Research Article |
Corresponding author: Tomáš Lackner ( lackobelansky@mac.com ) Academic editor: Michael Caterino
© 2018 Salman Shayya, Nicolas Dégallier, André Nel, Dany Azar, Tomáš Lackner.
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:
Shayya S, Dégallier N, Nel A, Azar D, Lackner T (2018) Contribution to the knowledge of Saprinus Erichson, 1834 of forensic relevance from Lebanon (Coleoptera, Histeridae). ZooKeys 738: 117-152. https://doi.org/10.3897/zookeys.738.21382
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Many histerid beetles are necrophilous on carrion during both active and advanced stages of decomposition. In this study, 13 species of Saprinus were recorded on carrion from Lebanon, containing eight that are new for the Lebanese fauna. The following Saprinus species are newly recorded from Lebanon: 1) Saprinus (S.) caerulescens caerulescens (Hoffmann, 1803); 2) S. (S.) calatravensis Fuente, 1899; 3) S. (S.) chalcites (Illiger, 1807); 4) S. (S.) godet (Brullé, 1832); 5) S. (S.) maculatus (P. Rossi, 1792); 6) S. (S.) strigil Marseul, 1855; 7) S. (S.) submarginatus J. Sahlberg, 1913; and 8) S. (S.) tenuistrius sparsutus Solsky, 1876. The peak activity was recorded, key for the species is provided, and habitus images and male genitalia are illustrated in order to facilitate their taxonomic identifications. Saprinus species are diverse and common on animal carcass; they were likewise collected from a human cadaver in Lebanon. Preliminary comments on biology and distribution of the studied species are given. Our paper represents the first faunistic study on Histeridae from Lebanon. A rigorous research program regarding the biology of Saprinus in Lebanon and the neighbouring countries would greatly improve the knowledge of the diversity, activity, and possible forensic value of Saprinus.
Carrion, key to species, Lebanon, peak activity, Saprinus
Coleoptera (beetles) have proven to be useful in forensic investigations when fly larvae cannot be obtained or when they left the carcass and only beetles could be sampled (
Histerids are unusual among beetles in having just two larval instars. In the first instar the head capsule and sclerites harden and the larva becomes capable of killing and feeding on soft-bodied insects. Mandibles and terminal palpal segments darken and become visible through the egg chorion just before eclosion (
In the Palaearctic Region, 357 species of the Saprininae subfamily have been reported so far (
The majority of specimens were collected on a daily basis during the decomposition process (Table
Stages of decomposition | ||||||
Locality | Coordinates | Season | Fresh | Bloat | Decay | Advanced decay |
Badghan | 33°46'4.24"N, 35°41'14.78"E | Early summer | Day 1 | Day 2–4 | Day 5–9 | Day 10–30 |
Fanar | 33°52'44"N, 35°34'04"E | Spring | Day 1–4 | Day 5–6 | Day 7–16 | Day 17–30 |
Deir El-Ahmar | 34°7'30.96"N, 36°7'22.04"E | Summer | Day 1 | Day 2–3 | Day 4–5 | Day 6–18 |
Naas | 33°54'42.4"N, 35°40'32.7"E | Spring | Day 1–2 | Day 3–6 | Day 7–14 | Day 15–30 |
Hasbaya | 33°23'52.35"N, 35°41.6'6.59"E | Summer | Day 1 | Day 2–3 | Day 4–6 | Day 7–30 |
Hammana | 33°50'N, 33°44'E | Summer | One day sampling | |||
Sin El-Fil | 33°52'N, 35°32'E | Spring | One day sampling | |||
Arsal | 34°10'N, 36°25'E | Autumn | Autopsy done in Beirut |
Four stages of decomposition were reported on the carcasses: 1) fresh stage; 2) bloat stage; 3) active decay stage; and, 4) advanced decay stage. Based on
General observation and dissection were carried out using stereomicroscope Nikon SMZ1500. Without genital extraction, the males of Saprinus species can be usually recognized through the examination of the anterior tarsal setae, which are expanded and lamellate, whereas they are unexpanded and pointed in female. Often the males possess a longitudinal depression on the metaventrite and occasionally also a single or two tiny tubercles on the apical metaventral margin. Male genitalia were first macerated in 10 % KOH solution for about 3 hours, cleared in 80 % ethanol and macerated in lactic acid with fuchsine, incubated at 60 °C for another 30 minutes, and subsequently cleared in 80 % ethanol and then observed in α-terpineol in a small dish. Digital photographs of male genitalia were taken by a Nikon 4500 Coolpix camera and edited in Adobe Photoshop CS5. Genitalia drawings based on the photographs or direct observations were produced with the aid of Hakuba klv-7000 light box. Habitus photographs were taken by F. Slamka (Bratislava, Slovakia). Specimens were measured with an ocular micrometre. Due to the lack of phylogenetic information regarding the genus Saprinus, the taxa in our present paper are arranged alphabetically. For the morphological terminology the reader is referred to
Our field experiments on pig cadavers in five localities across Lebanon (Fig.
