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Meriympäristöjen Hyönteiset Meriympäristöjen hyönteiset Aurelia Suvitie Pro gradu -tutkielma Akvaattiset tieteet Bio- ja ympäristötieteellinen tiedekunta Helsingin yliopisto Lokakuu 2020 Tiedekunta Koulutusohjelma Bio- ja ympäristötieteellinen tiedekunta Akvaattisten tieteiden koulutusohjelma Tekijä Aurelia Suvitie Työn nimi Meriympäristöjen hyönteiset Oppiaine Akvaattiset tieteet Työn laji Aika Sivumäärä Pro gradu -tutkielma Lokakuu 2020 95 +15 Tiivistelmä Hyönteiset (Insecta) ovat tunnetulta lajistoltaan maapallon runsaslukuisin luokka. Valtavan monimuotoisuuden ansiosta hyönteisiä löytyy lähes kaikenlaisista ympäristöistä. Yli 1 000 000 tunnetusta hyönteislajista n. 90 % on kuitenkin terrestrisiä. Vaikka meret kattavat maapallon pinta-alasta yli 70 %, mereisiä hyönteisiä tunnetaan alle 2 000 lajia. Tämän työn tarkoitus on antaa mahdollisimman kokonaisvaltainen käsitys mereisestä hyönteislajistosta, sekä tekijöistä, jotka ovat aiheuttaneet hyönteisten vähäisyyden merissä. Vastaavaa monitahoista katsausta aiheeseen ei ole aiemmin tehty. Katsauksen ensimmäinen tavoite on määritellä ja listata hyönteistaksonit, jotka sisältävät mereisiä lajeja, sekä tarkastella näiden lajien habitaattivalintoja ja elintapoja. Tässä tarkoituksessa mereisten hyönteisten lajikirjo ja elintavat on kartoitettu World Register of Marine Species -tietokannan (WoRMS), sekä mereisiä hyönteisiä käsittelevien yleisteosten, katsausartikkeleiden ja tutkimusartikkeleiden avulla. Pääasiassa WoRMS-tietokannan perusteella mereisiksi on määritelty 1 318 hyönteislajia kymmenestä lahkosta. Näiden lisäksi seitsemästä lahkosta tavataan tuntematon määrä mereisiä lajeja. Lajiston lahkokohtaisen tarkastelun lisäksi työssä käydään läpi mereisten hyönteisten asuttamat elinympäristöt litoraalissa ja pelagiaalissa, ja millaisia sopeumia näissä ympäristöissä eläminen hyönteisiltä vaatii. Katsauksen toinen tavoite on käsitellä mereisten hyönteisten vähäisyyttä ilmiönä. Aiheen pohjustamiseksi lukijalle selvitetään työn alussa miksi ja miten hyönteisten valtava lajirikkaus on syntynyt. Työn loppupuolella siirrytään tarkastelemaan tekijöitä, jotka ovat estäneet vastaavan rikastumisen merissä, kuten osmoottiseen säätelyyn ja hengitysjärjestelmiin liittyvät ongelmat, hyönteisten lisääntymisen ja elinkierron ominaispiirteet, sekä yksinkertaisesti se, että siirtymät fysiologisesti erilaisten habitaattien välillä ovat harvinaisia. Tätä varten käydään läpi myös hyönteisten kehityshistoria, jonka tuloksena lahkon levittäytymistä meriin haittaavat ominaisuudet ovat syntyneet. Pohjimmiltaan mereisten hyönteislajien vähäisyys johtuu siitä, että hyönteiset ovat Pancrustacea-kladin maalle suuntautunut kehityslinja. Vaikka mereisyys on lahkossa poikkeuksellista, ei se ole läheskään ennenkuulumatonta. Mereisiä lajeja on syntynyt monessa hyönteisten kehityslinjassa, ja nämä lajit sopeutuvat ympäristöönsä lukuisin eri tavoin. Hyönteisten levittäytymistä meriin koskevien jatkoselvitysten ja analyysien kannalta mereiset hyönteislajit, sekä niiden elintavat ja sopeumat meriympäristöihin tulisi kartoittaa vielä yksityiskohtaisemmin. Tämä katsaus tarjoaa toivottavasti tulevaisuudessa perustan tarkemmille selvityksille mereisestä hyönteislajistosta, sekä mereisten hyönteisten vähäisyyttä selittävistä tekijöistä. Avainsanat Hyönteiset, Insecta, monimuotoisuus, evoluutio, Halobates, sopeumat, Pancrustacea Ohjaajat Jaanika Blomster, Lauri Kaila Säilytyspaikka Helsingin yliopiston digitaalinen arkisto HELDA Muita tietoja Tiivistelmä suomeksi ja englanniksi Faculty Degree Programme Faculty of Biological and Environmental Sciences Aquatic Sciences Author Aurelia Suvitie Title Insects of marine environments Subject Aquatic sciences Level Month and year Number of pages Master’s degree thesis October 2020 95 +15 Abstract Insects (Insecta) are the most species-rich class of organisms on earth. Due to their vast biodiversity, insects are found in almost every environment. However, approximately 90 % of the over 1 000 000 insect species described are terrestrial. The ocean covers over 70 % of the earths surface, but less than 2 000 marine insect species are known. The purpose of this thesis is to give the reader a comprehensive understanding of marine insect biodiversity and what has caused its sparceness in the seas. Such a multi-faceted review of the topic has not previously been written. The first aim of this review is to catalogue all insect taxons containing marine species, and to examine the habitat choices and ecology of these species. For this purpose, marine insect species have been researched using the World Register of Marine Species (WoRMS) and reference literature, as well as reviews and other articles. Based mostly on WoRMS, the number of marine insect species is determined to be 1 318, divided between 10 insect