Space colonization by branching trachea explains the morphospace of a simple respiratory organ.

Branching morphogenesis helps increase the efficiency of gas and liquid transport in many animal organs. Studies in several model organisms have highlighted the molecular and cellular complexity behind branching morphogenesis. To understand this complexity, computational models have been developed with the goal of identifying the "major rules" that globally explain the branching patterns. These models also guide further experimental exploration of the biological processes that execute and maintain these rules. In this paper we introduce the tracheal gills of mayfly (Ephemeroptera) larvae as a model system to study the generation of branched respiratory patterns. First, we describe the gills of the mayfly Cloeon dipterum, and quantitatively characterize the geometry of its branching trachea. We next extend this characterization to those of related species to generate the morphospace of branching patterns. Then, we show how an algorithm based on the "space colonization" concept (SCA) can generate this branching morphospace via growth towards a hypothetical attractor molecule (M). SCA differs from other branch-generating algorithms in that the geometry generated depends to a great extent on its perception of the "external" space available for branching, uses few rules and, importantly, can be easily translated into a realistic "biological patterning algorithm". We identified a gene in the C. dipterum genome (Cd-bnl) that is orthologous to the fibroblast growth factor branchless (bnl), which stimulates growth and branching of embryonic trachea in Drosophila. In C. dipterum, this gene is expressed in the gill margins and areas of finer tracheolar branching from thicker trachea. Thus, Cd-bnl may perform the function of M in our model. Finally, we discuss this general mechanism in the context of other branching pattern-generating algorithms.

Revealing the diversity of Cloeodes Traver, 1938 (Ephemeroptera: Baetidae) in the Neotropics: description of eleven new species from Brazilian mountain ranges.

In the present work, based on material from distinct mountain ranges in Brazil, 11 new species of Cloeodes with hind wings or hind wing pads are described, illustrated and discussed. Among the new species, in C. aiuruoca, C. amantykyra, C. atlanticus, C. boldrinii and C. ioachimi, the apex of the fore femora in the nymphs is extremely projected (a characteristic previously found only in two species of the genus). Cloeodes guara and C. tracheatus share the presence of unusual large and dark gills. Cloeodes melanotarsus is readily distinguished by conspicuous blackish tarsi, C. lucifer by the bright yellow coloration of tergum I, and C. xyrognathos by blade-like incisors. Cloeodes magnus, besides being the largest species of the genus, with a body size reaching 12 mm, possess short maxillary palp. Comments on the presence of C. irvingi and C. opacus in Brazil are also provided. An interactive online key is provided for the nymphs of all the species of Cloeodes in which hind wing pads are present.

Contribution to the knowledge of Tenuibaetis Kang & Yang 1994, Nigrobaetis Novikova & Kluge 1987 and Labiobaetis Novikova & Kluge 1987 (Ephemeroptera: Baetidae) from the Western Ghats (India).

As part of ongoing exploration of the mayflies of hill streams of southern Western Ghats, we describe two new species: Nigrobaetis paramakalyani Kubendran & Balasubramanian n. sp. and Labiobaetis jacobusi Kubendran & Balasubramanian n. sp. The genus Nigrobaetis is reported for the first time from India. Tenuibaetis frequentus (Müller-Liebenau & Hubbard 1985) n. comb. was also collected in this area and we provide the first description of this species at the imaginal stage. Brief ecological notes of the three species are added.