyphagous; (ii) to the food plant type: He-herbaceous, Sc-shrubs and Tr-trees, Ot-lichens, mosses or decaying organic material. Additionally, species related with Con-conifers were distinguished [67,68,69,70,71,72].

Statistical Analysis

Analysis of variance (ANOVA) was performed to test whether there were statistical differences between the distribution of the number of species and individuals per families among sites with a 0.05 significant level of confidence. The Pearson’s correlation coefficient was used to measure the strength and direction of the relationship between two variables. A Mantel test was run to evaluate the relationship between two matrices: (i) the distances (kilometers) among the studied sites and (ii) the values of the Morisita similarity index among the sites (based on the number of species according to their larval food necessities). Based on the species larval food necessities, Cluster Analysis and non-metric multidimensional scaling (NMDS) were performed. Cluster Analysis was executed using the abundance-based Morisita index (CLM) values of distribution of the number of species among sites, based on their larval food necessities. Each pair of sites was evaluated considering the degree of similarity, then combined sequentially into clusters to form a dendrogram with the branching point, representing the measure of similarity. The Morisita index is not influenced by species richness or sample size but it is sensitive to the abundance of the dominant species. This index is likely to be resistant to under-sampling because the influential abundant species are always present in the samples. The NMDS were rendered using the Bray-Curtis similarity index values [74] of distribution of the number of species among sites, based on their larval food necessities. The Bray-Curtis index is particularly useful for assemblages containing a large number of rare species. These statistical analyses were performed using the free software Past, version 4.11 [75].
The Shannon index (H’) and the Fisher’s alpha (F’s) index were used to evaluate the Macroheteroceran biodiversity. The Evenness index was used to estimate how equally abundant the species were in each site [76]. The Chao1 estimator was used to calculate the estimated true species diversity of each site [77,78].

3. Results

A total of 11,502 Macroheterocera individuals belonging to 10 families and 284 species, Appendix B, were collected during 2018 and 2019. Most of the individuals (90.5%) and species (84.9%) belonged to three families: Erebidae (2716 individuals, 39 species), Geometridae (5858 individuals, 105 species), and Noctuidae (1833 individuals, 97 species). The sites SQPb, SRN, and QP3 were the ones where the highest number of species was recorded. The site RN4 was the one with the highest number of individuals recorded followed by the sites SRN and SQPb. The total number of individuals and species per family in each site is presented in Table 3. No differences were found in the distribution of the total number of individuals (F = 0.528, df = 9, p = 0.85) nor in the total number of species (F = 0.156, df = 9, p = 0.997) per family among sites (ANOVA test). The Mantel test did not detect a significant relationship between the distances among sites and their Morisita values of similarity (R = −0.136, p = 0.636) (Table 1 and Figure 3). A high correlation between the total number of individuals and the number of species per family was observed (R = 0.875, p < 0.001).
According to the host plant specificity, only in the o1 group, the abundance clearly represented a higher portion (18%), the double of its species richness (9%) (Figure 4). In the other groups, the distribution of individuals and species was equally represented. The collected fauna was mainly composed of polyphagous and 2nd degree oligophagous species, both in abundance and richness.
Considering the plant type in terms of abundance and species richness, all assemblages were characterized by species whose larvae fed mainly on herbaceous plants and then on trees and shrubs. The highest abundance of Con was documented in the sites that were mainly covered by P. sylvestris, although moth richness, except for the sites RN3 and RN4, do not strongly vary among sites. There were no differences in the distribution of the total number of species (F = 0.289, df = 9, p = 0.974) and individuals (F = 0.9349, df = 9, p = 0.506) per family among sites according to their host plant necessities (ANOVA).
The abundance of the Erebidae family weighted more in the RN sites than in the SC and QP sites, and the share of the Geometridae and Noctuidae families was lower. There were no huge differences in the abundance among families between the SC and QP sites. Their abundance was clearly lower in the RN sites than in the SC and QP sites, although the difference in the species was almost none. The weight of Sc species was lower in the SC sites but not very different from RN and QP sites; however, the share of individuals dependent of the Ot species was equal among study sites. The abundance of p species was clearly lower in the RN site than in the SC and QP sites. The share of o2 species was more significant in the RN sites than in all the others. No significant differences were found in the weight of the m1, m2 or o1 among the RN, SC, and QP sites (Figure 5).
Both NMSD and CLM grouped the moth assemblages according to forest biotopes by RN, QP, and SC; they clearly segregated the RN from QP and SC and brought together the latter two. According to the CLM and NMSD, the sites QP2, QP3, SQPb, and SRN studied in 2019 were separated from those that were studied in 2018 (Figure 6A and Figure 7A).
The total number of species that were reported as singletons or doubletons (SD) at least in one site was 233 (82%), although their abundance in all the sites was 5622 individuals, representing 49% of the collected specimens. The number of SD species, per site and per family, is stated in Table 4. The highest number of SD species was reported at the sites QP3, SQPb, and RN. From these 233 species, 45 were considered unique: once, only one individual was collected. No differences in the distribution of SD species number among sites (F = 0.201, df = 9, p = 0.99) were disclosed (ANOVA test). The distribution of SD species according to their larval plant necessities was highly correlated with the total fauna (R = 0.997, p = 0.0002). Most of the SD species, 67%, were 2nd degree oligophagous and polyphagous (Figure 8). There was a strong correlation between the total number of species and the number of SD species collected per month in each site (Figure 9).
Another analysis was performed. The SD species were eliminated from the total number of species present in each site, and CLM was performed again using the same methods and similarity measures as well as the NMSD (Figure 6B and Figure 7B). The CLM and the NMSD show similar results, but they do not clearly join the sites according to the habitats existing in each site. The NMSD continued to join the sites SQPb–SRN, both with SC biotopes, the QP2 and QP3, both with QP biotopes, although all the sites are equally distributed. The CLM continued to reveal a connection between the sites QP2–QP3, both with QP biotopes as well as the sites RN3 and RN4 both with RN biotopes, while the other habitats were completely mixed up. The CLM joined the sites SQPa and SQPb but it was not able to differentiate the assemblages recorded in 2018 and 2019. The areas RN1 and RN2, with both system of analysis, with or without the use of SD species, were separated from the rest of the sites.
As a complementary analysis, the SD species were submitted to CLM and NMSD (Figure 6C and Figure 7C, respectively), and both systems of analysis, except for site RN1, clearly separated the sites according to their moth assemblages’ similarities and hence the QP and SC biotopes from the RN.
Apart from the site RN2, the Shannon index (H’), when all the species were evaluated, indicated that the sites with the SC habitats had the highest Macroheterocera biodiversity, followed by the sites with QP and RN. The Fisher’s alpha (F’s) index disclosed similar results. When the SD species were removed from the analysis, the H’ and the F’s indexes values decreased in all sites, although the E’ index values increased. The Chao1 index had the higher values in the sites RN2, RN3, RN4, and SQPa, showing that these were the most under sampled sites (Table 5).

