Terrestrial arthropods of Steel Creek, Buffalo National River, Arkansas. II. Sawflies (Insecta: Hymenoptera: "Symphyta")

Abstract Background This is the second in a series of papers detailing the terrestrial arthropods collected during an intensive survey of a site near Steel Creek campground along the Buffalo National River in Arkansas. The survey was conducted over a period of eight and a half months using twelve trap types – Malaise traps, canopy traps (upper and lower collector), Lindgren multifunnel traps (black, green, and purple), pan traps (blue, purple, red, white, and yellow), and pitfall traps – and Berlese-Tullgren extraction of leaf litter. New information We provide collection records for 47 species of "Symphyta" (Insecta: Hymenoptera), 30 of which are new state records for Arkansas: (Argidae) Sterictiphora serotina; (Cimbicidae) Abia americana; (Diprionidae) Monoctenus fulvus; (Orussidae) Orussus terminalis; (Pamphiliidae) Onycholyda luteicornis, Pamphilius ocreatus, P. persicum, P. rileyi; (Pergidae) Acordulecera dorsalis, A. mellina, A. pellucida; (Tenthredinidae) Caliroa quercuscoccineae, Empria coryli, Hoplocampa marlatti, Macrophya cassandra, Monophadnoides conspiculatus, Monophadnus bakeri, Nematus abbotii, Neopareophora litura, Pachynematus corniger, Paracharactus rudis, Periclista marginicollis, Pristiphora banski, P. chlorea, Strongylogaster impressata, S. remota, Taxonus epicera, Thrinax albidopictus, T. multicinctus, Zaschizonyx montana; (Xiphydriidae) Xiphydria tibialis.


Introduction
The Interior Highlands is a mountainous physiogeographic region in the central United States that has remained unsubmerged and unglaciated since it was upthrust 270 mya (Bretz 1965, Robison and Allen 1995, Guccione 2008, U.S. Geological Survey 2014. It has acted as a refugium during times of inhospitable climate and as a result is a region of high biodiversity and endemism (Skvarla et al. 2015). However, terrestrial arthropods of the region have been understudied and undersurveyed compared to similar areas, such as the Southern Appalachains (Skvarla 2015). This paper, which focuses on "Symphyta", is the second in a series that intends to begin to rectify this situation (see Skvarla et al. 2015 for select Coleoptera).
"Symphyta", which are commonly referred to as sawflies and woodwasps because of the serrated or saw-shaped ovipositor present in many taxa and the habit of some taxa to bore into wood, is a paraphyletic grade of basal hymenopterans (Heraty et al. 2011). The majority of species are phytophagous, feeding both externally on leaves and internally as leaf miners and wood-and stem-borers on both monocots and dicots, though Orussidae, which are parasitic on wood-boring Hymenoptera and Coleoptera, are the notable exception. Historically, relatively few local or regional surveys have been reported in North America north of Mexico, though a number of studies conducted in eastern states have been published in recent years (e.g., Braud et al. 2003, Barrows and Smith 2014, Smith 2006b, Smith 2008c, Smith 2008a).
Herein we provide collection records for 47 species of sawflies collected in Arkansas.

Sampling methods
Sampling description: The sampling protocol was covered in detail by Skvarla et al. 2015.
The following summary is provided for convenience.
The following traps were maintained within a 4 ha site at Steel Creek, Buffalo National River, Arkansas (see Geographic coverage for a description of the site): five Malaise traps, twenty-five pan traps (five of each color: blue, purple, red, yellow, white); fifteen Lindgren multi-funnel traps (five of each color: black, green, purple); four SLAM (Sea, Land, and Air Malaise) traps with top and bottom collectors placed in or just below the canopy; and seventeen pitfall trap sets. Additionally, ten leaf litter samples were collected for Berlese extraction when traps were serviced.
Trap placement began on 8 March 2013 and all traps were set by 13 March 2013, except Lindgren funnels, which were set on 1 April 2013. Traps set earlier than 13 March were reset on that date in order to standardize trap catch between traps. Traps were serviced approximately every two weeks. The last collection of pitfall traps and pan traps occurred on 6 November 2013; Malaise, SLAM, and Lindgren funnel traps were run for an additional month, with the final collection on 4 December 2013. In total, 1311 samples were collected.
Propylene glycol (in the form of Peak RV & Marine Antifreeze) was used as the preservative in all traps as it is non-toxic, inexpensive, and preserves specimens reasonably well (Skvarla et al. 2014). Insect escape was impeded by the addition of unscented, hypoallergenic dish detergent to the propylene glycol to act as a surfactant. Trap catch was sieved in the field and stored in Whirl-Pak bags in 90% ethanol until sorting.
Quality control: Samples were coarse-sorted to suborder using a Leica MZ16 stereomicroscope illuminated with a Leica KL1500 LCD light source and a Wild M38 stereomicroscope illuminated with an Applied Scientific Devices Corp. Eco-light 20 fiber optic light source. After sorting, specimens were stored in 2 mL microtubes in 70% ethanol.
Specimens were identified by author Smith, who is an internationally recognized sawfly expert. Exemplar specimens are deposited in the National Museum of Natural History, Smithsonian Institution (USNM), with the remainder deposited in the University of Arkansas Arthropod Museum (UAAM). Distribution information is primarily from Smith 1979a; additional distribution references are indicated by species in "Notes on Select Species" below.

