Research Article |
Corresponding author: Romain Nattier ( nattier@mnhn.fr ) Academic editor: Michael Thomas
© 2018 Séverin Jouveau, Mathilde Delaunay, Régine Vignes-Lebbe, Romain Nattier.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Jouveau S, Delaunay M, Vignes-Lebbe R, Nattier R (2018) A multi-access identification key based on colour patterns in ladybirds (Coleoptera, Coccinellidae). ZooKeys 758: 55-73. https://doi.org/10.3897/zookeys.758.22171
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An identification key based on French ladybird colouration is proposed for the tribes Chilocorini, Coccinellini, and Epilachnini. These tribes were chosen based on their relatively limited species diversity, as well as their large size and high colour diversity, making them easy to observe and collect. The identification key runs on Xper3 software, which allows the building of structured knowledge bases and online free-access keys. The online interactive Xper key is available at http://french-ladybird.identificationkey.fr.
Citizen science, Coccinellidae , Coccinellini , Chilocorini , Epilachnini , France, interactive identification keys, ladybirds, Xper
The identification of species is central in ecology, conservation biology, systematics, and related disciplines (species inventories and community studies, ecosystem management, establishment and improvement of environmental public policies, taxonomic reviews, and management of natural history collections) (
If the identification of large and charismatic animals may be easy, the majority of organisms require expert skills for accurate identification and the inability to identify species represents a major challenge known as the Taxonomic Impediment (
Coccinellidae is a family of beetles popular and appreciated by naturalists and the general public. Because these animals have ecological and economic values as predators of pest insects (e.g. aphids, scale insects), their identification may be of importance for naturalists, amateurs and professionals (
Single-access identification keys consist of a series of identification steps that form a single and unique identification path for a given taxon. Although it is a very powerful tool for identifying species, the user cannot choose the character to be observed (the answer for every single step must be known), and the identification is impossible if some characters are missing (e.g., if the specimen is poorly preserved). Moreover, this type of keys cannot be modulated or adapted to various kinds of publics, environmental conditions, season, or geographical location.
Most North American or European ladybird identification keys are single-access and difficult to use for non-specialists (
Modern tools developed along with digital technologies and data processing make identification easier for the user. In this perspective, several interactive identification keys (IIK) are available online (e.g., http://www.ladybird-survey.org/bbc/spotter.php, http://www.discoverlife.org/20/q?guide=Ladybug), but most of them are only digital versions of single-access keys and maintain the same difficulties for the user.
A multi-access interactive key is a computer-aided identification tool that makes it possible to find correct names of species where the user enters attributes (character-state values) of the specimen (
This study aims to i) release the first multi-access digital interactive identification key for French ladybirds based on colour that takes into account intraspecific variability; and ii) study and discuss the discriminating power of the characters: can we identify species by colour pattern only? What are the most discriminating characters?