4. Discussion

In temperate forests, several authors have reported that the families Erebidae, Geometridae, and Noctuidae are the ones that normally dominate the moth assemblages [53,79,80,81,82], which is totally in agreement with our results. We found no differences in the distribution of the number of species and individuals between sites and a strong correlation between the number of species and individuals, similar to that reported previously [23,27,53,83,84]. This relationship between moth species richness and abundance is probably a marker of how undisturbed and undegraded these forest complexes are [85,86]. The similarity of species richness and abundance between sites, as Tikoca et al. (2017) [53] suggest, may be a consequence of two circumstances: first, these forest complexes are geographically close to each other, and second, all of them, in recent years, have suffered low disturbance burdens. In agreement with this statement, we found no significant differences in the distribution of the number of species between sites either according to their host plant specificity, or to the plant type in which their larvae inhabit. These results reveal a solid and balanced moth species distribution in all the forest biotopes under study.
Choi (2007) [87] states that disturbed habitats are preferred by polyphagous species feeding on herbaceous and weedy plants, while undisturbed sites are favored by species that feed on woody plants and trees. We found that the abundance of polyphagous species in undisturbed sites, RN forests, is 33% and in QP forests is 40%. In SC forests with build-up under the pressure of human activity, the corresponding abundance was 43% (Figure 2C). Hence, our results support the remarks of Choi (2007) [87] and that QP forests, even if this is not documented, are most probably built up under some influence of human activity. Considering that the o2 and p species are mostly generalists, our results, Figure 4, are similar to those obtained by Summerville et al. (2013) [57], where more than 75% of the sampled species were generalists.
In our study sites, the abundance and species richness of the groups He and Sc, understory species, are clearly higher than the abundance and species richness of the moths belonging to the Tr canopy group, as Hirao et al. (2009) and Horwáth et al. (2016) [27,47] reported. These results corroborate the assumption that the forest understory is as important as the forest overstory for Macroheteroceran biodiversity [83] and the possibility of species richness of the vegetation below the forest canopy is responsible for most of the variation in moth assemblages [45,47].
Usher and Keiller (1998) [88] do not distinguish differences in moth assemblages in coniferous, deciduous, and mixed forest areas. On the other hand, some authors based on moth assemblages such as Summerville and Crist (2003) [83] distinguish forest stands according to the season, region, and size of the stand; Greco et al. (2018) [60] was able to group stands according to the time elapsed from the last human intervention. With similar methods, based on the Macroheteroceran assemblages, we were able to segregate nine forest sites according to their forest biotopes, to the year of study and to their level of degeneration (Figure 6A and Figure 7A). Both CLM and NMSD assembled most of the sites according to their biotopes and by year of study. In Figure 5, it is possible to observe that the QP and SC forest biotopes had very similar moth assemblages, which explains why both CLM and NMSD were not able to clearly separate sites to the corresponding biotopes. According to the Forest Management Manual (2012) [64], used by the Polish State Forests, the SC biotopes have more than 60% of ecological and more than 30% of geographically alien species. We can therefore conclude that the Anthropocene pressure may be observed in forest habitats by the evaluation of moth assemblages because the SRN site was never joined with its most similar forest biotope, the RN. The moth fauna of the SRN site was clearly more similar to the QP and SQP biotopes. Both CLM and NMSD joined sites SQPa and QP, and CLM joined sites SQPb-QP2 and QP3-SRN, while the NMSD only joined the SRN-SQPB sites, which were most probably set ensembled since the QP and SC sites had similar moth assemblages. Applying this analysis, the other sites were grouped according to their biotopes.
From a totally different angle, these results confirm that weak light Heath traps, as they attract moths from a shorter range, provide a very local sampling that enabled to understand the differences in moth populations in a highly mosaic forest ecosystem [20]. This can also be confirmed and supported by the fact that the distance between the sites under research did not influence the similarity of moth assemblages between areas. The connection of the sites QP2 and QP3 and SQP was not driven by their proximity but due to the similarities that these biotopes induced in the moth assemblages associated with them.
The number of singletons and doubletons, collected in our study, does not diverge from data recorded in other studies [30,50,51,52,53,54,55,89]. In our study sites, most of the SD species were generalists (67%), probably because they were either colonists, which were attracted and caught during dispersal flights, or low-density specialist species (of groups m1, m2, and o1) that occupy special niches of the forest [17,50,87]. Generalist species, as well as specialist ones, can occupy special niches, where they find small patches of their host plants, and so, in some sites, their populations have low densities. These hypotheses may explain why there were no significant differences in the distribution of the number SD species between sites. The distribution of the total number of species per family per site, due to its similarity, demonstrates that, in this area of study, the number of SD species was not dependent of the biotopes. Considering that RN forests in the NNP suffer no human disturbance, while QP most probably did, and that SC was created under the pressure of human activity, the distribution of SD species did not depend of the disturbance. Therefore, it is possible to conclude that SD species were not necessarily rare (low abundance); they may have had low abundance in one specific site but not in the overall study area [90]. The 45 unique species that were collected should not be considered as rare (low abundance) in the total forest ecosystem under study, since this may be a consequence of incomplete sampling or of the distinct ecological characteristics of species, such as season and small habitat ranges, which led to small population expressiveness [48,53,59,91,92]. As far as our understanding goes, to overcome the above-mentioned study limitations, increasing the temporal extent of the inventory will be necessary to comprehend the peculiarities of the size of the small populations [47].
The analysis of the SD species, on their own, revealed how important they were for the moth assemblages that they represented, on average 43% of them, once they were able to segregate the study sites based on their moth assemblage similarities. The RN1 site, as a result of CLM and NMSD analysis, was near to the QP and SC moth assemblages, strongly suggesting that it was the one with the higher level of degeneration and on the other hand that the site RN4 was the less degenerated one.
In several studies, the SD species were not taken into consideration [48,56,57,93]; however, it has been suggested that if they are not included in the analysis, then the significance of ecological comparisons will be questionable [46,59]. In our studies, when the SD species were excluded from the analysis, it was not possible to distinguish sites according to the year of study, showing that without their use, moth assemblages became uniform [53] and had no seasonal variation, hence compromising other analysis and monitoring systems (Figure 7 and Figure 9). These results point out the relevance of including SD species sampled using weak light Heath traps in the studies of moth population diversity in forest sites. Our results are strongly supported by these statements: (i) these systems of analysis joined the QP2 and QP3 sites, both within the QP biotope; (ii) joined the RN1-RN2 and RN3-RN4 sites, all within the RN biotope; (iii) both systems joined the QP and SC sites, due to their similar moth assemblages; (iv) both systems of analysis were able to distinguish the sites according to the year of study; and (v) by themselves, they could clear segregate the study sites according to the similarities of their Macroheteroceran moth assemblages, indicating that SD species sampled using weak light Heath traps were very important and they should be included in ecological analysis. Further analysis of all the above-presented data opened up a different approach to how such a high number of SD species appeared as well as to their important contribution to the analysis of Macroheteroceran assemblages in these forest biotopes. The number of SD species did not depend of the disturbance degree of either of the different forest biotopes. There were no differences in the distribution of the number of SD species among sites. With the use of SD species in the analysis, it was possible to distinguish sites according to the year of study, and without their use, the moth assemblages were uniform and had no seasonal variation. With the use of SD species, we were able to segregate nine forest sites according to their biotopes and to their level of degeneration based on the Macroheteroceran assemblages. Tikoca et al. (2016) [51] report that weak light traps have up to 38% more SD species than strong light methods. If we take the above statements into consideration, we can point out that, although further studies are required, the high number of SD species was an effect of the low sample rates of the weak light Heath traps. Following this hypothesis, the SD species should not be considered as colonists, low-density specialists, or generalist species at the start of the analysis, but rather as an important and consistent (Figure 7) element of the samples that these traps are capable of obtaining when used. Moreover, SD species can be crucial to the understanding of the seasonal variations of the moth assemblages, since they contain much of the faunal information, as Hilt and Fiedler (2005) [46] suggest, and so are clearly important for long-term monitoring systems.
Slight differences, although clearly important, in the flora of the sites RN1 and RN2 separated them from all the other sites with or without the use of SD. They are separated even from their most similar ones, the RN3 and RN4, since no Cossidae or Hepialidae species were recorded and the number of Noctuidae species was lower than that in all the other sites.
Both biodiversity indices, S’ and Fs’, presented clear differences when SD species were used, decreasing their value when they were not included, as expected. The E’ index increased its values in all areas when SD species were absent from the analysis, once they became more uniform. The Chao 1 estimated that all the areas were under sampled [77], although if we take into consideration the above statements, that SD species were a result of the weak light Heath traps, these values cease to be trustworthy (Table 5).

