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Cjb-2020-0163.Pdf Botany Preliminary observations on the flower visitors of Nasa colanii (Loasaceae), a poorly known species endemic to northern Peru Journal: Botany Manuscript ID cjb-2020-0163.R2 Manuscript Type: Note Date Submitted by the 12-Apr-2021 Author: Complete List of Authors: Wong Sato, Akira Armando; Universidad Nacional Agraria La Molina, Department of Forest Management; CORBIDI, Plant Ecology Division Villanueva-Espinoza, Rosa; CORBIDI, Plant Ecology Division Revilla, Italo;Draft Instituto científico Michael Owen Dillon Fernandez-Hilario, Robin; CORBIDI, Plant Ecology Division ; Universidade Federal do Paraná, Programa de Pós-Graduação em Botânica Keyword: Andes, cloud forest, pollination syndrome, syrphid flies, beetles Is the invited manuscript for consideration in a Special Not applicable (regular submission) Issue? : © The Author(s) or their Institution(s) Page 1 of 20 Botany Preliminary observations on the flower visitors of Nasa colanii (Loasaceae), a poorly known species endemic to northern Peru Akira A. Wong Sato1,2, Rosa Villanueva-Espinoza2, Italo Revilla3, Robin Fernandez-Hilario2,4 1. Departamento de Manejo Forestal, Universidad Nacional Agraria La Molina, Av. La Molina s/n, La Molina, Lima, Peru. 2. Plant Ecology Division – CORBIDI, Calle Santa Rita 105 Of. 2, Urb. Huertos de San Antonio Monterrico, Surco, Lima, Peru. 3. Instituto científico Michael Owen Dillon (IMOD). Av. Jorge Chávez 610, Arequipa, Peru. 4. Programa de Pós-Graduação em Botânica, Universidade Federal do Paraná. Caixa Postal 19031, CEP 81531-970, Curitiba, PR, Brazil. Akira A. Wong Sato: [email protected] Rosa Villanueva-Espinoza: [email protected] Italo Revilla: [email protected] Draft Robin Fernandez-Hilario: [email protected] Corresponding author: Akira Armando Wong Sato Telephone: +51-926-149-267 E-mail: [email protected] © The Author(s) or their Institution(s) Botany Page 2 of 20 Abstract Nasa is the most diverse genus among the family Loasaceae, and it is mainly distributed in Peru. Most species of this genus are microendemics, with distributions restricted to a very specific location; such is the case of Nasa colanii. This species–previously only known from the type collected in 1978 from the Amazonas region of Peru–lacks information about its biology and ecology. To improve our understanding of the reproductive biology and pollination ecology of N. colanii, we observed the flower development and insect floral visits in a cloud forest in the Amazonas region. The flowers of N. colanii are nectariferous and protandrous. The typical stamen movement to the flower center of the subfamily Loasoideae appears to be autonomous. Only well-developed (elongated) pistils showed positive stigmatic receptivity to 3% hydrogen peroxide. Syrphid flies (Baccha sp.) and small beetles (Baridinae sp. and Alticini sp.) were the only observed flower visitors. The syrphid flies visited the flower and foragedDraft for pollen, whereas the small beetles were found already in the flowers and seemingly used it as a shelter. Results suggest that N. colanii might be adapted to fly pollination. Andes, cloud forest, pollination syndrome, syrphid flies, beetles © The Author(s) or their Institution(s) Page 3 of 20 Botany Introduction The family Loasaceae Juss. comprises 20 genera and is mainly restricted to the New World, ranging from southern Mexico to southern Chile and extends from sea level up to the Puna of the Andes (4800 m altitude) (Weigend et al. 2003). In Peru, Loasaceae is represented by eight genera and about 112 species (Brako and Zarucchi 1993; Ulloa et al. 2004). Almost all Peruvian regions have records of this family, with the exception of the low jungle regions of Loreto, Madre de Dios, and Ucayali (Brako and Zarucchi 1993; Rodriguez and Weigend 2006; Ackermann 2012). Peru has an exceedingly high Loasaceae endemism level (52 species) and is considered the most important center of diversity of this family (Weigend et al., 2003; León et al. 2006). This high endemism is related to the diversity of the genus Nasa Weigend, the most diverse genus in Loasaceae, with about 100 described species (Weigend 2001). Nasa species are found between elevations of 1400 and 4700 m in northernDraft and central Andes (Weigend et al. 2003; Rodriguez and Weigend 2006), in the Mesoandean Humid and Dry Puna, and in Very Humid Montane Forest regions (Rodriguez and Weigend 2006). Most Nasa species are microendemic, with distributions restricted to specific locations, likely as a consequence of the continuously changing topography and climate of the Tropical Andes (Mutke et al. 2014; Acuña Castillo et al. 2019), which suggests that this genus have been able to adapt to various specific microhabitats (see Rodriguez and Weigend 2006). Cross-pollination and self-pollination systems are found in Loasaceae, but the latter is rare and mostly found in basal taxa of the family (Weigend 2004a; Strelin et al. 2016). Facultative autogamy has been reported for various genera, including Nasa, Loasa Adans. and Blumenbachia Schrad., while cleistogamy