TY - JOUR
T1 - Phylogenetic relationships and convergent evolution of ocean-shore ground beetles (Coleoptera: Carabidae: Trechinae: Bembidion and relatives)
AU - Maddison, David R.
AU - Maruyama, Munetoshi
N1 - Funding Information:
This project would not have been possible without the generous help of those who collected specimens for our DNA studies. In particular, we would like to thank Agustín Aguiar, Antonio Machado, Toshihide Kato, Kazunari Nakano, Showtaro Kakizoe, Jun Nakajima, Mitsuru Moriguchi, Michael Caterino, J. Rykken, Konjev Desender, Paolo Bonavita, Werner Marggi, Igor Sokolov, P. Moravec, Kojun Kanda, Luca Toledano, and G. Giovagnoli. For their help in collecting, we thank H.M. Amerongen, A.E. Arnold, J. Galián, Shigehisa Hori, Takasuke Miyata, Toshie Miyata, Taku Tsukada, and Yutaka Sugawara. For their assistance in identifying voucher specimens, we thank all those listed in Table. In addition, we thank Joachim Schmidt for identifying the specimen of Bembidion leptaleum. For providing habitat information from their extensive experience collecting carabid beetles, we thank Luca Toledano, Paolo Neri, Paolo Bonavita, Joachim Schmidt, Marcus Toribio, Maurizio Pavesi, Werner Marggi, Jim LaBonte, David Kavanaugh, Luca Toledano, and M. Moriguchi. We also thank Takashi Komatsu for allowing us to use his pictures of living adults (Fig.), Werner Marggi for allowing us to use his photograph of a paratype of Bembidion eichleri (Supporting Information Fig. S2), and Michael Caterino for permission to use his image of the habitat of B. palosverdes (Fig.). For help with PCR, we thank Austin Baker, Joseph J. Dubie, Regina Kurapova, Kaitlyn Traynor, Kalyn M. Hansen, Danielle L. Mendez, and Estany M. Campbell-Dunfee. We are very thankful for James M. Pflug's help in making the genomic library preparation of the specimen of Orzolina. For help with data entry, we thank Tiana S.L. Week. For help in translating literature we thank R. Antonio Gomez. Thanks as well are due to John S. Sproul and Olivia F. Boyd for valuable comments on the manuscript. This project was funded in part by the Harold E. and Leona M. Rice Endowment Fund at Oregon State University.
Funding Information:
This project would not have been possible without the generous help of those who collected specimens for our DNA studies. In particular, we would like to thank Agustín Aguiar, Antonio Machado, Toshihide Kato, Kazunari Nakano, Showtaro Kakizoe, Jun Nakajima, Mitsuru Moriguchi, Michael Caterino, J. Rykken, Konjev Desender, Paolo Bonavita, Werner Marggi, Igor Sokolov, P. Moravec, Kojun Kanda, Luca Toledano, and G. Giovagnoli. For their help in collecting, we thank H.M. Ameron-gen, A.E. Arnold, J. Galián, Shigehisa Hori, Takasuke Miy-ata, Toshie Miyata, Taku Tsukada, and Yutaka Sugawara. For their assistance in identifying voucher specimens, we thank all those listed in Table 2. In addition, we thank Joachim Schmidt for identifying the specimen of Bembidion leptaleum. For providing habitat information from their extensive experience collecting carabid beetles, we thank Luca Toledano, Paolo Neri, Paolo Bonavita, Joachim Schmidt, Marcus Toribio, Maurizio Pavesi, Werner Marggi, Jim LaBonte, David Kavanaugh, Luca Toledano, and M. Moriguchi. We also thank Takashi Komatsu for allowing us to use his pictures of living adults (Fig. 2), Werner Marggi for allowing us to use his photograph of a paratype of Bembidion eichleri (Supporting Information Fig. S2), and Michael Caterino for permission to use his image of the habitat of B. palosverdes (Fig. 5). For help with PCR, we thank Austin Baker, Joseph J. Dubie, Regina Kurapova, Kaitlyn Traynor, Kalyn M. Hansen, Danielle L. Mendez, and Estany M. Campbell-Dunfee. We are very thankful for James M. Pflug’s help in making the genomic library preparation of the specimen of Orzolina. For help with data entry, we thank Tiana S.L. Week. For help in translating literature we thank R. Antonio Gomez. Thanks as well are due to John S. Sproul and Olivia F. Boyd for valuable comments on the manuscript. This project was funded in part by the Harold E. and Leona M. Rice Endowment Fund at Oregon State University.