As the aim of the key is to provide an identification tool for the general public in the perspective of citizen science programs, we have restricted the taxonomic coverage to the tribes Chilocorini, Coccinellini and Epilachnini (Table
Coccinellinae Latreille, 1807 | Chilocorini Mulsant, 1846 | Chilocorus bipustulatus Linnaeus, 1758 |
Chilocorus renipustulatus Scriba, 1791 | ||
Exochomus octosignatus Gebler, 1830 | ||
Exochomus quadripustulatus Linnaeus, 1758 | ||
Parexochomus nigromaculatus Goeze, 1777 | ||
Coccinellini Latreille, 1807 | Halyzia sedecimguttata Linnaeus, 1758 | |
Psyllobora vigintiduopunctata Linnaeus, 1758 | ||
Vibidia duodecimguttata Poda, 1761 | ||
Anisosticta novemdecimpunctata Linneaus, 1758 | ||
Coccinula quatuordecimpustulata Linnaeus, 1758 | ||
Coccinula sinuatoinarginata Faldermann, 1837 | ||
Tytthaspis sedecimpunctata f. duodecimpunctata | ||
Adalia bipunctata f. annulata | ||
Adalia bipunctata f. pantherina | ||
Adalia bipunctata f. quadrimaculata | ||
Adalia bipunctata f. sexpustulata | ||
Adalia bipunctata f. typica | ||
Adalia conglomerata f. decas | ||
Adalia decempunctata f. decempustulata | ||
Adalia decempunctata f. guttatopunctata | ||
Adalia decempunctata f. lutea | ||
Adalia decempunctata f. quadripunctata | ||
Adalia decempunctata f. scribai | ||
Adalia decempunctata f. subpunctata | ||
Adalia decempunctata f. terna | ||
Adalia decempunctata f. typica | ||
Anatis ocellata Linnaeus, 1758 | ||
Aphidecta obliterata f. typica | ||
Calvia decemguttata Linnaeus, 1767 | ||
Calvia quatuordecimguttata Linnaeus, 1758 | ||
Calvia quindecimguttata Fabricius, 1777 | ||
Ceratomegilla alpina Villa A. & Villa G. B., 1835 | ||
Ceratomegilla notata Laicharting, 1781 | ||
Ceratomegilla rufocincta Mulsant, 1850 | ||
Ceratomegilla undecimnotata Schneider D.H. 1792 | ||
Coccinella venusta Weise, 1879 | ||
Coccinella hieroglyphica Linnaeus, 1758 | ||
Coccinella magnifica Redtenbacher, 1843 | ||
Coccinella quinquepunctata Linnaeus, 1758 | ||
Coccinella septempunctata Linnaeus, 1758 | ||
Coccinella undecimpunctata Linnaeus, 1758 | ||
Harmonia axyridis f. conspicua | ||
Harmonia axyridis f. novemdecimsignata | ||
Harmonia axyridis f. spectabilis | ||
Harmonia conformis Boisduval, 1835 | ||
Harmonia quadripunctata f. sedecimpunctata | ||
Harmonia quadripunctata f. typica | ||
Hippodamia septemmaculata f. cestiva | ||
Hippodamia tredecimpunctata Linnaeus, 1758 | ||
Hippodamia variegata f. quinquemaculata | ||
Hippodamia variegata f. undecimpunctata | ||
Hippodamia variegata f. carpini | ||
Hippodamia variegata f. constellata | ||
Coccinellinae Latreille, 1807 | Coccinellini Latreille, 1807 | Hippodamia variegata f. velox |
Myrrha octodecimguttata Linnaeus, 1758 | ||
Myzia oblongoguttata Linnaeus, 1758 | ||
Oenopia conglobata Linnaeus, 1758 | ||
Oenopia doublieri Mulsant, 1846 | ||
Oenopia impustulata Linnaeus, 1767 | ||
Oenopia lyncea Olivier, 1808 | ||
Propylea quatuordecimpunctata f. weisei | ||
Sospita vigintiguttata Linnaeus, 1758 | ||
Epilachnini Chevrolat in Dejean, 1837 | Henosepilachna argus Geoffroy in Fourcroy, 1785 | |
Henosepilachna elaterii Rossi, 1794 | ||
Subcoccinella vigintiquatuorpunctata f. limbata | ||
Subcoccinella vigintiquatuorpunctata f. typica |
The current taxonomy (
Specimens were examined in the collection of the Muséum national d’Histoire naturelle, Paris, France (
A list of 21 morphological characters based on colour and shape is defined, mainly from existing identification keys (
Pronotum |
1. Pronotum colours |
2. Pattern on pronotum |
3. Number of pronotum patterns |
4. Type of pronotum patterns |
Elytra |
5. Elytra main colour (background) |
6. Elytra markings |
7. Colour of elytra markings |
8. Number of elytra markings |
9. Type of elytra markings |
10. Number of lateral lines in the elytra markings |
11. Number of longitudinal lines in the elytra markings |
12. A spot in the first third of the elytra |
13. One of the spots reaches the rim of the elytra |
14. Cream ring around dots |
15. Dark sutural elytra band |
16. Scutellar spot |
17. Shape of the scutellar spot |
18. White marks between the scutellar spot and the elytra basis |
19. Distinct rim around the edge of the elytra |
20. Elytra covered with short hairs |
Underside |