has been observed only in species of Nasa and Loasa (Weigend 2004a). Most Loasaceae species can be grouped into one of three pollination syndromes: bees, hummingbirds and rodents, among which the first and the last one are the most and least common, respectively (Cocucci and Sérsic 1998; Ackermann and Weigend 2006; Strelin et al. 2016). In © The Author(s) or their Institution(s) Botany Page 4 of 20 particular, Nasa species displays both bee and hummingbird pollination syndromes (Strelin et al. 2016; Henning et al. 2018). In flowers of the subfamily Loasoideae, pollen presentation occurs by the movement of mature stamens over the center of the flower (Weigend et al. 2010). This movement is typically thigmonastic (i.e., triggered by the manipulation of the nectar scale by the flower visitor), but it can also be autonomous (Cocucci and Sérsic 1998, Henning et al. 2018). Henning et al. (2018) showed that from 11 Nasa species, all but one autogamous species display the stamen thigmonastic movement. Interestingly, N. macrothyrsa (Urb. & Gilg) Weigend displays both movement types, perhaps to ensure pollination in the absence of pollinators (Weigend et al. 2010). The flowers of Nasa can be classified into two different floral types: “tilt-revolver” and “funnel-revolver”. In both floral types, nectarDraft is produced in the base of the nectar scales (structure composed of three fused staminodes). In tilt-revolver flowers, flower visitors must tilt the scales outward to extract the nectar (Weigend and Gottschling 2006), whereas in funnel-revolver flowers, the nectar is easily accessible (Weigend 2004b). Both floral types have been associated with bee- and bird-pollination, respectively (Weigend and Gottschling 2006). The present study is focused on Nasa colanii Dostert and Weigend, a rare species endemic to the Amazonas region of Peru. This species has been collected only once, more than 40 years ago, and virtually nothing is known about its biology, ecology and life history. Here, we present the first observations on the floral biology and flower visitors of this rare species. Methodology Study site and species Nasa colanii is a cloud forest endemic species, found only in the Cordillera de Colán National Sanctuary (SNCC) in the Amazonas region of Peru. Its distribution area has been calculated to be about 54 km2 (Young 2007). Rodriguez and Weigend (2006) commented that by the year 2006 the © The Author(s) or their Institution(s) Page 5 of 20 Botany habitat of N. colanii had not been altered due to the lack of scientific exploration. Furthermore, according to the head of the SNCC, only six scientific explorations have taken place near the “Refugio Lechuza” in the period of 2012 – 2020, including the one for this study (C. Olivera, personal communication, 2021). N. colanii is a small annual herb that grows up to 80 cm tall; however, smaller individuals are common. Flowers are star-shaped, 5-merous, pendent, with five nectar scales, numerous stamens and one pistil. The five petals are cymbiform, greenish-white. The nectar scales are yellow and white with red transversal stripes with two acuminate nectar sacs. As commonly found in Loasaceae, N. colanii presents numerous stamens grouped in epipetalous fascicles and an inferior ovary (Weigend 2004a). This species belongs to the Nasa triphylla complex, which is mainly distributed in the Andes mountains of the north of Peru (Dostert and Weigend 1999). Draft According to Rodriguez and Weigend (2006), N. colanii has not been collected since 1978, and data on its current state of conservation are virtually non-existent, as well as information on its biology and ecology, due to insufficient exploration of its distribution area. The study site was a creek habitat in a cloud forest ecosystem located in the buffer zone of the SNCC (5°37'50.97"S, 78°15'20.85"W) at an altitude of 2605 m, near the “Refugio Lechuza” of the SNCC (Fig. 1A). We visited the study site during December 19–22, 2019. Two voucher specimens are housed in the MOL and two others will be sent to the HSP (herbarium acronyms follow Thiers 2021). Insect visits We observed insect visits to six N. colanii flowers during a period of 8–17 h. on December 20–21, 2019. We also noted the behavior of the insect flower visitors, and morphospecied them by their phenotypic characteristics. © The Author(s) or their Institution(s) Botany Page 6 of 20 Reproductive biology To identify nectar production, we measured the volume of floral nectar from bagged flowers using 2 μl glass micropipettes and sugar concentration of the nectar using a refractometer (V- Resourcing, model VLT032). Stigmatic receptivity was determined with 3% hydrogen peroxide. Finally, to identify whether the stamen movement was thigmonastic, we tried to manually trigger it by using glass micropipettes, simulating insect nectar-foraging behavior. Results Flowering biology A single flower lasted for approximatelyDraft 2–3 days (Figs. 1B–E). A mature flower bud opens fully within 2 hours, and within the next hours the stamens move to the flower center.
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