5. Conclusions

Our results demonstrate that weak light Heath traps are suitable for building up an inventory scheme of moth assemblages in small mosaic forest areas. We were able to distinguish the variation of the Macroheteroceran moth assemblages inhabiting different forest biotopes in a mosaic landscape as well as according to the year of study. Our data also indicate that singletons and doubletons, sampled using weak light Heath traps, should be included in the ecological analysis of Macroheteroceran moth assemblages.

Author Contributions

J.M.d.C. designed the studies and conducted the field work and analyses under the supervision of M.S.; J.M.d.C. wrote the manuscript with the input of M.S.; J.M.d.C. acquired funds for studies. All authors have read and agreed to the published version of the manuscript.

Funding

This research was supported by Polish State Forests’ by the “Forest Fund”, grant number EZ.0290.1.17.2017.

Institutional Review Board Statement

Not applicable.

Data Availability Statement

Not applicable.

Acknowledgments

I am very grateful to the Direction of the NNP for allowing me to join this adventure. Three anonymous reviewers made valuable comments on the manuscript. I truly appreciate the help of Natercia Caneira throughout the revisions of this manuscript. Sarah Luczaj made linguistic improvements on the manuscript.

Conflicts of Interest

The authors declare no conflict of interests.

Appendix A

Figure A1. Representative pictures of the studied forest biotopes. A—QP biotope, B—SQP biotope, C—RN biotope and D—SRN biotope.
Figure A1. Representative pictures of the studied forest biotopes. A—QP biotope, B—SQP biotope, C—RN biotope and D—SRN biotope.
Diversity 15 00508 g0a1aDiversity 15 00508 g0a1b

