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Review of the Observations of Aggregates of Steninae Repor- Ted Since 1856 (Coleoptera Staphylinidae)

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Review of the Observations of Aggregates of Steninae Repor- Ted Since 1856 (Coleoptera Staphylinidae) Biodiversity Journal, 2017, 8 (1): 123–144 MONOGRAPH Review of the observations of aggregates of Steninae repor- ted since 1856 (Coleoptera Staphylinidae) Giulio Cuccodoro Muséum d’histoire naturelle, C. P. 6434, CH-1211 Genève 6, Switzerland; e-mail: [email protected] ABSTRACT The nine aggregates of Steninae documented since 1856 are reviewed, completed with seventeen new reports. All the pictures available on the topic are presented. Considering the ubiquity and megadiversity of the subfamily, this total of 26 observations reported over the last 150 years appears strikingly low, clearly indicating that the phenomenom is exceptional. These observations were all made in the Palaearctic and Oriental realms, and refer to both the extant genera of Steninae Dianous and Stenus. Six observations refer to Dianous species, all from China, mainly D. banghaasi and D. freyi. Of the twenty observations referring to Stenus ten were made in the Mediterranean area, mainly on S. cordatus, S. elegans and S. turk. This collection of observations seems a composite of several kinds of behavioural patterns, such as hibernation, aestivation, reproductive swarming, and possibly hilltopping, with some most likely intermixed. Aggregation in compact multilayer masses of individuals as well as occasional recurrence of Stenus aggretates at precise locations over days or year are probably controlled by pheromone signals. The reasons driving these slender, one centi- meter long rove beetles to occasionally swarm in compact masses of well over hundred thou- sand individuals remain nevertheless as enygmatic as how such a spectacular phenomenom performed by members of the second most diverse genus of animals on earth can remain so rarely observed. KEY WORDS Ethology; aggregation; behaviour; reproduction; hilltopping; Mediterranean Region. Received 16.01.2017; accepted 04.03.2017; printed 30.03.2017 Proceedings of the 3rd International Congress “Biodiversity, Mediterranean, Society”, September 4th-6th 2015, Noto- Vendicari (Italy) INTRODUCTION capture mouth apparatus unique within insects, which is probably the key innovation responsible Steninae are very distinctive slender rove beetles for the impressive radiation of the group (Betz, with broad globular compound eyes. The subfamily 1996, 1998). Partly because of the marked attrac- is distributed virtually worlwide except Antarctica tion of their favoured preys for humid microhabitas, and New Zealand. It consists of some 2750 extant Steninae are mainly found in moist forest leaf litter, species grouped in the genera Dianous Leach, 1819 near swamps, along river banks or lake shores, but and Stenus Latreille, 1779, the latter being the also walking on the foliage of tropical rainforests. second most diverse genus of animals with approx- In Europe their greatest diversity is to find in moist imately 2550 species (Herman, 2001; Thayer, 2005; environments such as reeds, where some species Cai et al., 2014). Feeding essentially on collembola forage on humus or plants debris near the ground, Stenus possess a chameleon-like protrusive prey- while others prefer to climb on plants, or are surface 124 GIULIO CUCCODORO runners on bare ground (Betz, 1999a, 1999b). in the rubric Von Kollege zu Kollege on the recto Although they usually possess fully functional of the backcover of volumes 73 (1977) to 77 (1981) wings, they all are very reluctant to flight, and move of the journal Entomologishe Blätter. Several un- almost only walking or running. Hence another of published observations were also forwarded to me their evolutionary traits appears to be a trend by the other specialists of Steninae Guillaume de towards significant widening of tarsi, resulting in Rougemont, London, and Liang Tang, Shanghai. the apparently repeated emergence of water-gliding The rest was addressed to me in feedback to my and swimming habilities (Betz, 2002). presentation on the topic at the “22nd International In 2005 I was casually confronted to another Meeting on Biology and Systematics of Staphylin- striking pecularity of Steninae unique within rove- idae” in Stuttgart (Cuccodoro, 2007) and to my call beetles, which is their ability to occasionally con- for informations addressed in June 2011 to all the gregate in very large numbers and form dense email list of the announcement of 26th edition of multilayer masses of individuals (Cuccodoro, 2007; the same meeting, or gathered by myself over the Puthz, 2008). Submitting my few pictures and field internet. When still possible I also contacted the au- notes to the Stenus specialist Volker Puthz, I was thors of previously published reports for additional very suprized to learn that such an impressive phen- informations. onenom was still documented in the literature with only a handfull of short notes since first reported some 150 