The monophyletic (based on the literature references as well as on the on-going molecular studies by T.L.) subfamily Saprininae Blanchard, 1845 is characterised by the presence of distinctive sensory apparatus situated inside their antennal clubs (for more details on this, as well as on the general diagnosis of the subfamily see
= Saprinus incognitus Dahlgren, 1964
= Saprinus therondi Auzat, 1931
Greece.
Slovakia, Southern Europe, Mediterranean subregion, Armenia, Iran (
The biology of S. aegialius is not adequately known, but the species is often found on carcasses. This species is often confused with Saprinus immundus (Gyllenhal, 1808), and it was absent from our sampling (therefore not shown on a distributional map).
Hister caerulescens Hoffmann, 1803
= Hister semipunctatus Fabricius, 1792
= Saprinus chobauti Auzat, 1926
Germany: Baden-Württemberg.
Southern Europe, Mediterranean subregion, Portugal (including the Azores Archipelago), Cape Verde Islands, Central Asia, introduced to Peru (
This species is found frequently on carrion, with a preference for larger carcasses (e.g., those of dogs, sheep, cattle, camels, etc. – T. Lackner pers. observ.). According to
This species has another subspecies S. (S.) caerulescens punctisternus Lewis, 1900 that occurs in China, Mongolia, Korean peninsula and Russian Far East (
= Saprinus angoranus Bickhardt, 1911
Spain.
Mediterranean subregion, Arabian Peninsula, Central Asia (
Saprinus calatravensis is found on small and medium-sized carrion. According to
Hister chalcites Illiger, 1807
= Hister affinis Paykull, 1811
= Saprinus aerosus Normand & Thérond, 1952
= Saprinus melanocephalus Normand & Thérond, 1952
= Saprinus prolongatus Normand & Thérond, 1952
= Saprinus scapularis Normand & Thérond, 1952
Portugal.
Mediterranean subregion, Africa, Arabian Peninsula, Central Asia, India, Burma, Australia (
Saprinus chalcites is a widespread flying predator found on carcasses, rotting vegetable substances, as well as in dung. According to
Hister godet Brullé, 1832
= Saprinus bitterensis Marseul, 1862
= Saprinus godetii Marseul, 1857 [emendation]
= Saprinus pseudolautus Reitter, 1904
Greece: Peloponnesus.
Spain, Portugal, southern France, Turkey, Georgia, Kazakhstan, Turkmenistan, Saudi Arabia (
Found on carrion (
Hister maculatus P. Rossi, 1792
= Saprinus obscuripennis J. Müller, 1899
= Hister personatus Fischer von Waldheim, 1823
= Saprinus pseudocruciata Auzat, 1920
Italy: Etrusca.
Slovakia, Hungary, south Europe, Turkey, Georgia, Azerbaijan, Cyprus, Iraq, Iran, Turkmenistan, Kazakhstan, Afghanistan (
Iran.