21. Small white triangular marks on the underside below both the middle and front legs |
Digitalization of the 47 species was performed using Xper2 v.2.3.2 (
An Xper knowledge base is a set of items described using the same model and terminology, and documented by texts and images. In this key there are 66 items covering 47 species and 19 intraspecies colour forms. The descriptive model consists of a hierarchy of descriptors and a chosen terminology for expressing different possible values (states). The descriptors are the 21 morphological characters previously described. Some of them are consistent only if some conditions are true for another descriptor and these dependencies define a hierarchical structure of descriptors (Table
Xper3 was also used to compare species and genera. For each descriptor, the comparison tests are able to distinguish a pair of items. Three different measures are available (
The discriminatory power, which represents the quantitative assessments of the ability of a descriptor to distinguish taxa, is measured with the Xper original index (
Comparisons within and between genera are made with the “compare groups” and “compare items” options of Xper3. For the comparison between genera, we estimated the number of discriminating characters, weighted or not by the number of colour forms. A subset of descriptors sufficient to discriminate the total of descriptors with the same efficiency was also calculated with the “minset” tool (
Two types of keys are available: free-access keys and single-access keys (
A single-access identification key was generated by IKey+ under Xper3 with the default option and the Xper score method. In this case we show four statistics by taxon: the number of steps, the length of the shortest and the longest paths, and the average length of paths. This key was then compared with five single-access keys for European ladybirds (
The consistency of the knowledge base has been tested with the “Checkbase” functionality of Xper3: no items share the same description and all items are described. The base is 100% complete. Twenty-one descriptors are used: five do not have any dependence (either father or son), four are parent descriptors (for which two are also child descriptors) and 14 are child descriptors (for which two are also parent descriptors). Ninety-eight states are described (minimal/maximal/average number of states: 2/12/4.67).
List of the discriminating power efficiency of descriptors. Those written in bold are sufficient for discriminating all taxa.
Category | Descriptor | XPER index | Number of descriptor states |
---|---|---|---|
Pronotum | 4. Type of pronotum pattern(s) | 0.85 | 10 |
Elytra | 8. Number of elytra markings | 0.85 | 12 |
Elytra | 10. Number of lateral lines of elytra markings | 0.83 | 8 |
Elytra | 11. Number of longitudinal lines of elytra markings | 0.82 | 7 |
Pronotum | 3. Number of pronotum patterns | 0.71 | 9 |
Elytra | 17. Shape of the scutellar spot | 0.68 | 4 |
Elytra | 5. Elytra main colour (background) | 0.53 | 7 |
Elytra | 7. Colour of elytra markings | 0.52 | 6 |
Elytra | 13. One of the spots reaches the rim of the elytra | 0.49 | 2 |
Elytra | 18. White marks between the scutellary spot and the elytra basis | 0.47 | 2 |
Elytra | 16. Scutellar spot | 0.46 | 2 |
Elytra | 12. A spot in the first third of the elytra | 0.4 | 2 |
Elytra | 9. Type of elytra markings | 0.32 | 6 |
Elytra | 15. Dark sutural elytra band | 0.18 | 2 |
Elytra | 19. Distinct rim around the edge of the elytra | 0.14 | 2 |
Elytra | 6. Elytra markings | 0.12 | 2 |
Elytra | 20. Elytra covered in short hairs | 0.12 | 2 |
Pronotum | 2. Pattern on pronotum | 0.11 | 2 |
Pronotum | 1. Pronotum colours | 0.09 | 7 |
Elytra | 14. Cream ring around dots | 0.06 | 2 |
Underside | 21. Small white triangular marks on the underside below both the middle and front legs | 0.03 | 2 |
The four most discriminating characters (XPER index >0.8) are the type of pronotum patterns (#5), the number of elytra markings (#8), and the number of lateral (#10) and longitudinal (#11) lines of elytra markings. These characters can separate taxa in 7 to 13 groups. For example, the two most discriminating characters (#5 and #8) split all the remaining taxa in 10–13 different groups including 2–13 taxa per group.