Appendix B

Table A1. List of species and individuals recorded in each site (see text for details).
Table A1. List of species and individuals recorded in each site (see text for details).
Family CossidaeLarval Food NecessitiesQP1QP2QP3RN1RN2RN3RN4SC1aSc1bSc2
Phragmataecia castaneae (Hübner, 1790)m1, He135 48 2
Family Drepanidae
Drepana curvatula (Borkhausen, 1790)o2, Tr8111281111101821
Drepana falcataria (Linnaeus, 1758)o2, Tr9511242614148
Falcaria lacertinaria (Linnaeus, 1758)o2, Tr1 1 3 2
Habrosyne pyritoides (Hufnagel, 1766)m2, Sc 2 42124
Ochropacha duplaris (Linnaeus, 1761)o2, Tr31137194341
Tethea ocularis (Linnaeus, 1767)O1, Tr2 1
Thyatira batis (Linnaeus, 1758)o1, Sc 231 3 64
Watsonalla binaria (Hufnagel, 1767)O1, He, Tr 111
Family Erebidae
Arctia caja (Linnaeus, 1758)p, He, Sc, Tr1463618251347
Atolmis rubricollis (Linnaeus, 1758)p, Ot 22 73
Callimorpha dominula (Linnaeus, 1758)p, He, Sc64 92116544
Calliteara pudibunda (Linnaeus, 1758)o2, Sc, Tr411161110 1031
Catocala electa (Vieweg, 1790)o2, Sc, Tr 1 1 13
Catocala fraxini (Linnaeus, 1758)o2, Sc, Tr 2 64
Catocala nupta (Linnaeus, 1767)O1, Tr 1
Collita griseola (Hübner, 1803)O1, He89123611154716913
Colobochyla salicalis (Denis & Schiffermüller, 1775)O1, Tr 12 1 6324
Coscinia cribraria (Linnaeus, 1758)p, He 121 1
Cybosia mesomella (Linnaeus, 1758)p, Ot 41 2 2
Diacrisia sannio (Linnaeus, 1758)o2, He 1 1 124
Diaphora mendica (Clerck, 1759)p, He, Sc 1
Eublemma minutata (Fabricius, 1794)m1, He 131 3
Euproctis similis (Fuessly, 1775)p, Sc, Tr24512914822838
Herminia grisealis (Denis & Schiffermüller, 1775)p, He, Ot1311 1 39
Herminia tarsipennalis (Treitschke, 1835)p, Ot 124 6733
Hypena proboscidalis (Linnaeus, 1758)p, He, Sc713121471758222390
Hypena rostralis (Linnaeus, 1758)p, Ot 1
Katha depressa (Esper, 1787)p, Ot1121526 54191110
Laspeyria flexula (Denis & Schiffermüller, 1775)p, Ot42 3212563
Lithosia quadra (Linnaeus, 1758)m2, Sc, Tr 114 1 43
Lygephila pastinum (Treitschke, 1826)O1, He 2
Lymantria dispar (Linnaeus, 1758)o2, Sc, Tr3 1026 85116
Family ErebidaeLarval food necessitiesQP1QP2QP3RN1RN2RN3RN4SC1aSc1bSc2
Lymantria monacha (Linnaeus, 1758)p, Con, Tr1442136 517615
Eilema lurideola (Zincken, 1817)o2, He 13 52
Eilema complana (Linnaeus, 1758)p, He 6163 562
Eilema lutarella (Linnaeus, 1758)p, He 11 1
Miltochrista miniata (Forster, 1771)m1, He1166814210329
Orgyia antiquoides (Hübner, 1822)o2, Sc 1
Orgyia antiqua (Linnaeus, 1758)p, Con, Sc, Tr 11 1
Pelosia muscerda (Hufnagel, 1766)m1, He1315586017193913 9
Phragmatobia fuliginosa (Linnaeus, 1758)p, He, Sc53715 675103
Rivula sericealis (Scopoli, 1763)o2, He17361862570193375
Scoliopteryx libatrix (Linnaeus, 1758)o1, Sc1 21 1
Spilarctia lutea (Hufnagel, 1766)p, He, Sc1623149252049816
Spilosoma lubricipeda (Linnaeus, 1758)o2, He, Sc2391117128111815834
Spilosoma urticae (Esper, 1789)p, He 2 16
Wittia sororcula (Hufnagel, 1766)p, Ot 22 4
Family Geometridae
Abraxas grossulariata (Linnaeus, 1758)o2, Sc, Tr 1115 39
Abraxas sylvata (Scopoli, 1763)p, Sc, Tr15122524181141227
Aethalura punctulata (Denis & Schiffermüller, 1775)O1, Tr 57122 72825
Agriopis aurantiaria (Hübner, 1799)o2, Tr1
Alcis repandata (Linnaeus, 1758)o2, Sc, Tr1 5 27 1
Angerona prunaria (Linnaeus, 1758)p, Sc, Tr 1032 2 4819
Apeira syringaria (Linnaeus, 1758)o2, Sc, Tr 1
Biston betularia (Linnaeus, 1758)p, He, Sc, Tr1329316233
Bupalus piniaria (Linnaeus, 1758)p, Con, Tr9180163 7143
Cabera exanthemata (Scopoli, 1763)o2, Sc, Tr51624991232111211
Cabera pusaria (Linnaeus, 1758)o2, Sc, Tr 18221 13716
Campaea margaritaria (Linnaeus, 1761)o2, Sc, Tr 951 4211415
Camptogramma bilineata (Linnaeus, 1758)o2, He64 11 43614
Catarhoe cuculata (Hufnagel, 1767)m2, He1
Cepphis advenaria (Hübner, 1790)p, Sc, Tr2 1 1 44
Charissa ambiguata (Duponchel, 1830)p, He, Sc 1
Chiasmia clathrata (Linnaeus, 1758)O1, He2 13 1 45
Chloroclysta siterata (Hufnagel, 1767)o2, Tr22 1
