University of Nebraska - Lincoln DigitalCommons@University of Nebraska - Lincoln

Panhandle Research and Extension Center Agricultural Research Division of IANR

2002

Life History and Laboratory Rearing of Pseudometapterus umbrosus (Heteroptera: Reduviidae) with Descriptions of Immatures Stages

Jeffrey Bradshaw University of Nebraska-Lincoln, [email protected]

J. E. McPherson Southern Illinois University, [email protected]

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Bradshaw, Jeffrey and McPherson, J. E., "Life History and Laboratory Rearing of Pseudometapterus umbrosus (Heteroptera: Reduviidae) with Descriptions of Immatures Stages" (2002). Panhandle Research and Extension Center. 36. https://digitalcommons.unl.edu/panhandleresext/36

This Article is brought to you for free and open access by the Agricultural Research Division of IANR at DigitalCommons@University of Nebraska - Lincoln. It has been accepted for inclusion in Panhandle Research and Extension Center by an authorized administrator of DigitalCommons@University of Nebraska - Lincoln. SYSTEMATICS Life History and Laboratory Rearing of Pseudometapterus umbrosus (Heteroptera: Reduviidae) with Descriptions of Immatures Stages

1 J. D. BRADSHAW AND J. E. MCPHERSON

Department of Zoology, Southern Illinois University at Carbondale, Carbondale, IL62901

Ann. Entomol. Soc. Am. 95(2): 192Ð200 (2002) ABSTRACT The life history of the emesine reduviid Pseudometapterus umbrosus (Blatchley) was studied in southern Illinois from February 1999 to November 2000, and the immature stages were described. The bug also was reared from egg to adult under controlled laboratory conditions. This univoltine species occurred on the rock faces of sandstone bluffs often in association with plants and spider webs. Adults and nymphs preyed primarily on small ßies. Adults overwintered within fallen leaves on the bluff outcroppings, emerged in early April, and began feeding and reproducing shortly thereafter. Eggs were found from early May to late October and were deposited on the leaves and stems of Heuchera parviflora Bartling. Nymphs were found from early June to early September. New adults appeared in mid-July and remained active until early October. This species was reared in the laboratory on adult Drosophila sp. at 26 Ϯ 3.0ЊC under a photoperiod of 16:8 (L:D) h. The incubation period averaged 13.25 d; eyespots appeared in Ϸ7 d. The Þve stadia averaged 11.86, 11.77, 12.44, 15.35, and 18.88 d, respectively. The instars can be distinguished by differences in several anatomical features including body length and width.

KEY WORDS Pseudometapterus umbrosus, life history, laboratory rearing, descriptions, nymphs