Turkey, Syria, Iraq, Azerbaijan, Lebanon (
Reichardt (1941) reports this species from Azerbaijan as an inhabitant of mountain steppes (500 m); several specimens were also collected by pitfall trapping. In our samples, we collected two specimens from a human corpse that was in the active decay stage (Fig.
= Saprinus lindrothi Dahlgren, 1968
Syria.
Italy: Sardinia, Greece: Crete, Bulgaria, Turkey, Armenia, Jordan (
Saprinus prasinus has another subspecies S. (S.) prasinus aeneomicans G. Müller, 1960 that occurs in neighbouring Israel and Syria (
= Saprinus vermiculatus Dahlgren, 1964
Turkey: Ankara.
According to
Ethiopia.
Insufficiently known, according to our observations it is similar to its congeners. In our samples, we collected 149 specimens mainly during the active decay stage and several specimens in other stages of decomposition (Table
Syria.
Algeria, Turkey, Israel, Armenia, Azerbaijan, Afghanistan, Iran (
According to
= Saprinus fagniezi Auzat, 1912
= Saprinus lecontei Casey, 1916
= Saprinus meridionalis Ihssen, 1949
= Saprinus simulans J. Sahlberg, 1913
Hungary, Romania.
Central and south Europe, North Africa, Spain (including Canary Islands), Portugal (including Madeira), Turkey, central Asia, introduced to north America (
A typical free-living volant predator found on carrion as well as in dung. In our samples we collected 242 specimens. Saprinus (S.) subnitescens was the most abundant species on the carcasses during both the active and advanced stage of decomposition.
= Saprinus brunnensis A. Fleischer, 1883
Uzbekistan.
Central and South Europe, Central Asia, Mongolia, North China (
This species is found predominantly on carcasses. During our sampling eight specimens were collected.
16Saprinus (S.) calatravensis Fuente, 1899 male genitalia: 8th sternite and tergite, ventral view 17 8th sternite and tergite, dorsal view 18 8th sternite and tergite, lateral view 19 9th + 10th tergites, dorsal view 20 9th + 10th tergites, lateral view 21 spiculum gastrale (9th sternite), ventral view 22 spiculum gastrale (9th sternite), lateral view 23 aedeagus, dorsal view 24 aedeagus, lateral view.
26Saprinus (S.) chalcites (Illiger, 1807) male genitalia: 8th sternite and tergite, ventral view 27 8th sternite and tergite, dorsal view 28 8th sternite and tergite, lateral view 29 9th + 10th tergites, dorsal view 30 9th + 10th tergites, lateral view 31 spiculum gastrale (9th sternite), ventral view 32 spiculum gastrale (9th sternite), lateral view 33 aedeagus, dorsal view 34 aedeagus, lateral view.
35Saprinus (S.) godet (Brullé, 1832) male genitalia: 8th sternite and tergite, ventral view 36 8th sternite and tergite, dorsal view 37 8th sternite and tergite, lateral view 38 9th + 10th tergites, dorsal view 39 9th + 10th tergites, lateral view 40 spiculum gastrale (9th sternite), ventral view 41 spiculum gastrale (9th sternite), lateral view 42 aedeagus, dorsal view 43 aedeagus, lateral view.
44Saprinus (S.) robustus Krása, 1944 male genitalia: 8th sternite and tergite, ventral view 45 8th sternite and tergite, dorsal view 46 8th sternite and tergite, lateral view 47 9th+10th tergites, dorsal view 48 9th+10th tergites, lateral view 49 spiculum gastrale (9th sternite), ventral view 50 spiculum gastrale (9th sternite), lateral view 51 aedeagus, dorsal view 52 aedeagus, lateral view.
54Saprinus (S.) submarginatus J. Sahlberg, 1913 male genitalia: 8th sternite and tergite, ventral view 55 8th sternite and tergite, dorsal view 56 8th sternite and tergite, lateral view 57 9th+10th tergites, dorsal view 58 9th+10th tergites, lateral view 59 spiculum gastrale (9th sternite), ventral view 60 spiculum gastrale (9th sternite), lateral view 61 aedeagus, dorsal view 62 aedeagus, lateral view.