The characters #14 and #21 are the least discriminating as they both have an XPER index below 0.8. These characters are binary and split all taxa in two unequal groups (60 vs 2 for the character #14, 65 vs 1 for the character #21). Despite its weak discriminating power, the character #21 is the only one that can distinguish the two species Coccinella septempunctata and C. magnifica. Eleven descriptors are sufficient to separate all taxa (Table
Comparison within a genus: Coccinella (Table
Comparison within genus: Coccinella (six species), showing informative characters (in bold) and constant characters (in regular). The intersection column shows what is constant within the genus.
Coccinella venusta | Coccinella hieroglyphica | Coccinella magnifica | Coccinella quinquepunctata | Coccinella septempunctata | Coccinella undecimpunctata | UNION | INTERSECTION | |
---|---|---|---|---|---|---|---|---|
1. Pronotum colours | Black; White | |||||||
2. Elytra main colour (background) | Black; Orange | Black; Orange | Red; Orange | Red; Orange | Red; Orange | Red; Orange | Black; Orange; Red | Orange |
3. Pattern on pronotum | Present | |||||||
4. Number of pronotum patterns | 2 | |||||||
5. Type of pronotum patterns | Central structure (solid, trapezium) with 2 anterior-lateral white or orange marks | |||||||
6. Elytra markings | Present | |||||||
7. Colour of elytra markings | Red; Orange; Black | Black | Black | Black | Black | Black | Red; Orange; Black | Black |
8. Number of elytra markings | 1; 2 | 7 | 7 | 5 | 7 | 10–14 | 1; 2; 5; 7; 10–14 | |
9. Type of elytra markings | Other | Ovoid shape spot; Other | Regular dots | Regular dots | Regular dots | Regular dots | Ovoid shape spot; Regular dots; Other | |
10. Number of lateral lines of elytra markings | not applicable | 3 | 3 | 3 | 3 | 3 | 3 | |
11. Number of longitudinal lines of elytra markings | not applicable | 4 | 5 | 5 | 5 | 5 | 4; 5 | |
12. Scutellar spot | Absent | Absent | Present | Present | Present | Present | Absent; Present | |
13. Distinct rim around the edge of the elytra | absent | |||||||
14. Elytra covered in short hairs | No | |||||||
15. Small white triangular marks on the underside below both the middle and front legs | Absent | Absent | Present | Absent | Absent | Absent | Absent; Present | |
16. Cream ring around dots | Absent | |||||||
17. Dark sutural elytra band | Absent | |||||||
18. Shape of the scutellar spot | not applicable | not applicable | A drop (spot with a neck) | A drop (spot with a neck) | A drop (spot with a neck) | A drop (spot with a neck) | A drop (spot with a neck) | |
19. White marks between the scutellar spot and the basis of elytra | not applicable | not applicable | Present | Present | Present | Present | Present | |
20. One of the spots reaches the rim of the elytra | No | Yes | No | No | No | No | Yes; No | |
21. A spot in the first third of the elytra | Absent | Present | Present | Absent | Present | Present | Absent; Present |
Among the 21 characters, 12 are informative (in blue) whereas the other nine are constant and cannot discriminate within this genus (in red). The intersection column shows what is constant in Coccinella, therefore helping with the description of the genus: black and white pronotum with two patterns (Central structure - solid, trapezium with two anterior-lateral white or orange marks), elytra with different markings, but always devoid of rim around the edge, short down hairs, cream rings around dots, or dark sutural band.