THE EMESINE REDUVIIDS, a group of assassin bugs with emesineÕs previously known Florida distribution slender bodies and appendages, have a cosmopolitan (McPherson 1991a, 1991b). The only life history in- distribution (Wygodzinsky 1966). Although these pre- formation then available was that it had been beaten daceous insects occur throughout North America, from fallen leaves of a royal palm in a dense hammock they usually are not collected in large numbers. As a on Paradise Key, FL(Blatchley 1926), and collected result, the biology of most species is poorly known from Spanish moss (Wygodzinsky 1966). (McPherson 1991a). Emesines are known to occur Hagerty and McPherson (1999) reported on a sur- beneath loose bark; in bases of tufts of grasses; in vey of the assassin bugs of southern Illinois, includ- bunches of dead leaves; in barns, dwellings, cellars, ing P. umbrosus. They collected adults of this species and outhouses (Blatchley 1926); at considerable from spider webs and plants (i.e., Heuchera parviflora heights on bushes and trees; within caves; and in as- Bartling) on sandstone bluffs (n ϭ 41) and on spider sociation with “psocid” and spider webs (Wygodzin- webs on limestone bluffs (n ϭ 2) in Jackson and Union sky 1966). counties, respectively. Specimens were taken from 13 Pseudometapterus Wygodzinsky, an emesine genus, April to 24 November, but most (69.8%, n ϭ 30) were contains 15 species, all with a Nearctic or Neotropical collected during August and September; these data distribution (Wygodzinsky 1966). Of these, three oc- suggest this species is univoltine and overwinters as cur in America north of Mexico: P. butleri Wygodz- adults. insky in Arizona, P. wygodzinskyi (Elkins) in Texas, The population in Jackson County was discovered and P. umbrosus (Blatchley) in Florida (Froeschner in Little Grand Canyon, which lies 14.5 km SW of 1988) and Illinois (McPherson 1991a). Murphysboro within the Shawnee National Forest. The discovery of P. umbrosus in Illinois was based on Little Grand Canyon spans 555.2 ha and consists of two male adults deposited in the Southern Illinois ridge tops sloping to large Pennsylvanian sandstone University Entomology Collection. They had been bluffs. The bluffs, in turn, encompass a 1,676.4 m long collected on 27 July 1972 from the LaRue-Pine Hills canyon that reaches a depth of 33.5 m (Hopkins and Ecological Area (now LaRue-Pine Hills Research Nat- Simon 1975, Mohlenbrock 1977). Flora on the ridge ural Area), Union County, a dramatic extension of this tops and slopes consists mostly of a mesic upland community of white oak (Quercus alba L.), red oak 1 Current address: Department of Entomology, Iowa State Univer- (Quercus rubra L.), black oak (Quercus velutina sity, 103 Insectary, Ames, IA 50011Ð0001 (e-mail: mcpherson@ Lamarck), shagbark hickory [Carya ovata (Miller)], zoology.siu.edu). and pignut hickory (Carya glabra Miller) (Mohlen-

