SPECIES FACT SHEET Common Name: Marsh damsel bug Scientific Name: Nabis propinquus = Nabicula propinq ua (Reuter, 1872) Phylum: Mandibulata Class: Insecta Order: Hemiptera Family: Nabidae Subfamily: Nabinae Genus: Nabis Subgenus: Nabicula Taxonomic Note: The Nabicula “group” has been treated both as its own genus and as a subgenus of Nabis by various authors. Since the most recent treatments (Kerzhner 1996, Kerzhner & Henry 2008) present this group as a subgenus of Nabis, the current name for the marsh damsel bug is Nabis propinquus. Conservation Status: Global Status (2001): G5 - Secure National Status (United States): NNR State/Province Statuses: Oregon (S2), British Columbia (S1) (NatureServe 2012). Technical Description: Nabis propinquus is a slender brown terrestrial hemipteran in the Nabidae (damsel bug) family (Applegarth 1995). This family is comprised of a small group of important generalist predators best known for their frequent use in biological control programs to manage insect pests on agricultural crops (Kerzhner & Henry 2008). Like other heteropterans, the mouthparts are of a unique piercing-sucking type, and the basal portion of the front wing is thickened and leathery, while the apical portion is membranous. Members of this family are mostly dull-brownish, occurring in both short-winged (brachypterous) and long-winged (macropterous) forms (Maw et al. 2000). In North America, this family is distinct from other true bugs by the front femora being slightly enlarged (swollen), and the membrane of the hemelytra (when developed) having a number of small cells around the margin (Triplehorn & Johnson 2005, Arnett 2000). This family is further described as follows (Maw et al. 2000): Ocelli are present; the labium is four-segmented. The fore legs have a dense pad of hairs (fossula spongiosa) at the distal end of the tibia. The costal fracture of the hemelytra may be present or absent. The membrane of the hemelytra has two or three elongate cells, and usually has emanating veins 1 and a stub. The abdomen has three dorsal scent gland openings between terga 4/5, 5/6, and 6/7. The male genitalia are usually symmetrical (Maw et al. 2000). The Nabidae family contains two subfamilies, 11 genera, and 39 species in North American (Kerzhner & Henry 2008, Henry 2009). The marsh damsel bug belongs to the Nabinae subfamily, Nabis genus. Members of this genus are 6 to 9 mm in length, slender, light to dark brown, and have the front femora slightly enlarged (BugGuide 2013). The Nabis genus is further divided into subgenera, including the subgenus Nabicula, to which this species belongs. There are a total of six species in the Nabicula subgenus in North America (Henry & Latttin 1988). Among these, Nabis propinquus is highly divergent morphologically, to the extent that it has been separated from the other five species into its own subgroup, the former subgenus Limnonabis. (This subgenus became obsolete when all of the species in the former Nabicula genus were shifted to Nabis, and Nabicula was reduced to subgenus status (Kerzhner 1996)). Nomenclature aside, N. propinquus is distinguished from other North American members of the Nabicula subgenus by the following characters: diverging posterior lobe of the head; recurved connexivum of the male; and the presence of three or four sclerites in the aedeagus (Asquith & Lattin 1990). In addition, the yellowish, elongate, parallel-sided abdomen help identify this species among its close relatives (BugGuide 2013). According to Harris (1928), this species is the longest of the American Nabids, and is easily recognized by its form. A full description of this species is as follows (Harris 1928): Slender, greatly elongate, pubescent; testaceous, an interrupted median line extending from base of tylus to tip of abdomen and a broad stripe on either side of body for its full length fuscous. Femora fusco-maculate. Head long, more than a third longer than broad, faintly widened behind the eyes. Eyes only moderately large, the length of one equal to the width of vertex. Ocelli prominent. Antennae long, darkened distally, segment I much longer than the width of head through eyes. Rostrum with segments II and III subequal, each much shorter than antennal segment I. Pronotum with long collar, the anterior lobe with the cicatrices darkened; the posterior lobe finely punctulate. Scutellum with a prominent, semicircular, shiny spot on either side basally. Abdomen above with an interuppeted fuscous stripe to either side of the median dark line; the connexivum horizontal. Legs long, the front and mid femora