Comparison between the 24 genera included in the study (Table
Comparison between the 24 genera included in the study, showing the most constant or variable genus, weighted or not by the intraspecific variability taken into account in this study (number of colour forms).
Number of species studied | Number of colour forms studied | Number of discriminating characters within genus | ||
---|---|---|---|---|
with all colours forms | weighted by the number of colour forms | |||
Adalia | 3 | 14 | 16 (76%) | 1.1 |
Calvia | 3 | 3 | 5 (23%) | 1.7 |
Ceratomegilla | 4 | 4 | 13 (62%) | 3.3 |
Chilocorus | 2 | 2 | 4 (19%) | 2 |
Coccinella | 6 | 6 | 11 (52%) | 1.8 |
Coccinula | 2 | 2 | 1 (5%) | 0.5 |
Exochomus | 2 | 2 | 7 (33%) | 3.5 |
Harmonia | 3 | 6 | 14 (67%) | 2.3 |
Henosepilachna | 2 | 2 | 3 (14%) | 1.5 |
Hippodamia | 3 | 7 | 10 (48 %) | 1.4 |
Oenopia | 4 | 4 | 10 (48%) | 2.5 |
Subcoccinella | 1 | 2 | – | – |
Anatis | 1 | 1 | – | – |
Anisosticta | 1 | 1 | – | – |
Aphidecta | 1 | 1 | – | – |
Halyzia | 1 | 1 | – | – |
Myrrha | 1 | 1 | – | – |
Myzia | 1 | 1 | – | – |
Parexochomus | 1 | 1 | – | – |
Propylea | 1 | 1 | – | – |
Psyllobora | 1 | 1 | – | – |
Sospita | 1 | 1 | – | – |
Tytthaspis | 1 | 1 | – | – |
Vibidia | 1 | 1 | – | – |
Among the genera with at least two species studied, the most constant are Coccinula (5% of discriminating characters), Henosepilachna (14%) and Chilocorus (19%); the most variable are Adalia (76%) and Harmonia (67%). If weighted by the number of described colour forms per genera, the most constant are still Coccinula and Henosepilachna; whereas the most variable genera are Ceratomegilla and Exochomus.
Single-access identification key and comparison with standard keys
For each identification, the descriptive statistics of the generated key (Appendix
Representation of a part of the single-access identification key generated by IKey+ under Xper3 and the Xper score method (statistics detailed in Appendix
In comparison to other standard keys (Table
Comparison between five single-access keys from bibliography and the generated key by IKey+ under Xper3 for 13 ladybird species: average number of steps (number of paths, length of the shortest and the longest paths).
Generated Key - this study 47 species |
|
|
|
|
|
|
---|---|---|---|---|---|---|
Adalia bipunctata f. typica | 4 (1) | 5 (1) | 8 (1) | 11 (1) | 7 (1) | 9 (1) |
Anatis ocellata | 3 (2) | 6 (1) | 4.6 (3, 3–7) | 4 (1) | 8 (3, 6–10) | 8 (2, 7–9) |
Anisosticta novemdecimpunctata | 3 (1) | 5 (1) | 9 (1) | 5 (1) | 10 (1) | 5.5 (2, 3–8) |
Calvia quatuordecimguttata | 5 (1) | 11 (1) | 12 (1) | 10 (1) | 8 (1) | 11 (1) |
Chilocorus bipustulatus | 4 (1) | - | 7 (1) | 4 (1) | 4 (1) | 8 (1) |
Coccinella hieroglyphica | 4 (2) | 10 (1) | 6.5 (4, 4–9) | - | 8 (1) | 8 (2, 7–9) |
Coccinella quinquepunctata | 3 (1) | 12 (1) | 8 (1) | - | 10 (1) | 13 (1) |