0013-8746/02/0192Ð0200$02.00/0 ᭧ 2002 Entomological Society of America March 2002 BRADSHAW AND MCPHERSON:LIFE HISTORY OF P. umbrosus 193 brock 1977). The canyon ßoor ßora consists mostly of toweling (Ϸ11.4 cm long, 6.0 cm wide), folded longi- a mesic lowland community of beech (Fagus grandi- tudinally to prevent the strip from rolling up when folia Ehrhart) and tulip poplar (Liriodendron tulipif- damp, was suspended inside the jar and secured at the era L.) (Mohlenbrock 1977). upper end between the discs of toweling and screen- The study site, within a chute of Little Grand Can- ing. This strip increased surface area for walking and yon, is located in the Gorham quadrangle 7.5Ј series absorption of excrement. (topographic) (T10S, R3W, W1/2 of NW1/4 of sec- The cages were examined daily for eggs, which were tion 1). The ridge slopes at this location support a removed and placed on moistened Þlter paper (Eaton- mesic upland community of black oak, shagbark hick- Dikeman, grade 613, Mount Holly Springs, PA) on the ory, false shagbark [C. ovalis (Wangenheim)], and bottom of a petri dish (Ϸ9 cm diameter, 2.0 cm deep) black maple (Acer nigrum Michaux f.). The bluff faces and covered with the lid. support communities of alumroot (H. parviflora), Newly emerged Þrst instars were placed on moist- christmas fern [Polystichum acrostichoides (Michaux)], ened Þlter paper in petri dishes prepared similarly to marginal shield-fern [Dryopteris marginalis (L.)], those for eggs except that each dish was closed with a mosses [e.g., Leucobryum glaucum (Hedwig) Ång- piece of interlock cloth (2-mm2 mesh) secured with stro¨m, Atichum angustatum (Bridel) Buchner and the lid. The cloth provided additional surface area for Shimper, Tetraphis pullucida Hedwig, and Bryoander- walking and molting; nymphs were unable to molt sonia recognitum (Hedwig) Lindberg], and a leafy successfully unless suspended upside down from the liverwort (Isotachis sp.). The chute supports a tem- cloth. No more than two nymphs were placed in each porary rain-fed stream that sustains persistent water dish to reduce the risk of cannibalism. Nymphal pairs pools on the streambed. were separated at subsequent molts as necessary to In this article, we present further information on the allow individual records of development. Þeld life history of P. umbrosus in Little Grand Canyon The numbers of nymphs completing the fourth and and include information on laboratory rearing and Þfth stadia were low. Therefore, we supplemented descriptions of the immature stages. these data with Þeld-collected third instars that were reared to adults under the same conditions as nymphs reared from eggs to adults. Materials and Methods Eggs, nymphs, and adults were kept in incubators Life History. The Þeld study of P. umbrosus was maintained at 26 Ϯ 3.0ЊC and a photoperiod of 16:8 conducted from February 1999 to November 2000. (L:D)h (Ϸ2,800 lux). Filter paper in the ovipositional The Þrst year we determined the feasibility of the cages and petri dishes was moistened daily. Adults study. These insects are difÞcult to see because of their were fed 9Ð11, nymphs 5Ð10, wingless, debilitated slender bodies and appendages, relatively small size Drosophila sp. adults per day. (e.g., Þrst instars averaged Ͻ3.60 mm in length [un- Descriptions of Immature Stages. The description published data]), and general inactivity. However, we of each stage is based on 10 Þeld-collected individuals found that individuals tended to congregate on the preserved in 70% ethanol. Drawings were made with small sandstone outcroppings of the bluffs, making a Wild MSA drawing tube (E. Leitz, Rockleigh, NJ), their detection easier. Thus, we conducted a detailed measurements (in millimeters) with an ocular micro- study during 2000. The study site was restricted to two meter. Scanning electron micrographs were used as areas (14.0 by 2.4 m, 20 by 2.4 m) on the chute walls aids in the descriptions of the egg and Þrst instar. separated by 140 m. Statistics. Averages are expressed as means Ϯ SE; Counts of nymphs and adults and notes on insect standard errors of Ͻ0.005 are listed as 0.00. Field- activities were taken weekly from early February to collected and laboratory-reared third instars were early November, before the bugs emerged from and reared to adults and their fourth and Þfth stadia com- after they entered overwintering sites. Specimens pared with a Mann-Whitney U test; level of signiÞ- were collected by handpicking but generally were cance was 0.01. The statistical test was chosen primar- released to prevent overcollecting. Those that could ily because the samples were small and of unequal size. not be identiÞed to instar were preserved in 70% EtOH and examined in the laboratory. Prey (deÞned as in- Results and Discussion dividuals held by prothoracic legs) were collected and preserved in 70% EtOH. Potential overwintering sites Life History. Pseudometapterus umbrosus overwin- (e.g., leaf litter) were examined periodically from tered as adults within piles of leaves that had fallen early October 1999 to early April 2000. from the slope overhead and accumulated around the Laboratory Rearing. Seven adults were collected on bases of plants (e.g., P. acrostichoides) on bluff out- 6 November 1999 from beneath leaves at Little Grand croppings (n ϭ 3 adults [30 October 1999]; n ϭ 7[6 Canyon, brought to the laboratory, and placed in four November 1999]; n ϭ 2 [13 November 1999]; n ϭ 1[5 ovipositional cages (three cages, one male, one fe- February 2000]). They emerged in early April and male; one cage, one female). Each cage consisted of a began feeding and copulating shortly thereafter. Cop- 1-pt (Ϸ0.47-liter) Mason jar with a disc of moistened ulation (13 pairs) was observed from early May to Þlter paper on the bottom; it was closed with discs of early June on bare rock outcroppings, or on plants paper toweling and wire screening secured with the attached to outcroppings, on the bluff face. On one band of the two-piece Mason jar lid. A strip of paper occasion, a male attempted to copulate with a female 194 ANNALS OF THE ENTOMOLOGICAL SOCIETY OF AMERICA Vol. 95, no. 2

Table 1. Feeding records of P. umbrosus in Little Grand Canyon, Jackson County, IL, during 1999 and 2000