Publisher Copyright:
© 2018 The Royal Entomological Society
PY - 2019/1
Y1 - 2019/1
N2 - Through phylogenetic analysis of seven genes, we show that there have been at least six independent entries into intertidal habitats in the history of bembidiine carabids, in the ancestors of: (i) Orzolina Machado, (ii) Bembidion (Desarmatocillenus Netolitzky), (iii) Bembidion laticeps (LeConte) + palosverdes Kavanaugh & Erwin, (iv) Bembidion laterale (Samouelle), (v) Bembidion umi Sasakawa and Bembidion quadriimpressum (Motschulsky) (which may represent two separate entries), and (vi) B. nigropiceum (Marsham). The following lineages are widely separated within the subtribe Bembidiina: Orzolina is sister to the genus Ocys Stephens; subgenus Desarmatocillenus appears to be sister to all Bembidion Latreille excluding subgenus Phyla Motschulsky; B. laticeps + B. palosverdes is a clade in the Bembidion series; B. laterale is a member of the Princidium Motschulsky complex; B. umi and B. quadriimpressum are related to the Nearctic Clade of the Ocydromus Clairville complex; B. nigropiceum is sister to B. praeustum Dejean among sampled species. There are three separate lineages of ocean-shore bembidiines that are known to prey on amphipods, and adults of these lineages [Bembidion (Desarmatocillenus), B. laterale, and B. mandibulare Solier] have unusually wide heads with long mandibles. Also common among independent lineages restricted to ocean shores are prominent front angles of the prothorax, larger numbers of setae on the elytral disc, a notable sinuation in the margin of each elytron near its apex, and short, wide mesotarsi. The reasons for the repeated evolution of these features are not evident. Our results also suggest that inland species of the Nearctic Clade may have arisen from an ocean-shore ancestor. The close genetic similarities between the gravel river shore dwelling B. praeustum and the intertidal specialist B. nigropiceum suggest that the striking morphological adaptations of B. nigropiceum to the intertidal zone arose rapidly. We make one nomenclatural change: we resurrect the subgenus Lymneops Casey to accommodate B. palosverdes and B. laticeps.
AB - Through phylogenetic analysis of seven genes, we show that there have been at least six independent entries into intertidal habitats in the history of bembidiine carabids, in the ancestors of: (i) Orzolina Machado, (ii) Bembidion (Desarmatocillenus Netolitzky), (iii) Bembidion laticeps (LeConte) + palosverdes Kavanaugh & Erwin, (iv) Bembidion laterale (Samouelle), (v) Bembidion umi Sasakawa and Bembidion quadriimpressum (Motschulsky) (which may represent two separate entries), and (vi) B. nigropiceum (Marsham). The following lineages are widely separated within the subtribe Bembidiina: Orzolina is sister to the genus Ocys Stephens; subgenus Desarmatocillenus appears to be sister to all Bembidion Latreille excluding subgenus Phyla Motschulsky; B. laticeps + B. palosverdes is a clade in the Bembidion series; B. laterale is a member of the Princidium Motschulsky complex; B. umi and B. quadriimpressum are related to the Nearctic Clade of the Ocydromus Clairville complex; B. nigropiceum is sister to B. praeustum Dejean among sampled species. There are three separate lineages of ocean-shore bembidiines that are known to prey on amphipods, and adults of these lineages [Bembidion (Desarmatocillenus), B. laterale, and B. mandibulare Solier] have unusually wide heads with long mandibles. Also common among independent lineages restricted to ocean shores are prominent front angles of the prothorax, larger numbers of setae on the elytral disc, a notable sinuation in the margin of each elytron near its apex, and short, wide mesotarsi. The reasons for the repeated evolution of these features are not evident. Our results also suggest that inland species of the Nearctic Clade may have arisen from an ocean-shore ancestor. The close genetic similarities between the gravel river shore dwelling B. praeustum and the intertidal specialist B. nigropiceum suggest that the striking morphological adaptations of B. nigropiceum to the intertidal zone arose rapidly. We make one nomenclatural change: we resurrect the subgenus Lymneops Casey to accommodate B. palosverdes and B. laticeps.
UR - http://www.scopus.com/inward/record.url?scp=85051129842&partnerID=8YFLogxK
UR - http://www.scopus.com/inward/citedby.url?scp=85051129842&partnerID=8YFLogxK
U2 - 10.1111/syen.12307
DO - 10.1111/syen.12307
M3 - Article
AN - SCOPUS:85051129842
SN - 0307-6970
VL - 44
SP - 39
EP - 60
JO - Systematic Entomology
JF - Systematic Entomology
IS - 1
ER -