Cleora cinctaria (Denis & Schiffermüller, 1775)p, He, Sc, Tr 1
Colostygia pectinataria (Knoch, 1781)o2, He31032 10621427
Colotois pennaria (Linnaeus, 1761)o2, Sc, Tr7 113228741
Cosmorhoe ocellata (Linnaeus, 1758)O1, He 3 111
Family GeometridaeLarval food necessitiesQP1QP2QP3RN1RN2RN3RN4SC1aSc1bSc2
Crocallis elinguaria (Linnaeus, 1758)o2, Sc, Tr 1
Cyclophora linearia (Hübner, 1799)o2, Sc, Tr1
Cyclophora punctaria (Linnaeus, 1758)o2, Tr26614 3 77
Cyclophora albipunctata (Hufnagel, 1767)O1, He, Tr4642121 253
Cyclophora annularia (Fabricius, 1775)o2, Tr2 11 121
Cyclophora pendularia (Clerck, 1759)o2, Sc, Tr39521 3152
Deileptenia ribeata (Clerck, 1759)p, Con, Sc, Tr1 35
Dysstroma truncata (Hufnagel, 1767)p, He, Sc, Tr222137 41630
Ecliptopera capitata (Herrich-Schäffer, 1839)m1, He3748 9178148
Ecliptopera silaceata (Denis & Schiffermüller, 1775)o2, He49119 91632318
Ectropis crepuscularia (Denis & Schiffermüller, 1775)p, He, Sc, Tr245 2 4109
Electrophaes corylata (Thunberg, 1792)o2, Sc, Tr 73 13
Ematurga atomaria (Linnaeus, 1758)o2, Sc, Tr 1 1
Ennomos alniaria (Linnaeus, 1758)o2, Sc, Tr 1
Ennomos autumnaria (Werneburg, 1859)o2, Sc, Tr 111 1 72
Epione repandaria (Hufnagel, 1767)p, Sc, Tr1 1 237 1
Epirrhoe alternata (Müller, 1764)m2, He1215116533351868
Epirrhoe rivata (Hübner, 1813)m2, He2 743153
Epirrhoe tristata (Linnaeus, 1758)m2, He 11 1 12
Epirrita autumnata (Borkhausen, 1794)m2, He, Sc, Tr31 12167446361
Epirrita dilutata (Denis & Schiffermüller, 1775)p, Tr2 1 1072
Euchoeca nebulata (Scopoli, 1763)o2, Tr1142551922320176379876
Eulithis mellinata (Fabricius, 1787)m2, Sc 1112545115
Eulithis prunata (Linnaeus, 1758)m2, He 12 222
Eulithis testata (Linnaeus, 1761)o2, He, Sc, Tr 1214 12
Euphyia unangulata (Haworth, 1809)m2, He61481711141691520
Eupithecia exiguata (Hübner, 1813)o2, Sc, Tr 3 21
Eupithecia virgaureata (Doubleday, 1861)o2, He165561232511728
Eupithecia vulgata (Haworth, 1809)p, Ot7 438 5
Eustroma reticulata (Denis & Schiffermüller, 1775)m1, He 41 5135
Geometra papilionaria (Linnaeus, 1758)p, Sc, Tr 2334 4 46
Hydrelia flammeolaria (Hufnagel, 1767)O1, He 27266271751076
Hydriomena furcata (Thunberg, 1784)o2, Sc, Tr 13
Hydriomena impluviata (Denis & Schiffermüller, 1775)p, Tr 3614 10021
ylaea fasciaria (Linnaeus, 1758)o1, Con, TR 2
Hypomecis punctinalis (Scopoli, 1763)p, Con, Sc, Tr12 342 1154
Hypomecis roboraria (Denis & Schiffermüller, 1775)p, Tr811352158642352223
Idaea aversata (Linnaeus, 1758)p, He, Sc, Tr51454 25715
Idaea biselata (Hufnagel, 1767)p, He 1 41
Family GeometridaeLarval food necessitiesQP1QP2QP3RN1RN2RN3RN4SC1aSc1bSc2
Idaea straminata (Borkhausen, 1794)p, He 1 4
Ligdia adustata (Denis & Schiffermüller, 1775)m2, He1 4 232113
Lomaspilis marginata (Linnaeus, 1758)o2, Sc, Tr 354139469
Lomographa bimaculata (Fabricius, 1775)p, Sc, Tr 113
Lomographa temerata (Denis & Schiffermüller, 1775)p, Sc, Tr111 1 211
Lycia hirtaria (Clerck, 1759)o2, Tr 68 1510
Macaria alternata (Denis & Schiffermüller, 1775)o2, Sc, Tr24372 6433
Macaria artesiaria (Denis & Schiffermüller, 1775)m2, Sc, Tr 1 3
Macaria brunneata (Thunberg, 1784)o2, Sc, Tr 16 24
Macaria liturata (Clerck, 1759)o2, Con, Sc, Tr1989511 272114
Macaria notata (Linnaeus, 1758)o2, Tr281252 11725
Macaria wauaria (Linnaeus, 1758)m2, Sc1 17123117
Mesoleuca albicillata (Linnaeus, 1758)m2, Sc1111123236
Minoa murinata (Scopoli, 1763)m1, He1
Opisthograptis luteolata (Linnaeus, 1758)o2, Sc, Tr 1
Orthonama vittata (Borkhausen, 1794)m2, He 176436402513
Paradarisa consonaria (Hübner, 1799)o2, Sc, Tr 1
Pelurga comitata (Linnaeus, 1758)O1, He 13 11
Pennithera firmata (Hübner, 1822)p, Sc6 23731 20162
Peribatodes rhomboidaria (Denis & Schiffermüller, 1775)p, He, Sc, Tr 1381 522949
Perizoma alchemillata (Linnaeus, 1758)O1, He233131 3
Perizoma bifaciata (Haworth, 1809)O1, He 1
Petrophora chlorosata (Scopoli, 1763)m1, O 949957117
Philereme transversata (Hufnagel, 1767)m2, He 1