Geographic coverage
Description: The survey was conducted within a 4 hectare plot established at Steel Creek along the Buffalo National River in Newton County, Arkansas. The site is primarily 80-100 year old mature second-growth Eastern mixed deciduous forest dominated by oak (Quercus) and hickory (Carya), though American beech (Fagus grandifolia ) and eastern red cedar (Juniperus virginiana) are also abundant. A glade (10 m x 30 m) with sparse grasses was present within the boundaries of the site.

Results
We collected and identified 468 specimens representing 47 species and 31 genera during this study (Table 1). Thirty-one species (including Orussus minutus, which was recently reported from Arkansas by Skvarla et al. 2015), which represent 66% of the total species collected, are newly recorded in Arkansas. All trap types except Berlese extraction of leaf litter produced sawflies. While an in in-depth analysis of the data is in preparation, we include here the total number of species and specimens collected per trap type and the average number of specimens collected per trap (total specimens per trap type/total number traps per trap type) (  (Britton 1925, Middlekauff 1956, Furniss 1972, Still et al. 1974. Larvae feed on honeysuckle (Lonicera), including non-native species (Britton 1925, Middlekauff 1956, Baron and Bisdee 1984. ( Orussus terminalis (Orussidae) (new state record) occurs from New England and Ontario west to Iowa and Illinois (Middlekauff 1983).

Monoctenus fulvus
Onycholyda luteicornis (Pamphiliidae) (new state record) occurs from Nova Scotia and New Brunswick south to Maryland, Virginia, and Florida, west to Tennessee and Illinois (Middlekauff 1964). Larvae feed on leaf folds of Rubus (Smith 2008c (Walden 1907a, Middlekauff 1964, Shinohara 1985. Outbreaks in which larvae partially defoliate peach trees (Prunus persica (L.) Batsch 1801 nec Stokes 1812 nor (L.) Siebold & Zucc. 1845) and hundreds of adults can be collected via sweep netting within a few seconds have been recorded (Walden 1907b, Walden 1907a, Walden 1908a, Walden 1908b Pamphilius rileyi (Pamphiliidae) ( new state record) occurs from Connecticut south to Virginia and West Virginia, west to Michigan, Iowa, and Missouri (Middlekauff 1964, Shinohara 1985. The larval food host has been reported to be Amelanchier, though Shinohara 1985 indicated that this is based off a single male specimen that was reported to be ovipositing and so likely in error. Acordulecera dorsalis, A. mellina, and A. pellucida (Pergidae) (new state records) are widespread in eastern North America from Quebec, Maryland, Virginia, and Georgia, west to Oklahoma and Texas (Smith 1979a). Acordulecera dorsalis has been reported from Quercus, Carya, Juglans, and Castanea, while the larval food plants of A. mellina and A. pellucida are unknown (Smith 2008c).
Caliroa quercuscoccineae (Tenthredinidae) (new state record), commonly known as the scarlet oak sawfly, is widespread in eastern North America from Maine south to North Carolina, west to Minnesota, Illinois, Missouri, and Louisiana (Smith 1979a, Nordin and Johnson 1983, Boggs et al. 1999. Though the species is native, outbreaks that cause significant damage to large areas (up to 1.5 million acres) have been recorded (Matuszewski and Barry 1975, Matuszewski 1976, Matuszewski and Ward 1977, Nordin and Johnson 1983. Empria coryli (Tenthredinidae) (new state record) occurs from New Hampshire and New York, west to Wisconsin, Iowa, and Missouri (Smith 1979c). Larvae have been recorded from hazel (Corylus) (Smith 1979a, Smith 2006b).
Macrophya cassandra (Tenthredinidae) ( new state record) is known from Nova Scotia south to Georgia, west to Manitoba, South Dakota, and Montana (Gibson 1980). Larvae have been reported to feed on Carya (Smith 2006b).

Monophadnoides conspiculatus (Tenthredinidae) ( new state record) occurs from Nova
Scotia and Ontario, south to North Carolina and Tennessee (Smith 1969a).
Monophadnoides pauper (Tenthredinidae) (new state record) occurs from Labrador and Maryland west to Missouri, Kansas, Colorado, and Alberta (Smith 1969a).

