Mosaic phenotypic evolution underlies the adaptive success of water-surface colonization in Gerromorpha

Z Zezhong Jin (Institute of Entomology, College of Life Sciences, Nankai University) M Mu Qiao S Siying Fu (Institute of Entomology, College of Life Sciences, Nankai University) Z Zihe Li (Institute of Entomology, College of Life Sciences, Nankai University) B Boxiong Guo (Institute of Entomology, College of Life Sciences, Nankai University) H Hongjiao Li (Institute of Entomology, College of Life Sciences, Nankai University) Z Zhaoqi Leng (Institute of Entomology, College of Life Sciences, Nankai University) M Matthew R. Pintar (Institute of Environment, Florida International University) J Jakob Damgaard (Natural History Museum of Denmark, Zoological Museum) B Benjamin L. Makepeace (Institute of Infection, Veterinary and Ecological Sciences, University of Liverpool) S Silvia A. Mazzucconi (Laboratorio de Entomología, Instituto de Biodiversidad y Biología Experimental y Aplicada, Consejo Nacional de Investigaciones Científicas y Técnicas - Universidad de Buenos Aires, Departamento de Biodiversidad y Biología Experimental - Facultad de Ciencias Exactas y Naturales, Universidad de Buenos Aires) G Gavril Marius Berchi (Department of Biology, Faculty of Chemistry, Biology, Geography, West University of Timișoara) F Fabio Cianferoni (Research Institute on Terrestrial Ecosystems, National Research Council of Italy) D Dan A. Polhemus (Department of Natural Sciences, Bishop Museum) K Kohei Watanabe (Ishikawa Insect Museum) J Jun Nakajima S Seraphine Esemu (Laboratory for Emerging Infectious Diseases, Faculty of Science, University of Buea) C Chen Liu B Beichen Zhang (Institute of Entomology, College of Life Sciences, Nankai University) H Huanhuan Yang (School of Bioengineering, Qilu University of Technology (Shandong Academy of Sciences)) S Shujing Wang (Institute of Entomology, College of Life Sciences, Nankai University) H Huaijun Xue (Institute of Entomology, College of Life Sciences, Nankai University) W Wenjun Bu (Institute of Entomology, College of Life Sciences, Nankai University) Z Zhen Ye

Abstract

Understanding how the remarkable phenotypic diversity observed in organisms arises through shifts in macroevolutionary patterns and tempos is a fundamental challenge in evolutionary biology. Phenotypes often evolve in a mosaic pattern during adaptive transitions. For organisms that have invaded highly specialized habitats, such as the unique two-phase interface habitats (water surfaces), the macroevolutionary history of their phenotypic diversification remains only superficially understood. Semiaquatic bugs (Insecta: Heteroptera: Gerromorpha), which exhibit extensive habitat diversification and unparalleled phenotypic innovation, represent one of the most successful extant adaptive groups at this interface and provide an excellent system for study. By analyzing their adaptive transitions and phenotypic macroevolutionary history, we demonstrate that Gerromorpha experienced a single major adaptive transition from land to the water surface. Subsequently, semiaquatic bugs successfully colonized a wide range of distinct water-surface habitats. During this process, phenotypic space was explored under strong constraints and along pronounced mosaic trajectories; i.e., different body regions exhibited markedly distinct patterns of phenotypic space occupation and partitioning, as well as markedly different evolutionary rates. Furthermore, we showed that this mosaic pattern of phenotypic space occupation and rates of exploration had complex and critical effects on the invasion and colonization of water-surface habitats. Our study provides a typical case of macroevolutionary dynamics in species adapted to specialized air–water interface habitats, emphasizing the complex and significant roles of environmental context and functional demands in shaping patterns of phenotypic evolution.

Article Details

Volume / Issue Vol. 123, Issue 12
Published March 24, 2026
ISSN 0027-8424
Publisher National Academy of Sciences

Authors (24)

Z

Zezhong Jin

Institute of Entomology, College of Life Sciences, Nankai University

M

Mu Qiao

S

Siying Fu

Institute of Entomology, College of Life Sciences, Nankai University

Z

Zihe Li

Institute of Entomology, College of Life Sciences, Nankai University

B

Boxiong Guo

Institute of Entomology, College of Life Sciences, Nankai University

H

Hongjiao Li

Institute of Entomology, College of Life Sciences, Nankai University

Z

Zhaoqi Leng

Institute of Entomology, College of Life Sciences, Nankai University

M

Matthew R. Pintar

Institute of Environment, Florida International University

J

Jakob Damgaard

Natural History Museum of Denmark, Zoological Museum

B

Benjamin L. Makepeace

Institute of Infection, Veterinary and Ecological Sciences, University of Liverpool

S

Silvia A. Mazzucconi

Laboratorio de Entomología, Instituto de Biodiversidad y Biología Experimental y Aplicada, Consejo Nacional de Investigaciones Científicas y Técnicas - Universidad de Buenos Aires, Departamento de Biodiversidad y Biología Experimental - Facultad de Ciencias Exactas y Naturales, Universidad de Buenos Aires

G

Gavril Marius Berchi

Department of Biology, Faculty of Chemistry, Biology, Geography, West University of Timișoara

F

Fabio Cianferoni

Research Institute on Terrestrial Ecosystems, National Research Council of Italy

D

Dan A. Polhemus

Department of Natural Sciences, Bishop Museum

K

Kohei Watanabe

Ishikawa Insect Museum

J

Jun Nakajima

S

Seraphine Esemu

Laboratory for Emerging Infectious Diseases, Faculty of Science, University of Buea

C

Chen Liu

B

Beichen Zhang

Institute of Entomology, College of Life Sciences, Nankai University

H

Huanhuan Yang

School of Bioengineering, Qilu University of Technology (Shandong Academy of Sciences)

S

Shujing Wang

Institute of Entomology, College of Life Sciences, Nankai University

H

Huaijun Xue

Institute of Entomology, College of Life Sciences, Nankai University

W

Wenjun Bu

Institute of Entomology, College of Life Sciences, Nankai University

Z

Zhen Ye