Philereme vetulata (Denis & Schiffermüller, 1775)m2, He 1 23113 2
Plagodis dolabraria (Linnaeus, 1767)p, Tr 1
Pterapherapteryx sexalata (Retzius, 1783)p, Sc, Tr 1 3
Scopula (Ustocidalia) ternata Schrank, 1802p, He1 1 1
Scopula marginepunctata (Goeze, 1781)p, He 21
Scopula ternata (Schrank, 1802)o1, Sc1
Scopula immorata (Linnaeus, 1758)p, He 1 1
Scopula rubiginata (Hufnagel, 1767)p, He 1
Scotopteryx chenopodiata (Linnaeus, 1758)p, Sc3 12 1 1
Selenia dentaria (Fabricius, 1775)p, Tr 11 1 1
elenia lunularia (Hübner, 1788)p, Tr115 2 244
Selenia tetralunaria (Hufnagel, 1767)p, Tr 1 15
Siona lineata (Scopoli, 1763)o2, He 1 1 1
Thera juniperata (Linnaeus, 1758)m2, Sc 1 2
Timandra comae (Schmidt, 1931)o2, He32115136851294917
Family GeometridaeLarval food necessitiesQP1QP2QP3RN1RN2RN3RN4SC1aSc1bSc2
Xanthorhoe biriviata (Borkhausen, 1794)m1, He 13 807 34
Xanthorhoe designata (Hufnagel, 1767)p, He23 6 6202 6
Xanthorhoe ferrugata (Clerck, 1759)p, He35 13 162910831327
Xanthorhoe quadrifasiata (Clerck, 1759)p, He 1 12
Xanthorhoe spadicearia (Denis & Schiffermüller, 1775)p, He8 2 785413
Family Hepialidae
Triodia sylvina (Linnaeus, 1761)p, He, Ot5 3 73187
Family Lasiocampidae
Dendrolimus pini (Linnaeus, 1758)o1, Con, TR3892124 1416
Euthrix potatoria (Linnaeus, 1758)p, He63543105466187
Gastropacha quercifolia (Linnaeus, 1758)o2, Sc, Tr 1 1
Macrothylacia rubi (Linnaeus, 1758)p, He, Sc, Tr 1 11
Poecilocampa populi (Linnaeus, 1758)o2, Tr1 1011
Family Noctuidae
Abrostola tripartita (Hufnagel, 1766)m1, He1 2 1
Abrostola triplasia (Linnaeus, 1758)m1, He 1 5
Acontia trabealis (Scopoli, 1763)m1, He 1
Acronicta leporina (Linnaeus, 1758)p, Sc, Tr11 4211
Acronicta strigosa (Denis & Schiffermüller, 1775)o1, Sc, Tr 1 11741
Acronicta megacephala (Denis & Schiffermüller, 1775)o1, Sc, Tr 1 22113
Acronicta cuspis (Hübner, 1813)O1, Tr 11 11 1
Acronicta rumicis (Linnaeus, 1758)p, He2121295105541112
Agrochola litura (Linnaeus, 1758)p, He, Sc, Tr1
Agrochola lota (Clerck, 1759)p, He, Sc, Tr2 2 7511
Agrochola circellaris (Hufnagel, 1766)p, He, Sc, Tr 2 3
Agrotis cinerea (Denis & Schiffermüller, 1775)o2, He 1
Agrotis exclamationis (Linnaeus, 1758)p, He131 213109
Agrotis segetum (Denis & Schiffermüller, 1775)p, He1 21
Agrotis vestigialis (Hufnagel, 1766)p, He 5
Allophyes oxyacanthae (Linnaeus, 1758)O1, Tr3 13 5551010
Amphipoea lucens (Freyer, 1845)p, He 11 1
Amphipoea oculea (Linnaeus, 1761)O1, He1 2 54
Amphipyra berbera (Rungs, 1949)p, Sc, Tr 11
Amphipyra livida (Denis & Schiffermüller, 1775)o2, He 2
Anaplectoides prasina (Denis & Schiffermüller, 1775)p, He, Sc 1210 1812
Apamea sordens (Hufnagel, 1766)O1, He 1
Arenostola phragmitidis (Hübner, 1803)m1, He 1 111
Asteroscopus sphinx (Hufnagel, 1766)p, Tr 1 2 4
Axylia putris (Linnaeus, 1761)o2, He2 4
Family NoctuidaeLarval food necessitiesQP1QP2QP3RN1RN2RN3RN4SC1aSc1bSc2
Cerapteryx graminis (Linnaeus, 1758)O1, He1 22113
Charanyca ferruginea (Esper, 1785)p, Sc, Tr 5
Conistra rubiginea (Denis & Schiffermüller, 1775)p, He, Sc, Tr 1 1
Cosmia trapezina (Linnaeus, 1758)o2, Sc, Tr 2
Craniophora ligustri (Denis & Schiffermüller, 1775)o1, Sc, Tr 2
Cucullia artemisiae (Hufnagel, 1766)O1, He1
Deltote bankiana (Fabricius, 1775)p, He 2 2
Deltote deceptoria (Scopoli, 1763)p, He2 1 26
Deltote uncula (Clerck, 1759)o2, He 1 3
Denticucullus pygmina (Haworth, 1809)o2, He12139848341
Diachrysia chrysitis (Linnaeus, 1758)p, He 22 24
Diarsia brunnea (Denis & Schiffermüller, 1775)o2, Sc, Tr1 1
Diarsia rubi (Vieweg, 1790)p, He, Sc3151 12111
Diloba caeruleocephala (Linnaeus, 1758)o1, Sc1 1742
Dypterygia scabriuscula (Linnaeus, 1758)o2, He3212412555
Egira conspicillaris (Linnaeus, 1758)p, He, Sc 1
Eucarta virgo (Treitschke, 1835)o2, He 11 1
Euplexia lucipara (Linnaeus, 1758)p, He, Sc 2 1 212
Eupsilia transversa (Hufnagel, 1766)p, He, Sc 1
Gortyna flavago (Denis & Schiffermüller, 1775)o2, He 1 11 6
Hada plebeja (Linnaeus, 1761)p, He122 12573
Helotropha leucostigma (Hübner, 1808)O1, He 4 31