Nematus abbotii (Tenthredinidae) ( new state record) is widespread in eastern North
America and has been recorded from Ontario south to Alabama and Georgia, west to Minnesota, Kansas, Iowa, and Missouri (Smith 2008b). It has been reported to feed on black locust (Robinia pseudoacacia L.) and is suspected of being univoltine as adults have only been collected during the spring, but little else is known about the species' biology (Smith 2008b).
Neopareophora litura (Tenthredinidae) (new state record), commonly called the blueberry sawfly, has been recorded from New Brunswick and Newfoundland south to Virginia and west to Illinois. It feeds on Vaccinium L., especially low-bush blueberry (V. angustifolium) on which it is occasionally a minor pest, so is likely present wherever host plants are found (Neilson 1955, Neilson 1958, Smith 1979b, Collins et al. 1994.
Pachynematus corniger (Tenthredinidae) (new state record) occurs from Newfoundland and Quebec south to Georgia, west to Minnesota, Illinois and Kansas; it has also been found in Colorado (Marlatt 1896, Crevecoeur 1922, Smith 1979b, Environment Canada 2014. Larval food plants have been reported to be Carex and possibly grasses (Smith 2006b, Smith 2008a. Paracharactus rudis (Tenthredinidae) (new state record) occurs from Maine and Quebec south to Georgia, west to Saskatchewan and Colorado (Smith 1969b).
Periclista marginicollis (Tenthredinidae) ( new state record), commonly known as the pecan sawfly, occurs from Ontario and Connecticut south to Florida, west to Iowa, Kansas, Oklahoma, and Texas. Host plants include species of Carya Nutt., including pecan ( C. illinoinensis (Wangenh.) K.Koch), on which it can occasionally cause economic damage (Smith 1969a, Smith 1979a, Smith et al. 1996, Ree and Knutson 1997, Dickey and Medina 2010.
Pristiphora banski (Tenthredinidae) (new state record) occurs from New York and West Virginia south to North Carolina, as well as Wyoming, where is feeds on Vaccinium L. (Smith 1979b, Braud et al. 2003, Smith 2006b, Smith 2008c, Smith 2008a, Barrows and Smith 2014. It is likely the species will be found in North America wherever larval host plants are located. Pristiphora chlorea (Tenthredinidae) (new state record) is widespread in North America and has been recorded from New Brunswick, south to Florida, west to Manitoba, Oregon, and Texas (Frost 1969, Smith 1979b, Smith 2006b, Smith 2008c, Smith 2008a, Barrows and Smith 2014. Larvae feed on Quercus (Smith 2006b).
Strongylogaster impressata (Tenthredinidae) (new state record) is widespread in North America and has been recorded from New Brunswick and Quebec, south to North Carolina, west to Minnesota, Colorado, and New Mexico. It feeds on ferns, including Pteridium aquilinum, though it has been shown that cyanogenic compounds within some ferns impair feeding and development (Smith 1969b, Schreiner et al. 1984.
Taxonus epicera (Tenthredinidae) ( new state record) occurs in eastern North America from New Hampshire and Ontario west to Kansas and Texas (Smith 1979a, Smith 1979b. Thrinax albidopicta (Tenthredinidae) (new state record) occurs from Nova Scotia and New Brunswick, south to Alabama, and west to Minnesota, Iowa, and Kansas (Rohwer 1910, Viereck 1916, Smith 1979b, Smith 2006b, Smith 2008a). It has historically been placed in Hemitaxonus, which was recently synonymized with Thrinax by Blank 2002. Larvae have been recorded on the ferns Onoclea sensibilis and Osmunda (Smith 1969c). Larval host plants are ferns (Smith 2008a).
Xiphydria tibialis (Xiphydriidae) (new state record) is widespread in eastern North America and occurs from Nova Scotia and Quebec south to Florida, west to Wisconsin, Illinois, and Kansas (Smith 1976). Larvae bore in small limbs of a number of trees, including Acer, Betula, Fagus, Malus, Quercus, Tilia, and Ulmus (Smith 2008c).

Discussion
Thirty-one species are recorded as new to Arkansas. The majority (23 species) are wideranging species found throughout North America east of the Rocky Mountains and many have been collected in states bordering Arkansas. While it is unsurprising that they also occur in Arkansas, they serve to highlight how little collecting and survey work some groups have historically received in the state.
Specimens of eight species -Monophadnoides conspiculatus, Neopareophora litura, Orussus minutus, O. terminalis, Pamphilius ocreatus, P. rileyi, Strongylogaster remota, and Thrinax multicinctus -represent significant western range extensions of hundreds of kilometers from previously known localities. However, it is likely that they do not represent disjunct populations confined to the Interior Highlands and that increased sampling effort in eastern North America will uncover additional specimens from new localities.
Malaise traps collected the highest number of species (28). Green Lindgren funnel traps collected the second highest number of species (17), which while fewer than Malaise traps is higher than all other trap types; they also collected slightly more specimens per trap than Malaise traps. Most species were represented by 10 or fewer specimens, while only four species were represented by 25 or more specimens.