Larval morphology of the diving beetle Celina and the phylogeny of ancestral hydroporines (Coleoptera : Dytiscidae : Hydroporinae)
Mariano C. Michat A D , Yves Alarie B , Patricia L. M. Torres A and Yoandri S. Megna CA Laboratorio de Entomología, Departamento de Biodiversidad y Biología Experimental, Facultad de Ciencias Exactas y Naturales, Universidad de Buenos Aires, Av. Int. Güiraldes s/n, Ciudad Universitaria, C1428EHA, Buenos Aires, Argentina.
B Laurentian University, Sudbury, Ontario, Canada.
C Departamento de Biología, Universidad de Oriente. Patricio Lumumba s/n, Santiago, Santiago de Cuba, Cuba.
D Corresponding author. Email: marianoide@gmail.com
Invertebrate Systematics 21(3) 239-254 https://doi.org/10.1071/IS06037
Submitted: 10 September 2006 Accepted: 22 March 2007 Published: 27 June 2007
Abstract
Phylogenetic relationships within diving beetles (Dytiscidae) are imperfectly known. In particular, some authors have considered that the tribe Methlini is included in the subfamily Hydroporinae (a large group including about half of the dytiscid species worldwide), whereas others have argued in favour of excluding Methlini from the Hydroporinae and giving it subfamilial rank. Larval characters have been underutilised in phylogenetic studies, mainly because the larvae of many taxa within the family are still unknown. The larval morphology of the dytiscid tribe Methlini, in particular, remains poorly known. In this study, the phylogenetic relationships among ancestral lineages of the Hydroporinae are investigated based on a cladistic analysis of 34 taxa and 127 morphological larval characters. For this purpose, larvae of the Methlini (Celina parallela (Babington, 1841)) and C. imitatrix Young, 1979) are described and illustrated in detail for the first time, with particular emphasis on morphometry and chaetotaxy. The results show high support for a monophyletic origin of the Hydroporinae, including Methlini, based on eight unique character states. Giving Methlini subfamily rank would leave Hydroporinae with a single unique larval apomorphy. This supports the inclusion of Methlini as a tribe of Hydroporinae. Other interesting but less well supported results include: 1, the clade Laccornini + Hydrovatini + Canthyporus Zimmermann, 1919 (Hydroporini) resolved as the sister-group to the other Hydroporinae minus Methlini; and 2, Hydrovatini and Canthyporus resolved as sister-groups. The presence of a galea, albeit in a reduced form, in larvae of Methlini, Laccornini and Hydrovatini is of the utmost interest. The putative hypothesis of an ancestral position for these genera within Hydroporinae suggests that hydroporines lost the galea secondarily.
Acknowledgements
We thank the anonymous referees whose comments helped improve this manuscript. Field and laboratory work of M. C. Michat and P. L. M. Torres was supported by postgraduate scholarships and grant PIP 02541/00 from the Consejo Nacional de Investigaciones Científicas y Técnicas de la República Argentina (CONICET), and grant PIB X846 from the Universidad de Buenos Aires (UBA). Financial support was also provided by the Natural Sciences and Engineering Research Council of Canada in the form of an operating research grant to Y. Alarie.
Alarie Y.
(1989) The larvae of Laccornis Des Gozis 1914 (Coleoptera: Adepaga: Dytiscidae) with description of L. latens (Fall 1937) and redescription of L. conoideus (Leconte 1850). Coleopterists Bulletin 43, 365–378.
Alarie Y.
(1991a) Primary setae and pores on the cephalic capsule and head appendages of larval Hydroporinae (Coleoptera: Dytiscidae). Canadian Journal of Zoology 69, 2255–2265.
Alarie Y.
(1991b) Description of larvae of 17 Nearctic species of Hydroporus Clairville (Coleoptera: Dytiscidae: Hydroporinae) with an analysis of their phylogenetic relationships. Canadian Entomologist 123, 627–704.
Alarie Y.
(1995) Primary setae and pores on the legs, the last abdominal segment, and the urogomphi of larvae of Nearctic Colymbetinae (Coleoptera: Adephaga: Dytiscidae) with an analysis of their phylogenetic relationships. Canadian Entomologist 127, 913–943.
