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Department of Ecology and Evolutionary Biology University of Kansas Lawrence, KS 66045, U.S.A The Coleopterists Bulletin, 71(2): 315–328. 2017. THE MORPHOLOGY AND BEHAVIOR OF THE ENDEMIC MALAGASY WHIRLIGIG BEETLE HETEROGYRUS MILLOTI LEGROS, 1953 (COLEOPTERA:GYRINIDAE:HETEROGYRINAE) GREY T. GUSTAFSON Department of Ecology and Evolutionary Biology University of Kansas Lawrence, KS 66045, U.S.A. [email protected] JOHANNES BERGSTEN Department of Zoology Swedish Museum of Natural History Stockholm, SWEDEN TOLOTRA RANARILALATIANA,JACQUELIN HERISAHALA RANDRIAMIHAJA Departement d’Entomologie, Facult´e des Sciences B. P. 906, Universit´ed’Antananarivo Antananarivo, MADAGASCAR AND KELLY B. MILLER Department of Biology and Museum of Southwestern Biology University of New Mexico Albuquerque, NM 87131, U.S.A. ABSTRACT The Malagasy endemic whirligig beetle Heterogyrus milloti Legros, 1953 is redescribed. Jumping behavior of H. milloti is reported here for the first time with video recordings provided. Results of a behavioral experiment conducted in the field demonstrate H. milloti jumps in a targeted manner in a downstream direction. The unique habitat of H. milloti is described in detail with both image and video of the habitat included. Morphology of H. milloti is discussed in detail, revealing symplesiomorphies with Spanglerogyrus Folkerts, 1979, characters forming transitional series between Spanglerogyrus and the Gyrininae, and features unique to H. milloti. The potential adaptive significance of these peculiar morphological features in association with the habitat of H. milloti is discussed. Finally, an argument for the necessity of conservation of this species is made, and common names in English, French, Malagasy, and Swedish for H. milloti are proposed to aid conservation efforts. Key Words: Hydradephaga, aquatic beetles, Madagascar, conservation, jumping DOI.org/10.1649/0010-065X-71.2.315 Heterogyrus milloti Legros, 1953 is a little known galea. Brinck (1955) was next to treat Heterogyrus. but extraordinary whirligig beetle with a striking Noting the numerous unique characteristics of the yellow and black striped appearance (Fig. 1A). The genus, he decided to place it in a new monotypic species was originally described in 1953 by Legros, tribe, but he also recognized several features uniting after he received a batch of Malagasy Hydradephaga it with the Dineutini Desmarest (then Enhydrinae from Renaud Paulian (then Deputy Director of R´egimbart) and the Orectochilini R´egimbart (then l’Institut de Recherche Scientifique de Madagascar), Orectochilinae). Brinck (1955) provided a short containing four specimens (one male and three fe- description of the species in English as well as an males) of a distinctly new taxon (Legros 1953). The improved illustration of the aedeagus of the type original description was in French with several il- specimen. lustrations of some of the more interesting mor- Following discovery of Spanglerogyrus albi- phological features (Legros 1953). Legros (1953) ventris Folkerts, 1979, the classification of the compared Heterogyrus Legros to the known genera Gyrinidae changed considerably, as Folkerts (1979) of the Gyrinini Latreille (then the Gyrininae), placing placed his new genus into the newly erected it within this group due to the presence of maxillary monotypic subfamily Spanglerogyrinae Folkerts 315 316 THE COLEOPTERISTS BULLETIN 71(2), 2017 Fig. 1. Heterogyrus milloti. A) Dorsal habitus of male, scale bar 5 2 mm, B) Ventral habitus, lw 5 lateral wing of metaventrite, ma 5 metanepisternum, Roman numerals indicate abdominal sternites. and lowered all the former subfamilies to tribal rank more derived Gyrininae, supporting its position in the within the nominotypical subfamily Gyrininae, as phylogeny (Miller and Bergsten 2012). Miller and the species possessed numerous seemingly plesio- Bergsten (2012) were also the first to report briefly on morphic conditions relative to all other Gyrinidae. the unusual habitat of Heterogyrus relative to that of Spanglerogyrus Folkerts was indeed recovered as other Gyrinidae. sister to all other Gyrinidae in the first phylogenetic While collecting H. milloti during a 2014 expe- analysis of the family including molecular data dition to Madagascar, it was observed that live (Miller and Bergsten 2012), supporting its place- specimens jumped about in the aquatic net after ment in its own subfamily, and the plesiomorphic being removed from water. Furthermore, specimens nature of its morphology. Surprisingly, Miller and placed in white plastic pans were noted to jump Bergsten’s (2012) analysis similarly placed Hetero- adeptly. To observe the species’ behavior in a more gyrus in an isolated position as sister to all the natural setting, specimens were placed on a dried Gyrininae, resulting in elevation of