Climacia Californica Chandler, 1953 (Neuroptera: Sisyridae) in Utah: Taxonomic Identity, Host Association and Seasonal Occurrence
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AQUATIC INSECTS 2019, VOL. 40, NO. 4, 317–327 https://doi.org/10.1080/01650424.2019.1652329 Climacia californica Chandler, 1953 (Neuroptera: Sisyridae) in Utah: taxonomic identity, host association and seasonal occurrence Makani L. Fishera, Robert C. Mowerb and C. Riley Nelsona aDepartment of Biology and M. L. Bean Life Science Museum, Brigham Young University, Provo, UT, USA; bUtah County Mosquito Abatement, Spanish Fork, UT, USA ABSTRACT ARTICLE HISTORY We provide a record of spongillaflies (Sisyridae) with an associated Received 30 December 2018 host sponge from a population found in Spring Creek, Utah. We Accepted 15 May 2019 monitored the population for the 2016 field season to identify the First published online insect and its associated host sponge and to establish the sea- 30 September 2019 ’ sonal period of the insect s presence in the adult stage. We also KEYWORDS evaluated the commonly used sampling techniques of sweeping Aquatic Neuroptera; and light trapping and made natural history observations. We Sisyridae; freshwater identified the spongillaflies as Climacia californica Chandler, 1953 sponge; Utah; and the sponge to be Ephydatia fluviatilis (Linnaeus, 1758). We col- Intermountain West lected 1726 adults, and light trapping proved to be the superior collecting method. The population was characterised by a two- week emergence peak occurring at the end of July to beginning of August followed by a steep decline. Other habitats within the state contained E. fluviatilis and should be sampled using light traps at times when peak abundances are predicted to further understand the distribution of C. californica across the state and the Intermountain West. urn:lsid:zoobank.org:pub:F795281F-0042-4F5F-B15B-F6C584FACF2B Introduction Sisyridae, or spongillaflies, and freshwater sponges (Porifera: Spongillidae) are linked together in a parasite–host association (Parfin and Gurney 1956; Steffan 1967; Resh 1976; Pupedis 1980). Sisyrids feed on freshwater sponge with stylets used to pierce and suck the contents from individual sponge cells (Cover and Resh 2007; Parfin and Gurney 1956; Pupedis 1980; Figure 1a). Freshwater sponges are found globally and have a wide range of morphological forms (Manconi and Pronzato 2008). Spongillaflies, having adapted to many of the same habitats as their hosts, are also found worldwide (Cover and Resh 2007; Parfin and Gurney 1956). Within North CONTACT Makani Fisher [email protected] Color versions of one or more of the figures in the article can be found online at www.tandfonline.com/naqi. ß 2019 The Author(s). Published by Informa UK Limited, trading as Taylor & Francis Group This is an Open Access article distributed under the terms of the Creative Commons Attribution-NonCommercial-NoDerivatives License (http://creativecommons.org/licenses/by-nc-nd/4.0/), which permits non-commercial re-use, distribution, and reproduction in any medium, provided the original work is properly cited, and is not altered, transformed, or built upon in any way. Published online 30 Sep 2019 318 M. FISHER ET AL. Figure 1. Live Climacia californica Chandler, 1953 from Spring Creek, Springville, Utah: (A) larva on host sponge; (B) adult on leaf. America, there are two genera and six known species (Bowles 2006). Despite their occurrence in parts of western North America (Bowles 2006), sisyrids have only been recorded twice in the Intermountain West. First, a host association was made for them with the sponge Ephydatia mulleri (Lieberkuhn,€ 1856) in Idaho (Clark 1985). Second, a Utah record was made from a single female specimen collected at a light trap in 1979 from the state’s southern border (Bowles 2006). After this initial Utah record, sisyrids have been sought extensively throughout the state by both professionals and students, largely from Brigham Young University (BYU) and Utah State University (USU). These efforts were directed towards habitats with potential freshwater sponge hosts. Sponges have rarely been noted in Utah, but they do exist in some regions where they provide potential habitats for the insect. The junior author anecdotally observed them at Wellsville Reservoir in extreme northern Utah in the 1980s as well as in several side channels of Lake Powell begin- ning in 2012. Rader also identified Ephydatia fluviatilis (Linnaeus, 1758) in Utah AQUATIC INSECTS 319 Lake (Rader 1984; Rader and Winget 1985). Other researchers including Dennis Shiozawa, Russell Rader, Richard Baumann and various BYU aquatic entomology classes (personal communication, January 20, 2017) repeatedly searched many of these and other likely habitats throughout the state over the course of 35 years. None of those efforts yielded spongillaflies, but the second author (Utah County Mosquito Abatement, UCMA), observed sisyrid adults in mosquito light traps from Spring Creek, Springville, Utah in 2014 (Figure 1b). After repeatedly seeing sisryids in those traps, he alerted the junior author who began to survey the creek. In