Alarie Y.
(1997) Taxonomic revision and phylogenetic analysis of the genus Oreodytes Seidlitz (Coleoptera: Dytiscidae: Hydroporinae) based on larval morphology. Canadian Entomologist 129, 399–503.
Alarie Y.
(1998) Phylogenetic relationships of Nearctic Colymbetinae (Coleoptera: Adephaga: Dytiscidae) based on chaetotaxic and porotaxic analysis of head capsule and appendages of larvae. Canadian Entomologist 130, 803–824.
Alarie Y.,
Beutel R. G., Watts C. H. S.
(2004) Larval morphology of three species of Hygrobiidae (Coleoptera: Adephaga: Dytiscoidea) with phylogenetic considerations. European Journal of Entomology 101, 293–311.
Alarie Y., Bilton D. T.
(2005) Larval morphology of Aspidytidae (Coleoptera: Adephaga) and its phylogenetic implications. Annals of the Entomological Society of America 98, 417–430.
| Crossref | GoogleScholarGoogle Scholar |
Alarie Y., Challet G. L.
(2006) Larval description and phylogenetic placement of the South African endemic genus Andex Sharp (Coleoptera: Adephaga: Dytiscidae). Annals of the Entomological Society of America 99, 743–754.
| Crossref | GoogleScholarGoogle Scholar |
Alarie Y.,
Cuppen J. G. M.,
Hendrich L., Nilsson A. N.
(2001a) Description of larvae of Herophydrus musicus (Klug) and analysis of relationships with members of the genus Hygrotus Stephens (Coleoptera: Dytiscidae, Hydroporinae). Aquatic Insects 23, 193–207.
| Crossref | GoogleScholarGoogle Scholar |
Alarie Y., Harper P. P.
(1990) Primary setae and pores on the last abdominal segment and the urogomphi of larval Hydroporinae (Coleoptera: Adephaga: Dytiscidae), with notes on other dytiscid larvae. Canadian Journal of Zoology 68, 368–374.
Alarie Y.,
Harper P. P., Maire A.
(1990a) Primary setae and pores on legs of larvae of Nearctic Hydroporinae (Coleoptera: Dytiscidae). Quaestiones Entomologicae 26, 199–210.
Alarie Y.,
Harper P. P., Roughley R. E.
(1990b) Description of the larvae of eleven Nearctic species of Hygrotus Stephens (Coleoptera: Dytiscidae: Hydroporinae) with an analysis of their phyletic relationships. Canadian Entomologist 122, 985–1035.
Alarie Y., Michat M. C.
(in press) Primary setae and pores on the maxilla of larvae of the subfamily Hydroporinae (Coleoptera: Adephaga: Dytiscidae): ground-plan pattern reconsidered. Coleopterists Bulletin 61,
Alarie Y., Nilsson A. N.
(1997) Larvae of Stictonectes Brinck: generic characteristics, description of S. canariensis Machado, and analysis of phylogenetic relationships with other genera of the tribe Hydroporini (Coleoptera: Dytiscidae). Coleopterists Bulletin 51, 120–139.
Alarie Y.,
Nilsson A. N., Hendrich L.
(1999) Larval morphology of the palearctic genera Deronectes Sharp and Scarodytes Gozis (Coleoptera: Dytiscidae: Hydroporinae), with implications for the phylogeny of the Deronectes-group of genera. Entomologica Scandinavica 30, 178–195.
Alarie Y.,
Nilsson A. N.,
Hendrich L., Watts C. H. S.
(2000) Larval morphology of four genera of Laccophilinae (Coleoptera: Adephaga: Dytiscidae) with an analysis of their phylogenetic relationships. Insect Systematics and Evolution 31, 121–164.
Alarie Y.,
Wang L. J.,
Nilsson A. N., Spangler P. J.
(1997) Larval morphology of four genera of the tribe Hyphydrini Sharp (Coleoptera: Dytiscidae: Hydroporinae) with an analysis of their phylogenetic relationships. Annals of the Entomological Society of America 90, 709–735.
Alarie Y., Watts C. H. S.