Brinck’s(1955) stream bed and again found to jump, but in this Heterogyrini Brinck to subfamilial status. The study also situation, individuals performed targeted, seem- suggested the morphology of Heterogyrus is transitional ingly non-random hops in a downstream direction. between the early diverging Spanglerogyrus and the Thus, the authors performed a small behavioral THE COLEOPTERISTS BULLETIN 71(2), 2017 317 experiment to test the hypothesis that H. milloti hops on 2 November 2014. Numerous specimens were in a non-random, targeted downstream manner. collected at random using a large strainer, then In this paper, we provide a redescription of the placed in a water-filled plastic container for holding species, more information on the habitat of H. during experimentation. Water for the holding milloti, report undocumented behavior with results container was collected from the same pool prior to from a small behavioral experiment, and finally collecting the beetles. Each trial consisted of first discuss the plesiomorphic, transitional, shared, and scooping a specimen from the plastic container unique morphology of this species in relation to using a small tea-strainer, placing the specimen on other Gyrinidae. Finally, common names are pro- the dried stream bed adjacent to the stream, then posed for H. milloti to aid in communication and observing the specimen’s behavior. Each trial (n 5 conservation of this species. 17) was recorded in situ by a Canon Legria HFS100 camcorder. For each trial, the specimen was placed roughly in MATERIAL AND METHODS the center of a small (ca. 30 cm), circular arena of dried stream bed. Once in the arena, specimens were Morphology and Taxonomy. In total, 71 spec- not manipulated and were allowed to acclimate imens were examined for this study. Material was before jumping. Specimens that did not exhibit any examined from the following collections: movement after ca. 20 seconds were not included in the final sample. Specimens where allowed to jump unimpeded until exiting the arena, at which point GTGC Grey Gustafson research collection, currently at they were caught by the first author and removed Department of Ecology and Evolutionary Biology, University of Kansas, Lawrence, from the arena. To insure the same specimens were Kansas, U.S.A. not re-sampled, they were collected into ethanol at MSBA Museum of Southwestern Biology, Division of the end of each trial. Specimens whose movements Arthropods, University of New Mexico, were not tracked by the observers present at the time Albuquerque, New Mexico, U.S.A (K. Miller). of the trial were also excluded from the final sample. NHRS Swedish Museum of Natural History, Stockholm, These qualifications resulted in two of the trials Sweden (J. Bergsten). being excluded and a final sample size of 15 trials. DEUA Departement d’Entomologie, Universit´e d’Antananarivo, Antananarivo, Madagascar. RESULTS Morphology and Taxonomy For descriptive and comparative purposes, Heterogyrus milloti Legros, 1953 specimens were examined using a Zeiss Discovery. V8 SteREO microscope with attached drawing tube. Material Examined. MADAGASCAR: Fia- Illustrations were first prepared under the drawing narantsoa Province: 7 km W Ranomafana, tube then scanned and traced using the software 1,100 m, 8-21.x.1988, leg. W. E. Steiner (3 ex. Adobe Illustrator. Female genitalia were prepared MSBA); same as previous except 1,000 m, 1-7. following the methods outlined in Miller and iii.1990 (3 ex. MSBA); ;8 km W Ranomafana Bergsten (2012) and were illustrated in water. town, stream, 21°14.992’S, 47°24.332’E, 20. Smaller anatomical structures such as the metacoxa xi.2008, leg. K. B. Miller, KBM20110801 (18 ex. and genitalia were first cleared using KOH then MSBA); Ranomafana NP, 3.8 km SE of Vohiparara examined using a Zeiss AXIO Imager A2 com- village, 21.2498°S, 47.4041°E, 1,080 m, 2.xi.2014, pound microscope with attached Axiocam 506 forest stream by road, leg. G. Gustafson GTG110214D mono camera. (6 ex. GTGC). Ranomafana NP, 3.8 km SE of Habitus and other color images were taken on a Vohiparara village, 21.2498°S, 47.4041°E, 1,080 m, Visionary Digital BK1 light imaging system (www. 2.xi.2014, forest stream by road, Leg. J. Bergsten, T. visionarydigital.com, R. Larimer). Habitus images Ranarilalatiana & S. Holmgren, MAD14-08, NHRS- were edited using Adobe Photoshop CS5 to improve JLKB000027148 (3 ex. NHRS); Ranomafana NP, clarity and color and to add scale bars. Scanning 3.8 km SE of Vohiparara village, 21.2498°S, electron microscope images were taken at the KU 47.4041°E, 1,080 m, 1.xi.2011, small forest Microscopy and Analytical Imaging Laboratory, stream with pools, leg. J. Bergsten, R. Bukontaite, T. University of Kansas, Lawrence, KS, USA. Ranarilalatiana & J. H. Randriamihaja, MAD11-11, Behavioral Experiments. Specimens
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