64Saprinus (S.) subnitescens Bickhardt, 1909 male genitalia: 8th sternite and tergite, ventral view 65 8th sternite and tergite, dorsal view 66 8th sternite and tergite, lateral view 67 9th + 10th tergites, dorsal view 68 9th + 10th tergites, lateral view 69 spiculum gastrale (9th sternite), ventral view 70 spiculum gastrale (9th sternite), lateral view 71 aedeagus, dorsal view 72 aedeagus, lateral view.
73Saprinus (S.) tenuistrius sparsutus Solsky, 1876 male genitalia: 8th sternite and tergite, ventral view 74 8th sternite and tergite, dorsal view 75 8th sternite and tergite, lateral view 76 9th + 10th tergites, dorsal view 77 9th + 10th tergites, lateral view 78 spiculum gastrale (9th sternite), ventral view 79 spiculum gastrale (9th sternite), lateral view 80 aedeagus, dorsal view 81 aedeagus, lateral view.
S. (S.) tenuistrius has another subspecies, the nominotypical S. (S.) tenuistrius tenuistrius Marseul, 1855, which is known from Egypt, north Africa, Ethiopia and Arabian Peninsula (
Saprinus (S.) subnitescens was collected during the sampling of the months March, April, May, June, July, August, and September. It was collected during three stages of decomposition; bloat, active decay and advanced decay. The maximum amount of specimens was observed during the advanced decay stage of the carcasses. This species was dominant (83 specimens) in Naas (Bikfaya) during spring season at the mean temperature of 16.4 °C. Specimens were collected during both active decay and advanced stages of decomposition from Naas, Badghan, and Fanar, and less frequently from Deir El-Ahmar and Hasbaya, respectively. Moreover, this species was also collected during the bloat stage of decomposition from Badghan (Fig.
Saprinus (S.) strigil was likewise collected during the sampling events of the months March, April, June, July, August, and September, with the maximum of 104 specimens collected in August from Hasbaya, when the mean temperature was 26.9 °C. The specimens were collected mainly during the active decay stage, less frequent during the advanced decay and minimally present during the bloat stages. Unlike Saprinus (S.) subnitescens, this species was rarely present in Naas. A few specimens were collected during the bloat stage from Badghan (Fig.
Saprinus (S.) chalcites was collected during the sampling events of the months of April, June, July, August, September and October. In general, it was dominant during the active decay stage of the carcasses. Peak abundance of 76 specimens was collected in August from Hasbaya during bloat, active decay and advanced decay stages of pig carcass decomposition. Similar amounts were collected during the active and advanced decay stages from Badghan. More specimens were collected during active decay stage than during the advanced decay stage from Deir El-Ahmar. S. (S.) chalcites was present in Naas only during the advanced decay stage (Fig.
Saprinus (S.) caerulescens caerulescens was captured in March, April, June, July and August; a maximum of 17 specimens was collected during the spring season from Fanar, when the mean weather temperature was 17.6 °C. In general, more specimens were collected during the active decay stages than those collected during the advanced decay ones. Similar amounts were collected during those stages from Badghan. It was collected during the decay stage mainly from Deir El-Ahmar and less frequently from Hasbaya. Regarding the locality Naas, this species was there found to be present mainly during the advanced stage of decomposition and less frequent during the active decay stage (Fig.
Saprinus (S.) prasinus prasinus – the peak of its abundance of 27 specimens was likewise in Fanar during spring (March and April). They were mainly captured during the active decay stage and found to be less numerous in the advanced decay stage. This species was likewise collected from Naas during same season during the advanced decay stage only (Fig.
Saprinus (S.) godet – the peak abundance of this species was in spring in Fanar. It is mainly present during the active decay stage of carcasses decomposition and in lesser numbers during the advanced decay stage. More specimens were collected during the active decay stage than from the advanced decay stage from Fanar and Badghan. It was only present during the active decay stage in Deir El-Ahmar and during the advanced decay stage in Hasbaya (Fig.