Coccinella septempunctata | 5 (1) | 13 (1) | 8 (1) | 12 (1) | 11 (1) | 14 (1) |
Coccinella undecimpunctata | 5 (1) | 11 (1) | 8.5 (2, 8–9) | 11 (1) | 10 (1) | 10.5 (2, 10–11) |
Coccinula quatuordecimpustulata | 6 (2) | 10 (1) | 5 (1) | 14 (1) | - | 11 (1) |
Myzia oblongoguttata | 5.5 (2, 5–6) | 6 (1) | 6 (1) | 6 (1) | 7 (1) | 7 (1) |
Oenopia conglobata | 4 (1) | 10 (1) | 11 (1) | 9 (1) | - | 6 (1) |
Psyllobora vigintiduopunctata | 3 (1) | 8 (1) | 11 (1) | 12 (1) | 12 (1) | 7 (1) |
The work presented in this study led to the release of the first multi-access interactive digital identification key for French ladybirds. The adaptability and great number of possibilities provided by this new generation tool are unparalleled for this group, and make the key very flexible and abundantly illustrated and described, thanks to images and texts. Since it is available online and open to experts for modification, the identification key can easily be improved. It will be possible to add ladybird taxa and to extend the geographic area (e.g., a key to all European ladybirds).
Most classical and single-access keys share characters that are quite difficult to observe for students, naturalists and the general public (e.g. for ladybirds in
Identification in the field is traditionally realised with paper-printed keys, but recent developments of mobile devices make it possible to use portable version of digital keys. In this perspective, the project “KeyToNature” (www.keytonature.eu) aims to develop new, more convenient and paper-free identification tools, for use within schools and universities across Europe and available on a variety of mobile platforms (laptops, smartphones). However these keys are single access, e.g., MobileKey (
Both experienced and inexperienced users are likely to succeed in identifying problematic species (
We thank Clémentine Dantard for the illustrations, and Frédéric Chevaillot, Fabrice Duvivier, Jean-François Godeau, David Gonzales, Donald Hobern, Guillaume Jacquemin, Julien Aït El Mekki, Andgelo Mombert, and Gilles San Martin for providing field pictures of the species. We also thank Sylvain Bouquin for his technical help with the Xper system, and Estelle Bourdon and Donald Davesne for proofreading.
STATISTICS | ||||
---|---|---|---|---|
Taxon | Number of paths leading to taxon | Length of the shortest path leading to taxon | Average length of paths leading to taxon | Length of the longest path leading to taxon |
Adalia bipunctata f. annulata | 1 | 4 | 4 | 4 |
Adalia bipunctata f. pantherina | 1 | 4 | 4 | 4 |
Adalia bipunctata f. quadrimaculata | 2 | 5 | 5.5 | 6 |
Adalia bipunctata f. sexpustulata | 2 | 3 | 3.5 | 4 |
Adalia bipunctata f. typica | 1 | 4 | 4 | 4 |
Adalia conglomerata f. decas | 1 | 3 | 3 | 3 |
Adalia decempunctata f. decempustulata | 2 | 7 | 7 | 7 |