years ago from France (Dufour & Perez, RESULTS 1857). Here I review the nine observations of aggreg- The observations of aggregates of Steninae are ates of Steninae recorded hitherto in the litterature listed below organized chronologically, and con- (Dufour & Perez, 1857; Rougemont, 1980; Abdul- sequently numbered. Nour & Kallab, 1989; Lecoq, 1991, 1993; Cooter, 1997; Puthz, 2000; Zhao & Li, 2004; Cuccodoro, 1) September 1856, Saint Sever (France) 2007), as well as sixteen additional observations reported to me by colleagues, or gathered over The first observation of an aggregate of Sten- the internet. All the pictures available to me are inae was made near Saint Sever (Landes France) presented, with some pertaining to previous in the beginning of September by a local teacher records published here for the first time. The struc- named Perez (first name unknown). He brought ture of these observations as well as possible causes back to Léon Dufour for identification a sample and adaptative interests of the phenomenom are containing two thousands and several hundreds of briefly discussed. Hopefully this will stimulate in- individuals of Stenus rusticus Erichson, 1840 (= vestigations on this phenonomenom, which is def- Stenus picipes Stephens, 1833), which were only a initely as spectacular as it remains enygmatic. fraction of those forming the aggregate he wit- nessed. Curious to see the phenomenom by himself L. Dufour went there some days later with M. MATERIAL AND METHODS Perez, and they could observe it again exactly at the same place. It was on leaves of a chestnut Most unpublished observations were forwarded stump cut at level with the ground located on the to me by Volker Puthz, Schlitz. He obtained them edge of a small, dry ravine. The leaves were com- mainly in feedback to his repeated call for informa- pletely blackened by the beetles piled one on each tions on the topic “Stenus-Ansammlungen im Mit- other. He estimated that they were more that twenty telmeerraum: wer hat solche Massenvorkommen thousands individuals within an area of 50 cm in von Stenus (Staphylinidae) auf einer kleine Fläche diameter. Dufour shaked the leaves and heard “like (unter Stein, o. ä.) beobachtet? Mitteilung Dr. V. a rain of sand”, so he put his hand underneath, Puthz (….)” [Stenus - aggregations in the Mediter- which was readily filled with thousands of Stenus. ranean area: who observed such mass occurences These were etherized, put into a glass vial and of Stenus (Staphylinidae) on a small surface (under given to his friend Joseph Alexandre Laboulbène, stone, or similar)? Contact Dr. V. Puthz] published with charge to present that sample together with a Review of the observations of aggregates of Steninae reported since 1856 (Coleoptera Staphylinidae) 125 report of the observation at the Entomological So- was experienced during the process. This sample ciety of France in Paris. It was finally Léon Fair- totalized 1,217 specimens of Stenus turk Puthz, maire, in the name of Laboulbène, who read the 1972 with a sex ration of 36 females for 64 males note of Dufour at the 22 October 1956 meeting of in a random sample of 100 individuals (det. L. the Entomological Society of France (Dufour & Hromádka). Additional masses were found under Perez, 1857). some other stones, thought only those located at a Note. The note is referred to as “Stenus rusticus few steps of the first one, as well as in a few shad- rencontré en immense quantité (note sur le), L. owed and wet places at base of the cliff. Reported Dufour et Perez” in the table of contents (alpha- to V. Puthz by J. Král in August 1976 (V. Puthz betical by taxa) at page CLXIV of tome IV of the pers. comm.). Annales de la Société Entomologique de France (3rd series), in the section containing the reports 4) 29 September 1972, Kreta (Greece) the meetings of this society in the last trimester of 1856; as the last meeting reported was on 24 The aggregate was observed by Hans Malicky December 1856, it is very unlikely that the volume in the Yeropotamos river valley near the bridge was published before the end of that year, hence below Phaistos palace (Kreta, Greece). It occurred my quotation of the reference not following Lecoq in the late afternoon of a sunny day in a place (1993), who credited the publication of the note to already at shade, and consisted of more than 10,000 Fairmaire in 1856. individuals of Stenus picipes (det. V. Puthz) ag- gregated in compact clusters hanging on a few 2) 12 June 1972, Rosas (Spain) - Figs. 1, 2 blades of grasses near the river shore, from far look- ing like bee swarms. There were no stones in the The aggregate was observed by Alfred Elbert in area. Reported to V. Puthz by H. Malicky in August montains West of Rosas (Catalogna, Spain). It oc- 1977 (V. Puthz pers. comm.). curred under a large stone, and consisted of several thousands individuals of Stenus sp. massed in a few 5) 3 July 1977, Trimiklini (Cyprus) multilayer masses (Figs. 1, 2). Reported to V. Puthz by A. Elbert in January 1973 (V. Puthz pers. The aggregate was observed by Stanislav Vit comm.).
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