Hoplodrina blanda (Denis & Schiffermüller, 1775)o2, He 11 23
Hydraecia micacea (Esper, 1789)o2, He 961 341325
Ipimorpha subtusa (Denis & Schiffermüller, 1775)o1, Sc, Tr 1
Lacanobia contigua (Denis & Schiffermüller, 1775)p, He, Sc 311 221015
Lacanobia oleracea (Linnaeus, 1758)p, He 23 17
Lacanobia splendens (Hübner, 1808)o2, He 121 2 1
Lacanobia w-latinum (Hufnagel, 1766)p, He, Sc 412 2021
Leucania obsoleta (Hübner, 1803)m1, He 2 1
Lithophane furcifera (Hufnagel, 1766)o2, Tr2 5 1221
Lithophane ornitopus (Hufnagel, 1766)o2, He, Tr1 1
Macdunnoughia confusa (Stephens, 1850)O1, He 1
Melanchra persicariae (Linnaeus, 1761)p, He 2
Mniotype satura (Denis & Schiffermüller, 1775)p, He, Sc, Tr58411 2 2539
Moma alpium (Osbeck, 1778)o2, Tr 1 2 21
Mythimna albipuncta (Denis & Schiffermüller, 1775)p, He 21 1 2
Mythimna impura (Hübner, 1808)p, He 2 1 1
Mythimna pallens (Linnaeus, 1758)p, He 11 57
Mythimna straminea (Treitschke, 1825)p, Sc, Tr11 13
Family NoctuidaeLarval food necessitiesQP1QP2QP3RN1RN2RN3RN4SC1aSc1bSc2
Mythimna turca (Linnaeus, 1761)O1, He2 1221 6
Noctua fimbriata (Schreber, 1759)p, He, Sc1 2 4
Noctua janthina (Denis & Schiffermüller, 1775)p, He, Sc 1 8
Noctua pronuba (Linnaeus, 1758)o2, He, Sc1269 1 23917
Ochropleura plecta (Linnaeus, 1761)o2, He2121273962194142130
Oligia latruncula (Denis & Schiffermüller, 1775)p, He 16
Panolis flammea (Denis & Schiffermüller, 1775)m1, He1
Panthea coenobita (Esper, 1785)o1, Con, TR 1
Phragmatiphila nexa (Hübner, 1808)o2, He11017 2323 3
Plusia festucae (Linnaeus, 1758)p, He 1 1
Deltote pygarga (Hufnagel, 1766)O1, He6941595752128
Pseudeustrotia candidula (Denis & Schiffermüller, 1775)p, He 11 7
Rhizedra lutosa (Hübner, 1803)m1, He 1 1
dina buettneri (E. Hering, 1858p, Tr 3
Senta flammea (Curtis, 1828)m1, He 1 7 3
Sideridis rivularis (Fabricius, 1775)O1, He 11 1 1 18
Simyra albovenosa (Goeze, 1781)p, He, Sc 12
Staurophora celsia (Linnaeus, 1758)O1, He 2 33
Thalpophila matura (Hufnagel, 1766)O1, He 1
Tholera cespitis (Denis & Schiffermüller, 1775)p, He6 1
Tholera decimalis (Poda, 1761)p, He 3 94
Trachea atriplicis (Linnaeus, 1758)p, He 1
Xanthia icteritia (Hufnagel, 1766)p, Sc, Tr 1 11 76
Xanthia togata (Esper, 1788)o2, Sc, Tr 1 111131
Xestia c-nigrum (Linnaeus, 1758)p, He435 65108614
Xestia baja (Denis & Schiffermüller, 1775)p, He, Sc, Tr1 1 135
Xestia sexstrigata (Haworth, 1809)o2, He522 71011
Xestia triangulum (Hufnagel, 1766)p, He, Sc, Tr1822 9101222222
Xestia xanthographa (Denis & Schiffermüller, 1775)o2, He 1
Family Nolidae
Earias vernana (Fabricius, 1787)m1, Tr 1
Pseudoips prasinana (Linnaeus, 1758)o2, Sc, Tr 11
Earias clorana (Linnaeus, 1761)m2, Sc, Tr 1 1 1 1
Family Notodontidae
Cerura erminea (Esper, 1783)m2, Tr 21
Clostera anastomosis (Linnaeus, 1758)o1, Sc, Tr 11 1 13
Clostera curtula (Linnaeus, 1758)o1, Sc, Tr 31 13
Clostera pigra (Hufnagel, 1766)o1, Sc, Tr 23 1 2121
Family NotodontidaeLarval food necessitiesQP1QP2QP3RN1RN2RN3RN4SC1aSc1bSc2
Furcula bicuspis (Borkhausen, 1790)O1, Tr1 14 2411
Gluphisia crenata (Esper, 1785)o1, Sc, Tr 1 1 1
Leucodonta bicoloria (Denis & Schiffermüller, 1775)m2, Tr111331 185
Notodonta dromedarius (Linnaeus, 1767)o2, Sc, Tr2 4383171 1
Notodonta torva (Hübner, 1803)o1, Sc, Tr 1
Notodonta ziczac (Linnaeus, 1758)o2, Sc, Tr 1 311
Odontosia carmelita (Esper, 1799)o1, Sc, Tr 1
Phalera bucephala (Linnaeus, 1758)o2, Sc, Tr45234313685
Pheosia gnoma (Fabricius, 1776)m2, Tr5116434 537
Pheosia tremula (Clerck, 1759)o1, Sc, Tr 214 27
Pterostoma palpina (Clerck, 1759)O1, He, Sc 3 1
Ptilodon capucina (Linnaeus, 1758)o2, Sc, Tr443221 288
Stauropus fagi (Linnaeus, 1758)o2, Sc, Tr 3113321
Family Sphingidae
Deilephila elpenor (Linnaeus, 1758)o2, He 11 1 1
Hyles gallii (Rottemburg, 1775)o2, He 1 2
Laothoe populi (Linnaeus, 1758)o1, Sc, Tr1 31461122
Mimas tiliae (Linnaeus, 1758)o2, Tr 1 1
Smerinthus ocellata (Linnaeus, 1758)o2, Sc, Tr131732142 3