Adalia decempunctata f. guttatopunctata | 4 | 7 | 7 | 7 |
Adalia decempunctata f. lutea | 1 | 2 | 2 | 2 |
Adalia decempunctata f. quadripunctata | 1 | 5 | 5 | 5 |
Adalia decempunctata f. scribai | 1 | 5 | 5 | 5 |
Adalia decempunctata f. subpunctata | 1 | 4 | 4 | 4 |
Adalia decempunctata f. terna | 1 | 3 | 3 | 3 |
Adalia decempunctata f. typica | 2 | 6 | 6 | 6 |
Anatis ocellata | 2 | 3 | 3 | 3 |
Anisosticta novemdecimpunctata | 1 | 3 | 3 | 3 |
Aphidecta obliterata f. typica | 2 | 2 | 3 | 4 |
Calvia decemguttata | 2 | 6 | 6 | 6 |
Calvia quatuordecimguttata | 1 | 5 | 5 | 5 |
Calvia quindecimguttata | 2 | 6 | 6 | 6 |
Ceratomegilla alpina | 2 | 5 | 5.5 | 6 |
Ceratomegilla notata | 2 | 6 | 7 | 8 |
Ceratomegilla rufocincta | 1 | 3 | 3 | 3 |
Ceratomegilla undecimnotata | 1 | 8 | 8 | 8 |
Chilocorus bipustulatus | 1 | 4 | 4 | 4 |
Chilocorus renipustulatus | 1 | 3 | 3 | 3 |
Coccinella hieroglyphica | 2 | 4 | 4 | 4 |
Coccinella magnifica | 1 | 5 | 5 | 5 |
Coccinella quinquepunctata | 1 | 3 | 3 | 3 |
Coccinella septempunctata | 1 | 5 | 5 | 5 |
Coccinella undecimpunctata | 1 | 5 | 5 | 5 |
Coccinella venusta | 2 | 4 | 5 | 6 |
Coccinula quatuordecimpustulata | 2 | 6 | 6 | 6 |
Coccinula sinuatomarginata | 1 | 5 | 5 | 5 |
Exochomus octosignatus | 1 | 2 | 2 | 2 |
Exochomus quadripustulatus | 1 | 4 | 4 | 4 |
Halyzia sedecimguttata | 1 | 5 | 5 | 5 |
Harmonia axyridis f. conspicua | 2 | 6 | 6 | 6 |
Harmonia axyridis f. novemdecimsignata | 2 | 4 | 4.5 | 5 |
Harmonia axyridis f. spectabilis | 2 | 5 | 5 | 5 |
Harmonia conformis | 3 | 3 | 4 | 5 |
Harmonia quadripunctata f. sedecimpunctata | 2 | 3 | 3 | 3 |
Harmonia quadripunctata f. typica | 1 | 5 | 5 | 5 |
Henosepilachna argus | 2 | 4 | 5 | 6 |
Henosepilachna elaterii | 1 | 4 | 4 | 4 |
Hippodamia septemmaculata f. cestiva | 2 | 3 | 3.5 | 4 |
Hippodamia tredecimpunctata | 1 | 4 | 4 | 4 |
Hippodamia variegata f. undecimpunctata | 2 | 5 | 6 | 7 |
Hippodamia variegata f. quinquemaculata | 1 | 3 | 3 | 3 |
Hippodamia variegata f. carpini | 1 | 4 | 4 | 4 |
Hippodamia variegata f. constellata | 2 | 3 | 3.5 | 4 |
Hippodamia variegata f. velox | 2 | 5 | 5.5 | 6 |
Myrrha octodecimguttata | 1 | 5 | 5 | 5 |
Myzia oblongoguttata | 4 | 5 | 5.5 | 6 |
Oenopia conglobata | 1 | 4 | 4 | 4 |
Oenopia doublieri | 1 | 4 | 4 | 4 |
Oenopia impustulata | 1 | 2 | 2 | 2 |
Oenopia lyncea | 4 | 5 | 5.5 | 6 |
Parexochomus nigromaculatus | 1 | 2 | 2 | 2 |
Propylea quatuordecimpunctata f. weisei | 2 | 4 | 4 | 4 |
Psyllobora vigintiduopunctata | 1 | 3 | 3 | 3 |
Sospita vigintiguttata | 1 | 3 | 3 | 3 |
Subcoccinella vigintiquatuorpunctata f. limbata | 1 | 2 | 2 | 2 |
Subcoccinella vigintiquatuorpunctata f. typica | 1 | 3 | 3 | 3 |
Tytthaspis sedecimpunctata f. duodecimpunctata | 2 | 5 | 5.5 | 6 |
Vibidia duodecimguttata | 1 | 5 | 5 | 5 |
AVERAGE | 1.53 | 4.197 | 4.364 | 4.53 |