thickly clothed beneath with short hooked hairs and more thinly so with shorter piceous spinules; front femora about six times as long as thick; posterior tibiae clothed with short hairs. Male narrower than female, the genital segments long, prominent, the clasper distinctive. Female with abdomen rather sharply angular at apex, the apical margin of last abdominal segment emarginated. Brachypterous form: pronotum longer than broad, the lobes indistinctly marked off, the basal margin 2 roundly emarginated (more strongly so in male). Hemelytra extending almost to apex of second abdominal segment, obtusely rounded behind, without membrane. Length 9-12 mm; width 1.32-1.68 mm (at abdomen, 1.32-2.82 mm). Macropterous form: pronotum broader than long, the posterior lobe arched. Hemelytra somewhat translucent, finely punctulate; membrane extending onto base of last abdominal segment. Length 12 mm; width 2.1 mm (Harris 1928). Although N. propinquus exhibits both macropterous (fully-winged) and brachypterous (short-winged) morphs, the macropterous condition is very rare relative to the brachypterous morph. For example, only 2 of the 45 females and 1 of the 29 males examined by Asquith & Lattin (1990) were macropterous. In addition to wing polymorphism, this species also exhibits sexual dimorphism in which the females are larger than males and differ in various body proportions (see Asquith & Lattin 1990 for details). See Attachment 4 in the Appendix for a photograph and illustrations of the male and female macropterous and brachypterous morphs of this species. The genitalia of this species are illustrated in Asquith & Lattin (1990). Immature: Immature nabids have a long, slender, 4 segmented labium arising from the anterior part of the head (Elvin & Sloderback 1984). They also have long, slender antennae, apical tarsal claws, and usually 3-4 pairs of scent glands present on the abdomen. The labium is not located in a rostral groove in the gular region (Elvin & Sloderback 1984). The nymphal instar of a particular immature nabid can be determined by assessing the amount of wing pad development (see Elvin & Sloderback 1984 for details). Within the family, there is only limited information on identifying nymphal stages to genus or species. Nymphs of the genus Nabis are distinguished from the related genus Hoplistoscelis largely by body shape (Nabis is generally narrower than Hoplistoscelis) (Elvin & Sloderback 1984). A description of the nymph of N. propinquus is not known. Life History: Nabid bugs are aggressive generalist predators, feeding on a wide variety of small arthropods including caterpillars, aphids, and plant bugs (Marshall 2006, Triplehorn & Johnson 2005). Nabid nymphs are active shortly after hatching and begin feeding immediately, often on prey considerably larger than themselves (Lattin 1989). The phenology of this species is poorly known. Known records in Oregon are from July and September (OSAC 2013, Asquith & Lattin 1990). Rangewide, this species has been collected from June to September, with the majority of records in July and August (Asquith & Lattin 1990). Nabids tend to overwinter as adults (Applegarth 1995, Lattin 1989). Members of this genus can be 3 univoltine or multivoltine, producing one to five generations per year, depending on location (BugGuide 2013). Dispersal behavior in N. propinquus has not been examined. The macropterous (fully-winged, flight capable) form of some adult nabids are known to move considerable distances (Lattin 1989), probably in response to instable habitats or food sources. Since this species is known primarily in the brachypterous (reduced wing) form (Asquith & Lattin 1990), its dispersal potential is probably limited by the reduced flight ability of this wing morph. Overall, many aspects of this species biology are in need of study, including dietary preferences, mating behavior, oviposition site selection, number of eggs laid, development time, life span, number of generations per year, dispersal behavior, and overwintering behavior and habitat. Range, Distribution, and Abundance: This species is widely distributed across North America. In Canada it is known from Alberta, British Columbia, Northwest Territories, Ontario, Quebec, Saskatchewan. Manitoba, New Brunswick, Prince Edward Island and Nova Scotia (Henry & Lattin 1988, Scudder 1994, Maw et al. 2000, Asquith & Lattin 1990). In the United States, it is reported from Illinois, Iowa, Maine, Maryland, Massachusetts, Michigan, North Dakota, Nebraska, Kansas, Oklahoma, New Jersey, New York, Ohio, Oregon, South Dakota, and Wisconsin (Henry & Lattin 1988, reviewed in Kerzhner & Henry 2008, Applegarth 1995, Asquith & Lattin 1990, GBIF 2013). Since this species has not yet been found in the arid Intermountain west region, the Oregon populations are considered disjunct from the main distribution (Asquith & Lattin 1990). See the Conservation Considerations section, below, for further discussion of the biogeography of this species. In Oregon, this species appears to be restricted to the Pacific Coast (Asquith & Lattin 1990). Known records in Oregon are from two separate areas along the coast, in Tillamook and Coos Counties (Applegarth 1995, OSAC 2013). This species has not been found in the Willamette Valley or Puget Trough areas of Oregon and Washington (Asquith & Lattin 1990). Asquith & Lattin (1990) note the possibility that this species may occur in coastal marshes of Washington. BLM/Forest Service lands: This species is Suspected on Siuslaw National Forest based on the presence of large coastal stretches of Siuslaw NF land between and in the vicinity of the known Oregon sites. It is also Suspected by Salem and Eugene BLM Districts, based on their coastal lands near the known sites. Abundance: Abundance estimates for this species have not been conducted. Most collections have been of one to two individuals, although one Oregon collection was of 16 individuals (12 male, 4 female) (Asquith & Lattin 1990). 4 Habitat Associations: This species lives in wet environments where it is found on the ground or in low vegetation (Asquith & Lattin 1990). Rangewide, it occurs in moist, riparian or marshy habitats (Asquith & Lattin 1990). According to Harris (1928), this species frequents the edges of marshes and ponds where reeds and sedges flourish. In Oregon, coastal salt marshes are the only known habitat for this species (Applegarth 1995). In British Columbia this species is also known from beach habitat (Scudder 1994). Threats: Threats to this species include conversion and fragmentation of coastal marshes and wetlands, loss and degradation of sand dune systems and riparian areas, and encroachment of exotic vegetation. Conservation Considerations: According to Asquith & Lattin (1990), the origin of the highly disjunct Oregon coast population is perplexing. Lattin (1966) suggested that this population might be an introduction from the eastern United States, but Asquith & Lattin (1990) believed this to be unlikely, considering the restricted habitat of this species and the fact that the Oregon population displays a slightly different genitalic structure than the eastern populations (see Asquith & Lattin 1990 for further details). In their thorough examination of the biogeography of this species, Asquith & Lattin (1990) present the following conclusions (slightly paraphrased): We hypothesize that this species (or its direct ancestor) was present in North America during the Eocene period, at which time there were few barriers to impede East-West movement and a contiguous distribution across the northern latitudes would have been possible. Beginning in the Miocene, orogenic activity in western North America resulted in topographical geographic barriers and, perhaps more importantly, increased aridity in the Intermountain region. Because N. propinquus lives in moist, riparian habitats, this dramatic change in climate and terrain would have restricted mobility and could have led to the extinction of many, or all of the intervening populations. This is merely a hypothesis of course, and we cannot actually date the separation of the west coast population and it is possible that it is a much more recent event. The climate of the northern United States during the Pleistocene periods was much more mesic than the present, with abundant, large shallow lakes and marshes throughout the Great Basin province. This would have provided abundant habitats for N. propinquus from the Rocky Mountains west [to the Pacific Coast]. With the advent of the Hypsithermal and higher temperatures, these habitats in this area largely disappeared, which could have left the coastal population isolated from those north and east of the Rockies. This hypothesis would be 5 corroborated if relictual populations of N. propinquus were found in isolated marshes in the northern Great Basin (Asquith & Lattin 1990). Inventory: Unlike some of the commonly encountered species of Nabis species that occur on vegetation in a variety of habitats, Nabis propinquus is rare in museum collections (Asquith & Lattin 1990), probably due to a combination of rarity, restricted habitat, and limited sampling effort. Further surveys