orders. In addition, seven insect orders are noted to include an unknown number of marine species. As well as examining marine insect biodiversity order-by-order, this work considers the littoral and pelagic habitats in which marine insects live, and the adaptations that are required of insects living in these environments. The second aim of this review is to investigate the lack of insects in marine environments as a phenomenon. The groundwork is laid at the beginning of the review with an examination of why and how insects have diversified so successfully. Later on, possible factors to prevent a similarily successful diversification in marine environments are presented, such as issues with osmoregulation and breathing systems, the typical features of insect reproduction and life cycles, and the fact that transitions between physically contrasting habitats are rare. Pertaining to this the evolutionary history of insects, which has produced the aforementioned features impeding radiation to the seas is also explored. Ultimately insect species in the oceans are few, because insects are a terrestrial evolutionary branch of Pancrustacea. Even though marine habits are uncommon among insects, they are not unheard of. Marine species have emerged in several insect clades and these species have adapted to their environment in numerous ways. For the purposes of further analyses and research on the radiation and biodiversity of marine insects, further surveying of the species and their habits and adaptations is recommended. This work will hopefully offer a point of reference for more detailed future studies and reviews of marine insect biodiversity and the factors that have led to the near-exclusion of insects from the seas. Keywords Insects, Insecta, biodiversity, evolution, Halobates, adaptations, Pancrustacea Supervisors Jaanika Blomster, Lauri Kaila Where deposited Digital Repository of the University of Helsinki HELDA Additional information Abstract in Finnish and English Sisällysluettelo 1 Johdanto 1 1.1 Työn tarkoitus ......................................................................................................... 2 1.2 Menetelmät ja kirjallisuus ....................................................................................... 3 2 Miksi hyönteisiä on niin paljon? 4 2.1 Proksimaattiset tekijät ............................................................................................ 6 2.1.1 Kladin ikä .......................................................................................................... 6 2.1.2 Monimuotoistumisnopeus............................................................................... 7 2.2 Ultimaattiset tekijät ................................................................................................ 8 2.2.1 Morfologiset ominaisuudet ............................................................................. 8 2.2.1.1 Siivet.......................................................................................................... 8 2.2.1.2 Täydellinen muodonvaihdos .................................................................. 10 2.2.1.3 Muut morfologiset ominaisuudet .......................................................... 11 2.2.2 Lajienväliset ja lajinsisäiset vuorovaikutukset ............................................... 12 3 Mereisen hyönteisen ja mereisen habitaatin määrittely 14 3.1 Yleisesti.................................................................................................................. 14 3.2 Mereisen hyönteisen määrittely tässä työssä ...................................................... 16 3.2.1 Pääsääntö ....................................................................................................... 16 3.2.2 Poikkeukset .................................................................................................... 18 4 Mereiset hyönteistaksonit 21 4.1 Siimahäntäiset (Archaeognatha) ........................................................................... 23 4.2 Päivänkorennot (Ephemeroptera) ........................................................................ 24 4.3 Sudenkorennot (Odonata) .................................................................................... 25 4.4 Suorasiipiset (Orthoptera)..................................................................................... 26 4.5 Kummitussirkat (Phasmida) .................................................................................. 27 4.6 Pihtihäntäiset (Dermaptera) ................................................................................. 27 4.7 Torakat ja termiitit (ent. Isoptera ja Blattaria, nyk. Blattodea) ............................ 28 4.8 Jäytiäiset (Psocoptera) .........................................................................................
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