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Figure 1. Localization of the sites where the inventory of the Macroheterocera fauna was carried out in the Narew National Park (NNP), Podlaskie Voivodeship, North-East Poland.
Figure 1. Localization of the sites where the inventory of the Macroheterocera fauna was carried out in the Narew National Park (NNP), Podlaskie Voivodeship, North-East Poland.
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Figure 2. Light spectrum of the actinic lamp (upper image) and of the ultraviolet lamp (bottom image) used in this study [65,66].
Figure 2. Light spectrum of the actinic lamp (upper image) and of the ultraviolet lamp (bottom image) used in this study [65,66].
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Figure 3. The Mantel test scattered plot of the distances between the studied forest sites in the NNP, Podlaskie, North-East Poland (kilometers) and the Morisita values of species similarity among sites.
Figure 3. The Mantel test scattered plot of the distances between the studied forest sites in the NNP, Podlaskie, North-East Poland (kilometers) and the Morisita values of species similarity among sites.
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Figure 4. Number of individuals and species according to their larval host plant specificity (A) and to the plant type (B). m1-1st degree monophagous, m2-2nd degree monophagous, o1-1st degree oligophagous, o2-2nd degree oligophagous (see text for details) and po-polyphagous; He-herbaceous, Sc-scrubs, Tr-trees and Ot-lichens, mosses or decaying organic material.
Figure 4. Number of individuals and species according to their larval host plant specificity (A) and to the plant type (B). m1-1st degree monophagous, m2-2nd degree monophagous, o1-1st degree oligophagous, o2-2nd degree oligophagous (see text for details) and po-polyphagous; He-herbaceous, Sc-scrubs, Tr-trees and Ot-lichens, mosses or decaying organic material.
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Figure 5. Share of individuals per family in each forest (NNP, Podlaskie, North-East Poland) plant community according to: the family they represent (A); the plant type (B) and to their plant specificity (C) m1-1st degree monophagous, m2-2nd degree monophagous, o1-1st degree oligophagous, o2-2nd degree oligophagous (see text for details) and po-polyphagous; He-herbaceous, Sc-scrubs, Tr-trees and Ot-lichens, mosses or decaying organic material. In all graphics, the inner circle represents the Ribesum-nigrum alnetum and the middle circle represents the Querco-Pinetum, while the outer circler represents the Substitute Communities.
Figure 5. Share of individuals per family in each forest (NNP, Podlaskie, North-East Poland) plant community according to: the family they represent (A); the plant type (B) and to their plant specificity (C) m1-1st degree monophagous, m2-2nd degree monophagous, o1-1st degree oligophagous, o2-2nd degree oligophagous (see text for details) and po-polyphagous; He-herbaceous, Sc-scrubs, Tr-trees and Ot-lichens, mosses or decaying organic material. In all graphics, the inner circle represents the Ribesum-nigrum alnetum and the middle circle represents the Querco-Pinetum, while the outer circler represents the Substitute Communities.
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Figure 6. The NMSD analysis of the total amount of species collected (A), excluding singleton and doubleton species (B) and SD species by their own (C) in the studied forest sites (NNP, Podlaskie, North-East Poland).
Figure 6. The NMSD analysis of the total amount of species collected (A), excluding singleton and doubleton species (B) and SD species by their own (C) in the studied forest sites (NNP, Podlaskie, North-East Poland).
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Figure 7. CLM dendrogram of the total amount of species collected (A) when the SD species were removed from the analysis (B) and the SD species by their own (C) in the studied forest sites (NNP, Podlaskie, North-East Poland).
Figure 7. CLM dendrogram of the total amount of species collected (A) when the SD species were removed from the analysis (B) and the SD species by their own (C) in the studied forest sites (NNP, Podlaskie, North-East Poland).
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Figure 8. Share of SD species according to their larvae plant specificity. m1-1st degree monophagous, m2-2nd degree monophagous, o1-1st degree oligophagous, o2-2nd degree oligophagous (see text for details) and po-polyphagous.
Figure 8. Share of SD species according to their larvae plant specificity. m1-1st degree monophagous, m2-2nd degree monophagous, o1-1st degree oligophagous, o2-2nd degree oligophagous (see text for details) and po-polyphagous.
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Figure 9. Total number of species collected (continuous lines) and SD species (dashed lines) collected per month in each site (NNP, Podlaskie, North-East Poland). The R value under each site represent the correlation between the total number of species collected per month in each forest site with the number of SD species collected per month in each site.
Figure 9. Total number of species collected (continuous lines) and SD species (dashed lines) collected per month in each site (NNP, Podlaskie, North-East Poland). The R value under each site represent the correlation between the total number of species collected per month in each forest site with the number of SD species collected per month in each site.
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Table 1. Distances between the studied forest sites in the NNP, Podlaskie, North-East Poland (kilometers) are present in the down triangle and the Morisita similarity values of the distribution of the number species among sites based on the species larval food necessities in the upper triangle.
Table 1. Distances between the studied forest sites in the NNP, Podlaskie, North-East Poland (kilometers) are present in the down triangle and the Morisita similarity values of the distribution of the number species among sites based on the species larval food necessities in the upper triangle.
QP1QP2QP3RN1RN2RN3RN4SQPaSQPbSRN
QP1-0.98570.98380.96980.95330.97630.96540.98620.98350.9877
QP210.46-0.98830.97410.95380.96140.96120.97130.9890.989
QP310.120.25-0.98130.96020.95690.9610.97810.9830.9893
RN14.875.485.66-0.97530.94180.93490.97610.96180.9697
RN28.531.451.663.98-0.94770.93970.98060.94550.9428
RN311.2317.8818.0812.716-0.97570.97670.97120.9659
RN44.329.839.664.318.28.51-0.96570.96950.9734
SQPa10.350.835.565.932.0318.5310.25-0.96830.9721
SQPb10.350.835.565.932.0318.5310.250-0.9879
SRN11.481.561.387.053.0319.3811.231.151.5-
Table 3. Number of species and individuals from the families recorded in each study site in the NNP, Podlaskie, North-East Poland.
Table 3. Number of species and individuals from the families recorded in each study site in the NNP, Podlaskie, North-East Poland.
QP1QP2QP3RN1RN2RN3RN4SQPaSQPbSRN
Spp.Ind.Spp.Ind.Spp.Ind.Spp.Ind.Spp.Ind.Spp.Ind.Spp.Ind.Spp.Ind.Spp.Ind.Spp.Ind.
Geometridae533005558163541574584321449600438635846572103273804
Noctuidae3514640130401402711923833611739305351355527453384
Erebidae18137241562420525403241071820719588262673024327403
Notodontidae61791911341023117681294392512431136
Drepanidae523317517528516485116631645639
Lasiocampidae34531332734621421524732133528
Sphingidae384847411311283163736510
Cossidae11131500001418000012
Hepialidae15001300001713111817
Nolidae00110000110011003311
Total1256821409281529791311088112522121978123199014195218516891801694
Table 4. Number of SD species among the studied forest sites (NNP, Podlaskie, North-East Poland) per family. SD-total amount of SD species in each site. % of SD-SD species share of the total number of species per site.
Table 4. Number of SD species among the studied forest sites (NNP, Podlaskie, North-East Poland) per family. SD-total amount of SD species in each site. % of SD-SD species share of the total number of species per site.
QP1QP2QP3RN1RN2RN3RN4SQPaSQPbSRN
Geometridae2918243014144242022
Noctuidae22272719142322152717
Erebidae591071464475
Notodontidae3675465575
Drepanidae2141221322
Sphingidae2233012224
Lasiocampidae1112001111
Cossidae1000000001
Nolidae0100101031
Hepialidae0000000100
Total SD65657667495240556958
% of SD52%46%50%51%44%43%33%39%37%32%
Table 5. Biodiversity indices values of each studied forest site (NNP, Podlaskie, North-East Poland). H’-Shannon index value, F’s-Fischer alfa index value and E’-Evenness index value. The ‘all’ refers to the values when all species were taken into consideration while the ‘NoSD’ represents the values when the SD species were excluded, T. Spp.-total number of species collected, Chao1-Chao 1 estimator of the true species diversity of a sample.
Table 5. Biodiversity indices values of each studied forest site (NNP, Podlaskie, North-East Poland). H’-Shannon index value, F’s-Fischer alfa index value and E’-Evenness index value. The ‘all’ refers to the values when all species were taken into consideration while the ‘NoSD’ represents the values when the SD species were excluded, T. Spp.-total number of species collected, Chao1-Chao 1 estimator of the true species diversity of a sample.
QP1QP2QP3RN1RN2RN3RN4SQPaSQPbSRN
H’ all4.24.034.13.74.33.9444.24.384.53
H’ NoSD3.83.673.73.43.93.663.83.94.174.36
F’s A all4545.850394436.429465350.9
F’s A NoSD1719.920152017.4182428.730.6
E’ all0.90.820.80.80.90.820.80.90.840.87
E’ NoSD0.90.850.90.810.860.90.90.880.91
T. Spp.125140152131112121123141185180
Chao1170180245198146145136165245252
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Matos da Costa, J.; Sielezniew, M. The Contribution of Singletons and Doubletons Captured Using Weak Light Heath Traps for the Analysis of the Macroheteroceran Assemblages in Forest Biotopes. Diversity 2023, 15, 508. https://doi.org/10.3390/d15040508

AMA Style

Matos da Costa J, Sielezniew M. The Contribution of Singletons and Doubletons Captured Using Weak Light Heath Traps for the Analysis of the Macroheteroceran Assemblages in Forest Biotopes. Diversity. 2023; 15(4):508. https://doi.org/10.3390/d15040508

Chicago/Turabian Style

Matos da Costa, João, and Marcin Sielezniew. 2023. "The Contribution of Singletons and Doubletons Captured Using Weak Light Heath Traps for the Analysis of the Macroheteroceran Assemblages in Forest Biotopes" Diversity 15, no. 4: 508. https://doi.org/10.3390/d15040508

APA Style

Matos da Costa, J., & Sielezniew, M. (2023). The Contribution of Singletons and Doubletons Captured Using Weak Light Heath Traps for the Analysis of the Macroheteroceran Assemblages in Forest Biotopes. Diversity, 15(4), 508. https://doi.org/10.3390/d15040508

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