are needed for this species in Oregon, and will be critical in evaluating its current status and habitat use in the region. Initial surveys are recommended in coastal salt marshes on Siuslaw National Forest in the area between and in the vicinity of known sites. This species was last documented at both the Tillamook and Coos County coastal sites in 1988 (OSAC 2013, Asquith & Lattin 1990). In British Columbia, this species is also considered a priority for inventory and descriptive research, due to being very rare or endangered at both the provincial and national levels (Scudder 1996). Management: Protect all known and potential sites from practices that would adversely affect this species or its habitat. Management designed to limit the encroachment of exotic vegetation may be important at some sites. Research: Many aspects of this species’ biology are in need of study including food preferences, mating behavior, oviposition site selection, number of eggs laid, development time, life span, number of generations per year, and overwintering behavior. Prepared by: Sarah Foltz Jordan, Xerces Society for Invertebrate Conservation Date: 9 April 2013 Edited by: Sarina Jepsen, Xerces Society for Invertebrate Conservation Date: 10 April 2013 6 ATTACHMENTS: (1) References (2) List of pertinent or knowledgeable contacts (3) Map of known records in Oregon (4) Photograph and illustrations of this species (5) Survey protocol for this species ATTACHMENT 1: References. Applegarth, J.S. 1995. Invertebrates of special status or special concern in the Eugene district. U.S. Department of the Interior, Bureau of Land Management. 126 pp. Arnett, R.H., Jr. 2000. American Insects, A Handbook of the Insects of America North of Mexico, 2nd edition, CRC Press, Boca Raton, FL. 1003 pp. Asquith. A. & J.D. Lattin. 1990. Nabicula (Limnonabis) propinquus (Reuter) (Heteroptera: Nabidae): dimorphism, phylogenetic relationships and biogeography. Tijdschrift voor Entomologie 133: 3-16. Blinn, R.L. 1995. Phorticus collaris Stål: A nabid new to Eastern North America (Heteroptera: Nabidae). Journal of the New York Entomological Society 103: 216-218. BugGuide 2013. Nabis. Available at: http://bugguide.net/node/view/12327 (Accessed 4 April 2013). Elvin, M.K. and P.E. Sloderbeck 1984. A key to the nymphs of selected species of Nabidae (Hemiptera) in the southeastern USA. Fla. Entomol. 67: 269-273. Global Biodiversity Information Facility (GBIF). Nabicula propinquus. Available at: http://data.gbif.org/occurrences/350330209 and: http://data.gbif.org/occurrences/350330207 (Accessed 4 April 2013). Harris, H.M. 1928. A monographic study of the Hemipterous family Nabidae as it occurs in North America. Entomologica Americana 9: 1-97. Henry, T.J. 2009. Biodiversity of Heteroptera. Chapter 10. Pp. 223–263 In: Foottit, R.G.; Adler, P.H. (eds) 2009: Insect Biodiversity: Science and Society. Wiley-Backwell, Oxford, U.K. 656 pp. Henry, T.J. and J. Brambila. 2003. First report of the neotropical damsel bug Alloeorhynchus trimacula (Stein) in the United States, with new records for two 7 other nabid species in Florida (Heteroptera: Nabidae: Prostemmatinae). Proceedings of the Entomological Society of Washington 105: 801-808. Henry, T.J. and J.D. Lattin. 1988. Family Nabidae Costa. 1853. The damsel bugs, pp. 508-520. In Henry. T. J. and R. C. Froeschner. eds. Catalog of the Heteroptera, or true bugs, of Canada and the continental United States. E.J. Brill, Leiden and New York. 958 pp. Kerzhner I.M. and T.J. Henry. 2008. Three new species, notes and new records of poorly known species, and an updated checklist for the North American Nabidae. Proceedings of the Entomological Society of Washington 110(4): 988– 1011. Kerzhner I.M. 1996. Family Nabidae. Catalogue of the Heteroptera of the Palaearctic Region 2: 84-107. Lattin, J.D. 1989. Bionomics of Nabidae. Annual Review of Entomology 34: 383–400. Marshall, S. 2006. Insects: their natural history and diversity: with a photographic guide to insects of eastern North America. Firefly Books, Buffalo, New York. 718 pp. Maw, H.E.L., Foottit, R.G., Hamilton, K.G.A. and G.G.F. Scudder. 2000. Checklist of the Hemiptera of Canada and Alaska. NRC Research Press. Ottawa, Ontario. 220 pp. NatureServe. 2013. “Nabicula propinquus.” NatureServe Explorer: An online encyclopedia of life [web application]. Feb. 2009. Version 7.1. NatureServe, Arlington, Virginia. Data last updated: October 2012. Available at: www.natureserve.org/explorer (Accessed 11 March 2013). Oregon State University Arthropod Collection (OSAC). 