No. of Taxon specimens Araneida Theridiosomatidae Theridiosoma gemmosum (L. Koch)a 1 Theridiidae Achaearanea sp.a 1 Collembola Entomobryidae Tomocerus minor (Lubbock)b 2 Lepidoptera Tineidae Chloropleca sp.c (species undescribed) 1 Diptera Ceratopogonidaec 3 Palpomyia sp.c 1 Mycetophilidae Trichonta sp.c 1 Cecidomyiidaec 12 Fig. 1. Field life cycle of P. umbrosus. Percentage in each Chironomidae Orthocladinae sample of total individuals of same stage collected during the c 2000 season at Little Grand Canyon, Jackson County, IL. Bar Thienemaniella xena Roback 1 Chironominae for eggs only indicates presence. Chironominic 1 Tanypodinae Pentaneurinic 1 already in copula. It attempted to dislodge the copu- Tipulidaec 1 lating male by nudging its abdomen; unsuccessful, the Limonia sp.c 2 intruding male left. Limonia (Dicranomyia) sp.c 1 Eggs were found from early May to late October Drosophilidae Drosophila sp.c 1 (Figs. 1 and 2). However, because fertile, infertile, and Sciaridae hatched eggs could not be distinguished easily, sea- Lycoriella sp.c 1 sonal changes in ovipositional frequency could not be Culicidae c determined. Eggs were deposited singly on the leaves Culex restuans Theobald 1 and stems of H. parviflora with an ovipositional time a Ϸ Immature. of 1Ð3 min. On one occasion, a female was observed b Subadult or adult. with her metathoracic legs on the bare rock face of the c Adult. bluff and mesothoracic legs grasping and pulling an attached H. parviflora leaf (the role of her prothoracic legs was unclear, although she appeared to be stroking of her abdomen to the underside of the leaf and de- the leaf with them). As she did so, she touched the tip posited an egg. Two adult males were found with an egg attached to their metathoracic legs. First instars were found primarily from early June to late July, second instars from early June to early Au- gust, third instars from late June to mid-August, fourth instars from late June to late August, Þfth instars from early July to early September, and adults from early April to late June and mid-July to early October (Figs. 1 and 2). Adults were found on cribellate (n ϭ 166) and noncribellate (n ϭ 7) spider webs, on leaves and stems of H. parviflora (n ϭ 238), within dead fronds of P. acrostichoides (n ϭ 6), and on the barren substrate interspersed with patches of mosses and lichens (n ϭ 308). Nymphs were observed on webs of cribellate spiders (n ϭ 847), bare substrate interspersed with patches of mosses and lichens (n ϭ 320), and leaves and stems of H. parviflora (n ϭ 251). Pseudometapterus umbrosus preyed on a variety of small arthropods but preferred ßies, particularly ce- cidomyiids (Table 1). On one occasion, a noncribel- Fig. 2. Field life cycle of P. umbrosus. Percentage in each late spider, Achaearanea tepidariorum (Koch), was sample of individuals of each stage collected during the 2000 observed feeding on a Þfth instar. On another occa- season at Little Grand Canyon, Jackson County, IL. Bar for sion, an adult male was observed with its beak inserted eggs only indicates presence. into a dead ßower of H. parviflora. March 2002 BRADSHAW AND MCPHERSON:LIFE HISTORY OF P. umbrosus 195