Both Saprinus (S.) tenuistrius sparsutus and Saprinus (S.) calatravensis were collected in July – August. The former was only collected during the active decay stages of Deir El-Ahmar and Hasbaya. It was more abundant in the locality Deir El-Ahmar in July; the mean temperature was 29.7 °C. The latter was mainly present during the active decay stage and less frequently during the bloat and the advanced decay stages (Figs
Saprinus (S.) robustus was collected from Fanar in March during the active decay stage only, and from Deir El-Ahmar in July during both the active and advanced decay stages. Only a single specimen of Saprinus (S.) submarginatus and of Saprinus (S.) maculatus, respectively, was collected during the summer season from one locality (Badghan); the mean temperature was 20 °C. A male and a female specimen of Saprinus (S.) magnoguttatus were collected from a human corpse that was infested by Diptera larvae (Fig.
This key includes only those species specifically recorded from Lebanon so far. For the Saprinus species found in the neighbouring countries (Israel, Palestine, Syria and Jordan), which are likely to occur also in Lebanon (see
1(4) | Elytra bicolored (Fig. |
2 |
2(3) | At least the entire lateral elytral margin orange-red, usually most part of the elytral disk orange-red with only a short band along the elytral suture black (Fig. |
Saprinus (S.) maculatus (P. Rossi, 1790) |
3(2) | Each elytron with a well-defined red macula, never occupying the entire lateral elytral margin (Fig. |
Saprinus (S.) magnoguttatus Reichardt, 1926 |
4(1) | Elytra unicolored, never with red macula (Fig. |
5 |
5(6) | Pronotal hypomeron setose, fourth dorsal elytral stria strongly reduced, often absent. A large, usually metallic species (PEL = 5.00–7.50 mm) (Fig. |
Saprinus (S.) caerulescens caerulescens (Hoffman, 1803) |
6(5) | Pronotal hypomeron asetose, fourth dorsal elytral stria usually not reduced, fully developed. Smaller species (PEL = 2.50–6.50 mm) | 7 |
7(8) | Elytra, especially on their apical halves with very dense punctation, punctures aciculate and striolate, elytral intervals punctured, third dorsal elytral stria well-developed (Fig. |
Saprinus (S.) strigil Marseul, 1855 |
8(7) | Elytra with variously dense punctation, but punctures usually not aciculate or striolate (some specimens of S. (S.) robustus can have striolate punctures, but then the third dorsal elytral stria is always strongly reduced) (Fig. |
9 |
9(12) | Elytra with well-defined polished areas ‘mirrors’, punctation of elytral disk very dense, punctures separated by less than their own diameter, third dorsal elytral stria reduced to absent (Fig. |
10 |
10(11) | Elytral ‘mirror’ with microscopic scattered punctation, light to dark brown species, without greenish or bronze metallic hue, third dorsal elytral stria reduced, but usually discernible; elytral punctation in fourth elytral interval reaches elytral half (Fig. |
Saprinus (S.) aegialius Reitter, 1884 |
11(10) | Elytral ‘mirror’ glabrous, third dorsal elytral stria usually strongly reduced to absent, dorsum with distinct greenish or bronze metallic hue; punctation in fourth elytral interval does not reach elytral half (Fig. |
Saprinus (S.) prasinus prasinus Erichson, 1834 |
12(9) | Elytra without well-defined polished areas (‘mirrors’), punctation of the elytral disk less dense, punctures usually separated by their own diameter or more (Fig. |
13 |
13(16) | Apices of carinal prosternal striae strongly divergent, laying on lateral sides of the prosternal process (Fig. |
14 |
14(15) | Pronotal depressions deep, third dorsal elytral stria usually not reduced, light to dark brown species with slight bronze metallic hue (Fig. |
Saprinus (S.) subnitescens Bickhardt, 1909 |
15(14) | Pronotal depressions shallow, third dorsal elytral stria usually strongly reduced, black species without metallic hue (Fig. |
Saprinus (S.) robustus Krása, 1944 |