2013. Specimen data gathered by Ashley Clayton, contractor for the Xerces Society. Reuter, 0.M., 1872. Nabidae novae et minus cognitae. Bidrag till Nabidernas kannedom. Ofversigt af Kongliga Vetenskaps -Akademiens Forhandlingar 29(6): 79-96. Scudder, G.G.E. 1994. An annotated systematic list of the potentially rare and endangered freshwater and terrestrial invertebrates in British Columbia. Entomological Society of British Columbia, Occasional Paper 2: 1-92. Scudder, G.G.E. 1996. Terrestrial and freshwater invertebrates of British Columbia: priorities for inventory and descriptive research. Res. Br., B.C. Min. 8 For., and Wildl. Br., B.C. Min. Environ., Lands and Parks. Victoria, B.C. Work. Pap. 09/1996. Triplehorn, C. and N. Johnson. 2005. Introduction to the Study of Insects. Thomson Brooks/Cole, Belmont, CA. 864 pp. ATTACHMENT 2: List of pertinent or knowledgeable contacts Michael Schwartz, Research Affiliate, Insect Biosystematics, Canadian National Collection of Insects. Thomas Henry, Smithsonian National Museum of Natural History, Washington DC. John D. Lattin, Oregon State University, Corvallis, Oregon. 9 ATTACHMENT 3: Map of known records in Oregon and Washington Known records of Nabis propinquus in Oregon, relative to BLM and Forest Service lands. 10 Known records of Nabis propinquus in Tillamook County, Oregon, showing proximity to Suislaw National Forest. 11 ATTACHMENT 4: Photograph of this species Nabis propinquus brachypterous adult female, dorsal view. Photograph by Ashley Clayton for The Xerces Society, used with permission. 12 Nabis propinquus brachypterous male adult, dorsal view. Illustration from Asquith & Lattin (1990). 13 Nabis propinquus adults, dorsal view. Macropterous female (A), macropterous male (B). Illustration from Asquith & Lattin (1990). ATTACHMENT 5: Survey Protocol for this species Survey Protocol: Nabis propinquus Where: This species is known from very few collections in Oregon, probably due (at least in part) to limited sampling effort. Further surveys are needed for this species in Oregon, and will be critical in evaluating its current status and habitat use in the region. Initial surveys are recommended on Siuslaw National 14 Forest in the area between and in the vicinity of known sites. This species was last documented at both the Tillamook and Coos County coastal sites in 1988 (OSAC 2013, Asquith & Lattin 1990). Surveys in Oregon should target coastal salt marshes, the only known habitat in the state. Rangewide, this species inhabits moist, riparian areas. When: Surveys for this species are recommended in summer. Known records in Oregon are from July and September (OSAC 2013, Asquith & Lattin 1990). Rangewide, this species has been collected from June to September, with the majority of records in July and August (Asquith & Lattin 1990). Nabids tend to overwinter as adults (Applegarth 1995, Lattin 1989). Members of this genus can be univoltine or multivoltine, producing one to five generations per year, depending on location (BugGuide 2013). How: Visual searches and sweep-netting in known habitat (coastal marshes) are probably the most appropriate survey method at this time. Window traps (e.g., plexiglass panels in tubs of soapy water) may also be useful, as these traps (placed 1 to 7 m off the ground) have been successful at collecting other members of this family (Lattin 1989). Ultraviolet light traps have also been used to collect nabids (Blinn 1995), and many nabid species are known to be attracted to visible light (BugGuide 2013). After capture, voucher specimens should be placed immediately into a kill jar until they can be pinned. A field catch can also be temporarily or permanently stored in 75% ethyl alcohol but the alcohol will cause some colors to fade (Triplehorn & Johnson 2005). Adult specimens should be pinned through the scutellum (Triplehorn & Johnson 2005). Juveniles are best preserved in vials containing 75% ethyl alcohol. Collection labels should include the following information: date, collector, detailed locality (including geographical coordinates, mileage from named location, elevation, etc.), and detailed habitat/host plant. Complete determination labels include the species name, sex (if known), determiner name, and date determined. Field identification is possible by those familiar with heteropteran taxonomy, and can be accomplished by non-experts who have examined and become familiar with museum specimens. This species is identified using characteristics provided in the Species Fact Sheet. Confirmation of field identifications should be done by taxonomic experts with experience identifying nabids. 15