Table 2. Duration (in days) of each immature stage of white, almost transparent upon emerging, and dark- P. umbrosus under controlled laboratory conditions ened only after feeding. The Þrst, second, third, fourth, and Þfth stadia No. Cumulative Stage completing Range Mean Ϯ SE averaged 11.86, 11.77, 12.44, 15.35, and 18.88 d, mean age stadium respectively. The total developmental period aver- Egga 122 9Ð15 13.25 Ϯ 0.09 13.25 aged 83.55 d (Table 2). Highest mortality occurred Nymphb during the second stadium, primarily resulting from 1st instar 72 7Ð24 11.86 Ϯ 0.39 25.11 incomplete ecdysis. 2nd instar 39 7Ð30 11.77 Ϯ 0.79 36.88 Comparisons of the fourth and Þfth stadia of labo- 3rd instar 27 7Ð21 12.44 Ϯ 0.77 49.32 4th instar 17 9Ð25 15.35 Ϯ 1.14c 64.67 ratory-reared and Þeld-collected third instars showed 5th instar 8 10Ð31 18.88 Ϯ 2.64d 83.55 that each stadium was shorter for Þeld-collected in- Nymphe dividuals (Table 2). This strongly suggests that Dro- 4th instar 16 6Ð18 10.81 Ϯ 0.98c Ñ sophila sp. was not an ideal food source, at least for Ϯ d 5th instar 16 9Ð17 11.44 0.56 Ñ young instars. a 359 eggs were laid. Adults appeared to drink water from the Þlter paper b Reared from egg. 2Ð5 min after the paper was moistened. They also fed c SigniÞcantly different; MannÐWhitney U ϭ 56.5, P ϭ 0.002. upon adults of Drosophila sp. within 15 min after ßies d SigniÞcantly different; MannÐWhitney U ϭ 18.5, P ϭ 0.003. had been introduced. e Field-collected as third instars. Precopulatory behavior was observed on the lid, paper toweling, and Þlter paper. Courtship began with Pseudometapterus umbrosus is univoltine in south- the male approaching the female from behind, peri- ern Illinois as evidenced by the peaks in abundance of odically antennating her abdomen and meta- and me- the adults and nymphs (Figs. 1 and 2). sothoracic legs before mounting her. He then grasped Laboratory Rearing. The three pairs of overwin- the lateral margin of her pro- or mesothorax with his tered adults reproduced, with the females depositing prothoracic legs. At this point, his abdomen crossed 86, 92, and 181 eggs of which 23, 46, and 53 hatched, hers diagonally. He lowered the tip of his abdomen so respectively. The unpaired female laid 41 eggs, but that his genitalia came in contact with hers, and cop- none hatched. ulation ensued. Copulation (n ϭ 11) sometimes con- Eggs were laid singly and afÞxed laterally, usually to tinued for more than 4 h. the lid or paper toweling but occasionally to the Þlter Descriptions of Immature Stages. Egg. (Fig. 3). paper or sides of the cages. The incubation period Length, 1.61 Ϯ 0.01; width at widest point (usually averaged 13.25 d (Table 2). The egg was yellowish near middle), 0.40 Ϯ 0.00. Egg curviform, comprised brown with a smooth chorion, usually with attached of egg proper (length, 1.46 Ϯ 0.01) and operculum longitudinal, white ßanges, and capped by a reticu- (length, 0.14 Ϯ 0.00; diameter, 0.26 Ϯ 0.00). Eggs laid lated operculum (Fig. 3). Red eyespots were visible singly, yellowish brown, glued laterally to substrate, after Ϸ7 d (range ϭ 4Ð9; n ϭ 129). this side convex, opposite side straight. Chorion gen- The Þrst instar emerged through a cephalic slit in erally smooth, usually with Ϸ12 attached, longitudinal, the chorion, pushing aside the operculum. It was white (translucent in alcohol) ßanges fused at base of egg and extending anteriorly beyond base of opercu- lum; operculum with reticulate pattern, central area projecting as distinct cone. Instars. The Þrst instar is described in detail, but only major changes from previous instars are recorded for subsequent instars. Length is measured from tip of anteclypeus to tip of abdomen, width across the pro- jecting posterior lobes of the metaepisterna. Addi- tional measurements are given in Table 3. First Instar (Fig. 4). Length, 3.31 Ϯ 0.03; width, 0.39 Ϯ 0.01. Body slender, elongate. General ground color of head (including beak), thorax, and abdomen white to yellowish white; body shiny, with brown dorsolateral and lateral longitudinal stripes on head and thorax; antennae, meso-, and metathoracic legs brown with white to yellowish white annuli, protho- racic legs with less brown. Head broadly fusiform, narrowed anteriorly and posteriorly, longer than wide. Antenniferous tubercles weakly divergent anteriorly; width of head across api- cal margin of tubercles subequal to synthlipsis. Pair of dorsolateral, longitudinal, discontinuous, brown Fig. 3. Scanning electron micrograph of egg of P. um- stripes extending from antenniferous tubercles to al- brosus (scale bar ϭ 0. 5 mm). most posterior margin of head; stripes interrupted at 196 ANNALS OF THE ENTOMOLOGICAL SOCIETY OF AMERICA Vol. 95, no. 2