16(13) | Apices of prosternal striae divergent, but never laying on lateral sides of the pronotal process (Fig. |
17 |
17(18) | Pronotal depressions absent, pronotal disk medially with distinct punctation, humeral elytral stria confluent with inner subhumeral one creating a supplementary dorsal elytral stria parallel to first (Fig. |
Saprinus (S.) godet (Brullé, 1832) |
18(17) | Pronotal depressions present, pronotal disk medially with only scattered fine punctation (Fig. |
19 |
19(20) | Entire elytral disk with punctation, punctures separated by twice or more their diameter, dorsal elytral striae thin, impunctate (Fig. |
Saprinus (S.) tenuistrius sparsutus Solsky, 1876 |
20(19) | At least the area between united sutural and fourth elytral striae without punctation (or punctures microscopic), punctures of elytral disk separated usually by less than twice their diameter (Fig. |
21 |
21(22) | Apical margin of metaventrite of male without tubercles. Male terminalia: 8th sternite with two rows of brush-like setae: one situated approximately medially and another apically, aedeagus constricted before apex; apex rounded (Figs |
Saprinus (S.) submarginatus J. Sahlberg, 1913 |
22(21) | Apical margin of metaventrite of male with two distinct tubercles (Fig. |
23 |
23(24) | Tubercles on the apical margin of metaventrite of male slightly removed from metaventral margin (Fig. |
Saprinus (S.) calatravensis Fuente, 1899 |
24(23) | Tubercles situated almost on the very apical metaventral margin (Fig. |
Saprinus (S.) chalcites (Illiger, 1807) |
In an experiment performed in Al-Baha Province (Kingdom of Saudi Arabia) Saprinus species were commonly found on rabbit carcasses during spring, summer, and autumn. In spring, they were found to be abundant during the active decay and the first two days of the dry decay stages. However, in autumn they were common ranging from the bloat stage to the dry decay stages (
In our survey, Saprinus (S.) caerulescens, Saprinus (S.) godet, Saprinus (S.) prasinus, and Saprinus (S.) robustus were attracted to cadavers during the both active decay and advanced decay stages of decomposition, when the Diptera larvae are active and feeding on the carcasses, and were also recovered from the soil after the departure of Diptera larvae to pupate. Saprinus (S.) strigil, Saprinus (S.) subnitescens, Saprinus (S.)calatravensis, and Saprinus (S.) chalcites were present on the carcasses earlier, in the bloat stage in the summer season, whereas in the spring months (as observed in Fanar and Naas) the arrival of Saprinus on the carcass was during the active decay stage (Figs
It is possible that different patterns of residency on carrion in Saprinus result from the differences in release patterns of the volatile organic compounds (VOCs) attracting them (
In the present study, 489 specimens of Saprinus were recorded, representing 13 species. Out of these, 8 are new records for the Lebanese fauna (Fig.
The first author would like to thank the CNRS (National Council of Scientific Research-Lebanon) and Walid Joumblatt Foundation for University studies for their financial support. This paper is a contribution of the team project “Biodiversity: Origin, Structure, Evolution and Geology” and to the activity of the laboratory “Advanced Micropalaeontology, Biodiversity and Evolution Researches (AMBER)” at the Lebanese University, the support of taxonomists and to the Institut de Systématique, Évolution, Biodiversité in the Muséum National d’Histoire Naturelle, Paris, France and the collaboration with the Bavarian State Collection of Zoology Munich, Germany. T. Lackner would like to thank the Alexander von Humboldt Foundation (Bonn, Germany), itself part of the Federal Ministry for Education and Research (Berlin, Germany), which provided funding for this study to T. Lackner. The reviewers Alexey K. Tishechkin (Sacramento, USA) and Michael S. Caterino (Clemson, USA) are thanked for their meticulous corrections and numerous suggestions that led to a greatly improved manuscript.