Glime, J. M. 2021. Aquatic and Wet Marchantiophyta, Class Jungermanniopsida, Orders Porellales: Jubulineae, part 1. Chapt. 1-7. In: 1-7-1 Glime, J. M. (ed.). Bryophyte Ecology. Volume 4. Habitat and Role. Ebook sponsored by Michigan Technological University and the International Association of Bryologists. Last updated 22 May 2021 and available at <http://digitalcommons.mtu.edu/bryophyte-ecology/>. CHAPTER 1-7 AQUATIC AND WET MARCHANTIOPHYTA, CLASS JUNGERMANNIOPSIDA, ORDER PORELLALES: JUBULINEAE, PART 1 TABLE OF CONTENTS Porellales – Suborder Jubulineae ........................................................................................................................ 1-7-2 Frullaniaceae ................................................................................................................................................ 1-7-2 Frullania asagrayana ........................................................................................................................... 1-7-2 Frullania riparia ................................................................................................................................... 1-7-4 Frullania tamarisci ............................................................................................................................... 1-7-5 Frullania teneriffae ............................................................................................................................... 1-7-7 Jubulaceae .................................................................................................................................................... 1-7-9 Jubula hutchinsiae ............................................................................................................................... 1-7-9 Jubula hutchinsiae subsp. pennsylvanica ........................................................................................... 1-7-12 Jubula hutchinsiae var. integrifolia .................................................................................................... 1-7-13 Lejeuneaceae .............................................................................................................................................. 1-7-13 Acanthocoleus aberrans ...................................................................................................................... 1-7-13 Bromeliophila ..................................................................................................................................... 1-7-13 Bromeliophila helenae ........................................................................................................................ 1-7-14 Bromeliophila natans .......................................................................................................................... 1-7-14 Cephalantholejeunea temnanthoides .................................................................................................. 1-7-15 Ceratolejeunea temnantha .................................................................................................................. 1-7-15 Cheilolejeunea clypeata ...................................................................................................................... 1-7-16 Cololejeunea biddlecomiae ................................................................................................................. 1-7-18 Cololejeunea calcarea ........................................................................................................................ 1-7-22 Cololejeunea hodgsoniae .................................................................................................................... 1-7-23 Cololejeunea madothecoides .............................................................................................................. 1-7-23 Cololejeunea microscopica ................................................................................................................. 1-7-23 Cololejeunea rossettiana .................................................................................................................... 1-7-24 Cololejeunea stotleriana ..................................................................................................................... 1-7-25 Colura ................................................................................................................................................. 1-7-26 Colura calyptrifolia ............................................................................................................................ 1-7-27 Colura irrorata ................................................................................................................................... 1-7-28 Diplasiolejeunea cavifolia .................................................................................................................. 1-7-30 Summary ........................................................................................................................................................... 1-7-32 Acknowledgments ............................................................................................................................................. 1-7-33 Literature Cited ................................................................................................................................................. 1-7-33 1-7-2 Chapter 1-7: Aquatic and Wet Marchantiophyta, Class Jungermanniopsida, Order Porellales: Jubulineae, part 1 CHAPTER 1-7 AQUATIC AND WET MARCHANTIOPHYTA, CLASS JUNGERMANNIOPSIDA, ORDER PORELLALES: JUBULINEAE, PART 1 Figure 1. Jubula hutchinsiae habitat. Photo by Michael Lüth, with permission. Nomenclature for this chapter is based primarily on aquatic habitats with which I am most familiar, should be Söderström et al. (2016). In addition, Lars Söderström given priority. provided me with correct names for species that I could not Many of the species in this subchapter are not typical link to the names on that list. TROPICOS also permitted wetland or aquatic species. They were, however, found in me to link names by tracking the basionym. I have ignored a wetland or aquatic study. Their relative frequency can be varieties, forms, and subspecies unless I could verify a inferred based on the number of references cited. current name for them. These unverifiable taxa have been included in the species. To develop this list, I used my own bibliography, Porellales – Suborder Jubulineae collected over the past 56 years, and Google Scholar. These papers soon led me to others. I do not pretend that Frullaniaceae this is complete. It includes streams, lakes, and a wide This family occurs predominantly on boulders and tree range of other wetlands. It deliberately ignores bogs trunks and branches. However, a few taxa seem at least (especially Sphagnum) and mostly ignores fens, but tolerant of more moist habitats and have been included nevertheless includes some species from these habitats here. because they were found in a wetland study. Bogs and poor fens have been treated in whole books and provide an extensive literature; fens seem somewhat less studied. They would require considerably more review and time. Frullania asagrayana (Figure 2) Thus I felt that less-reviewed topics, particularly the (syn. = Frullania tamarisci subsp. asagrayana) Chapter 1-7: Aquatic and Wet Marchantiophyta, Class Jungermanniopsida, Order Porellales: Jubulineae, part 1 7-1-3 Although Frullania asagrayana (Figure 2) has been Distribution treated as a subspecies of Frullania tamarisci (Figure 3), Frullania asagrayana (Figure 2) is distributed broadly Crandall-Stotler et al. (1987) found that both throughout eastern North America from Canada to the Gulf morphological and biochemical differences among these Coast (Ramaiya et al. 2010) and westward to the Ozarks subspecies indicated a greater genetic distance than that (Schuster 1992). The leaves have ocelli (Figure 4-Figure expected for subspecies. This includes unique flavonoid 5), a character absent in many Frullania species. profiles and a considerable level of terpenoid and phosphoglucoisomerase dissimilarity. Heinrichs et al. (2010) likewise supported its recognition as a separate species based on DNA markers, but found these were not as strongly supported by morphological differences. They considered this species complex to be "the most explicit example of a complex of semi-cryptic and cryptic liverwort species." Their findings "support frequent short-distance migration, rare successful long-distance dispersal events, extinction, and recolonization as an explanation for the range formation in these Frullania species." Figure 4. Frullania asagrayana leaf cells and ocelli (line of brown cells). Photo by Blanka Aguero, with permission. Figure 2. Frullania asagrayana (reddish) with Drepanolejeunea appalachiana (green) growing on it. Photo by Ken McFarland and Paul Davison, with permission. Figure 5. Frullania asagrayana ocelli and leaf cells showing oil bodies where secondary compounds are stored. Photo by Blanka Aguero, with permission. Aquatic and Wet Habitats Frullania asagrayana (Figure 2) occurs in crevices or the surface of drier cliffs in a ravine in Connecticut, USA, but Nichols (1916) lists it in his study along rivers and streams from these habitats. Otherwise, it does not seem to be truly aquatic or wetland. More frequently, Frullania asagrayana (Figure 2) is epiphytic (Davison 1997). Reproduction Frullania asagrayana is dioicous (Figure 6), so one is Figure 3.