Table 3. Measurements (mean ؎ SE mm) of P. umbrosus instars

Nymph 1st instar 2nd instar 3rd instar 4th instar 5th instar Body lengtha 3.31 Ϯ 0.03 5.41 Ϯ 0.05 7.49 Ϯ 0.05 10.51 Ϯ 0.16 15.35 Ϯ 0.32 Head lengtha 0.44 Ϯ 0.00 0.54 Ϯ 0.01 0.64 Ϯ 0.01 0.78 Ϯ 0.01 1.04 Ϯ 0.02 Anterior lobe 0.21 Ϯ 0.00 0.29 Ϯ 0.01 0.35 Ϯ 0.01 0.41 Ϯ 0.01 0.56 Ϯ 0.01 Posterior lobe 0.23 Ϯ 0.00 0.27 Ϯ 0.01 0.29 Ϯ 0.01 0.37 Ϯ 0.01 0.48 Ϯ 0.02 Width across eyes 0.26 Ϯ 0.00 0.30 Ϯ 0.00 0.33 Ϯ 0.00 0.41 Ϯ 0.01 0.53 Ϯ 0.01 Synthlipsis 0.22 Ϯ 0.00 0.25 Ϯ 0.00 0.27 Ϯ 0.00 0.30 Ϯ 0.01 0.39 Ϯ 0.01 Antennal segments 4.55 Ϯ 0.04 6.16 Ϯ 0.08 7.67 Ϯ 0.10 10.29 Ϯ 0.09 13.89 Ϯ 0.17 1st 1.83 Ϯ 0.02 2.57 Ϯ 0.03 3.26 Ϯ 0.04 4.49 Ϯ 0.05 6.15 Ϯ 0.09 2nd 1.23 Ϯ 0.01 1.91 Ϯ 0.03 2.64 Ϯ 0.04 3.77 Ϯ 0.04 5.28 Ϯ 0.08 3rd 0.17 Ϯ 0.00 0.19 Ϯ 0.00 0.21 Ϯ 0.00 0.24 Ϯ 0.00 0.31 Ϯ 0.01 4th 1.32 Ϯ 0.01 1.49 Ϯ 0.02 1.56 Ϯ 0.03 1.79 Ϯ 0.02 2.15 Ϯ 0.02 Beak segments 0.47 Ϯ 0.01 0.60 Ϯ 0.01 0.69 Ϯ 0.01 0.83 Ϯ 0.03 1.22 Ϯ 0.02 1st 0.13 Ϯ 0.00 0.18 Ϯ 0.01 0.21 Ϯ 0.01 0.27 Ϯ 0.01 0.42 Ϯ 0.02 2nd 0.12 Ϯ 0.00 0.16 Ϯ 0.00 0.19 Ϯ 0.00 0.23 Ϯ 0.00 0.35 Ϯ 0.00 3rd 0.22 Ϯ 0.01 0.26 Ϯ 0.00 0.29 Ϯ 0.00 0.33 Ϯ 0.01 0.45 Ϯ 0.01 Notal lengthsb 0.94 Ϯ 0.01 1.48 Ϯ 0.02 2.03 Ϯ 0.01 2.87 Ϯ 0.04 4.30 Ϯ 0.07 Pronotum 0.34 Ϯ 0.00 0.56 Ϯ 0.01 0.80 Ϯ 0.01 1.18 Ϯ 0.02 1.78 Ϯ 0.03 Mesonotum 0.33 Ϯ 0.00 0.52 Ϯ 0.01 0.68 Ϯ 0.01 0.91 Ϯ 0.02 1.32 Ϯ 0.02 Metanotum 0.27 Ϯ 0.00 0.40 Ϯ 0.00 0.55 Ϯ 0.01 0.78 Ϯ 0.01 1.20 Ϯ 0.02 Width across metaepisterna 0.39 Ϯ 0.01 0.42 Ϯ 0.01 0.46 Ϯ 0.01 0.58 Ϯ 0.01 0.77 Ϯ 0.02 Abdominal