fall of 2014, BYU’s aquatic entomology class collected larvae and adult sisyrids from this creek in rather large numbers (Figure 1). After BYU students regularly collected specimens from this site in 2014 and 2015, we monitored the population for the 2016 field season. Records and dates have been published for North American sisyrids by Bowles (2006), but no populations have been consistently followed through a season. These collective efforts produced a new distributional record of spongillaflies for Utah and their associated sponge host. This record further expands their known distribution in the Intermountain West. We also compared the two commonly used sampling techniques of sweeping and light trap- ping (Bowles 2008) and observed the population’s seasonality. In particular, we inves- tigated the effectiveness of the sampling techniques, length of the adult flight season and the frequency and magnitude of emergence events. Compared to sweeping, we hypothesised light trapping to be superior as spongillaflies are strongly attracted to a variety of lights (Bowles 2008). In regard to their natural history, we expected the sis- yrids to be active from May to November (Bowles 2006) and to be multivoltine (DeWalt, Resh, and Hilsenhoff 2010). Material and methods We monitored sisyrids in Springville, Utah at the Utah Division of Wildlife Resources (UDWR) Fish Hatchery. The study site is located at 401005600N, 1113604300W at an elevation of 1395 m. Spring Creek flows through the site providing habitat for the sponges and sisyrids we examined. We identified suitable freshwater sponge habitat within a 100 m stretch of the creek and sampled it intensively. We found many larvae present on the sponge (Figure 1a) and collected both larvae and sponge for taxonomic purposes. Collected sisyrid larvae were stored in 70% ethanol. We attempted to identify the larvae using keys as detailed in the work by Pupedis (1980) and Bowles (2006). We sampled four of these sponges for identification by scraping their surfaces using soft tip forceps and placing the scrapings in 50 ml vials filled with 70% ethanol. We extracted their spicules using several techniques to aid in identification of the species. We first used the technique described by Smith and Pennak (2001) for extracting spicules using heated HNO3. We used a section of sponge tissue containing gemmules to ensure finding all types of spicules. This yielded both the megascleres and gemmoscleres (Figure 2) needed for proper identification. We also left sponge tissue in 10% vol potassium hydroxide overnight and found this to yield a good num- ber of megascleres, but the gemmules did not reduce to gemmoscleres. We then 320 M. FISHER ET AL. Figure 2. Ephydatia fluviatilis (Linnaeus, 1758) spicules extracted and cleared from host sponge collected at Spring Creek, Springville, Utah: (A) megascleres and gemmoscleres; (B) isolated gemmosclere. explored treating the gemmules in cold HNO3 for several days, and many individual gemmoscleres were obtained using this approach. We mounted cleared megascleres, gemmoscleres and sponge tissue onto glass slides for species identification (Ricciardi and Reiswig 1993). We repeated this process for sponge we collected earlier the same year from the Provo River in Provo, Utah. This sponge did not have any sisyrid larvae, but its spi- cules were similar to those from Spring Creek. We compared both these sponge iden- tifications to sponge previously identified from Utah Lake (Rader 1984; Rader and Winget 1985). This was done to identify other nearby local habitats with host sponge that may be suitable for the spongillaflies. We monitored the adult sisyrid population along Spring Creek and identified them to species using Bowles (2006) guide. We used wing veins and male genitalia for definitive identification (Figure 3) as female characteristics tend to be more variable (David Bowles, personal communication, January 21, 2016). The population was monitored by simultaneously using quantitative sweep sampling and light trapping. AQUATIC INSECTS 321 Figure 3. Taxonomically relevant parts of adult Climacia californica Chandler, 1953 from Spring Creek, Springville, Utah: (A) forewing; (B) cleared male genitalia with visible gonarcus complex, lat- eral view; (C) cleared male genitalia with visible gonarcus complex, dorsal view. The sampling techniques were done together to directly compare the two. If a speci- men was collected by one technique, it could not be collected by the other i.e., if specimens were effectively attracted to light, they would be taken up by the trap and sweep counts would be low and vice versa. For sweep samples we used a 45-cm collapsible net to sweep the vegetation along the north side of the creek each week a light trap sample was taken. We started at the edge of unmown vegetation 3 m east of the trap to 97 m beyond the trap, for a total of a 100 m sweep. We removed all sisyrid adults (Figure 1b) from the net and placed them into 70% ethanol. This process was repeated back to the starting point resulting in vegetation being swept twice for a total sweep of 200 m. We checked our net for sisyrids every 15 sweeps.