(2004) Larvae of the genus Antiporus (Coleoptera: Dytiscidae) and phylogenetic implications. Invertebrate Systematics 18, 523–546.
| Crossref | GoogleScholarGoogle Scholar |
Alarie Y.,
Watts C. H. S., Nilsson A. N.
(2001b) Larval morphology of the tribe Matini (Coleoptera: Dytiscidae: Colymbetinae): descriptions of Batrachomatus daemeli, Matus bicarinatus, and Allomatus nannup and phylogenetic relationships. Canadian Entomologist 133, 165–196.
Crespo F. A.
(1994) Descripción del tercer estadio larval de Celina conspicua Zimmermann 1921 (Coleoptera, Dytiscidae). Physis, Seccion B 52, 1–7.
De Marzo L., Nilsson A. N.
(1986) Morphological variation and fine structure of some head structures in larvae of Dytiscidae (Coleoptera). Entomologica Basiliensia 11, 29–42.
Michat M. C.
(2006) The phylogenetic position of Hydrovatus Motschulsky: evidence from larval morphology of H. caraibus Sharp (Coleoptera: Dytiscidae: Hydroporinae). Insect Systematics and Evolution 37, 419–432.
Michat M. C., Alarie Y.
(2006) The larvae of Amarodytes duponti (Aubé) (Coleoptera: Dytiscidae: Hydroporinae), with comments on Bidessini larval morphology and chaetotaxy. Zootaxa 1351, 1–13.
Michat M. C., Archangelsky M.
(in press) Descriptions of larvae of Desmopachria Babington (Coleoptera: Dytiscidae: Hydroporinae): the D. vicina Sharp species group. Coleopterists Bulletin 61, 000–000.
Michat M. C.,
Archangelsky M., Torres P. L. M.
(2005) Descriptions of the preimaginal stages of Lancetes marginatus (Steinheil) and L. biremis Říha (Coleoptera: Dytiscidae), and comparative notes with other Lancetes larvae. Studies on Neotropical Fauna and Environment 40, 129–142.
| Crossref | GoogleScholarGoogle Scholar |
Michat M. C., Torres P. L. M.
(2005) Larval morphology of Macrovatellus haagi (Wehncke) and phylogeny of Hydroporinae (Coleoptera: Dytiscidae). Insect Systematics and Evolution 36, 199–217.
Michat M. C., Torres P. L. M.
(2006)
Hydaticus tuyuensis Trémouilles (Coleoptera: Dytiscidae): larval morphology and phylogenetic relationships within Dytiscinae. Hydrobiologia 563, 479–492.
| Crossref | GoogleScholarGoogle Scholar |
Miller K. B.
(2001) On the phylogeny of the Dytiscidae (Insecta: Coleoptera) with emphasis on the morphology of the female reproductive system. Insect Systematics and Evolution 32, 45–92.
Pederzani F.
(1995) Keys to the identification of the genera and subgenera of adult Dytiscidae (sensu lato) of the world (Coleoptera Dytiscidae). Atti dell’Accademia Roveretana degli Agiati 224, 5–83.
Shaverdo H. V., Alarie Y.
(2006) Description of the larva of Canthyporus kenyensis Bilardo & Sanfilippo (Coleoptera: Dytiscidae: Hydroporinae) with implication for the phylogeny of the Hydroporini. Aquatic Insects 28, 113–130.
| Crossref | GoogleScholarGoogle Scholar |
Spangler P. J.
(1973) A description of the larva of Celina angustata Aubé (Coleoptera, Dytiscidae). Journal of the Washington Academy of Sciences 63, 165–168.
Wolfe G. W.
(1985) A phylogenetic analysis of plesiotypic Hydroporinae lineages with an emphasis on Laccornis des Gozis (Coleoptera: Dytiscidae). Proceedings. Academy of Natural Sciences of Philadelphia 137, 132–155.
Wolfe G. W.
(1988) A phylogenetic investigation of Hydrovatus, Methlini and other plesiotypic hydroporines (Coleoptera: Dytiscidae). Psyche 95, 327–344.
Young F. N.
(1979) A key to the Nearctic species of Celina with descriptions of new species (Coleoptera: Dytiscidae). Journal of the Kansas Entomological Society 52, 820–830.