lengthb 1.94 Ϯ 0.03 3.48 Ϯ 0.04 4.82 Ϯ 0.04 6.81 Ϯ 0.12 10.01 Ϯ 0.23 Leg lengths Procoxa 0.58 Ϯ 0.01 0.79 Ϯ 0.01 0.99 Ϯ 0.01 1.39 Ϯ 0.01 2.06 Ϯ 0.04 Protrochanter 0.19 Ϯ 0.01 0.27 Ϯ 0.01 0.33 Ϯ 0.01 0.37 Ϯ 0.01 0.48 Ϯ 0.01 Profemur 0.88 Ϯ 0.01 1.15 Ϯ 0.02 1.45 Ϯ 0.02 1.97 Ϯ 0.02 2.92 Ϯ 0.05 Protibia 0.42 Ϯ 0.01 0.55 Ϯ 0.01 0.60 Ϯ 0.01 0.81 Ϯ 0.01 1.07 Ϯ 0.02 Protarsus 0.28 Ϯ 0.00 0.33 Ϯ 0.00 0.36 Ϯ 0.00 0.47 Ϯ 0.01 0.66 Ϯ 0.01 Mesocoxa 0.17 Ϯ 0.00 0.21 Ϯ 0.00 0.24 Ϯ 0.01 0.31 Ϯ 0.00 0.48 Ϯ 0.01 Mesotrochanter 0.17 Ϯ 0.01 0.21 Ϯ 0.01 0.24 Ϯ 0.01 0.29 Ϯ 0.00 0.39 Ϯ 0.01 Mesofemur 1.78 Ϯ 0.03 2.41 Ϯ 0.04 3.05 Ϯ 0.03 4.21 Ϯ 0.05 6.01 Ϯ 0.08 Mesotibia 2.53 Ϯ 0.02 3.37 Ϯ 0.05 4.10 Ϯ 0.06 5.61 Ϯ 0.08 7.80 Ϯ 0.09 Mesotarsus 0.29 Ϯ 0.01 0.32 Ϯ 0.00 0.32 Ϯ 0.01 0.38 Ϯ 0.01 0.47 Ϯ 0.02 1st 0.09 Ϯ 0.00 0.11 Ϯ 0.00 0.12 Ϯ 0.00 0.14 Ϯ 0.00 0.19 Ϯ 0.01 2nd 0.20 Ϯ 0.00 0.21 Ϯ 0.00 0.20 Ϯ 0.01 0.23 Ϯ 0.00 0.30 Ϯ 0.01 Metacoxa 0.18 Ϯ 0.00 0.23 Ϯ 0.00 0.27 Ϯ 0.01 0.35 Ϯ 0.01 0.49 Ϯ 0.01 Metatrochanter 0.18 Ϯ 0.01 0.23 Ϯ 0.00 0.25 Ϯ 0.01 0.33 Ϯ 0.01 0.44 Ϯ 0.01 Metafemur 2.10 Ϯ 0.02 2.89 Ϯ 0.05 3.70 Ϯ 0.05 5.22 Ϯ 0.07 7.56 Ϯ 0.10 Metatibia 3.11 Ϯ 0.02 4.12 Ϯ 0.06 5.15 Ϯ 0.07 7.26 Ϯ 0.15 10.31 Ϯ 0.10 Metatarsus 0.29 Ϯ 0.01 0.32 Ϯ 0.00 0.33 Ϯ 0.00 0.38 Ϯ 0.00 0.51 Ϯ 0.01 1st 0.09 Ϯ 0.00 0.11 Ϯ 0.00 0.12 Ϯ 0.00 0.15 Ϯ 0.00 0.19 Ϯ 0.01 2nd 0.20 Ϯ 0.00 0.21 Ϯ 0.00 0.21 Ϯ 0.00 0.24 Ϯ 0.00 0.32 Ϯ 0.01

Data based on 10 individuals per instar. a Meaured from tip of anteclypeus. b Measured at midline. posterior margin of middorsal transverse sulcus that segment 2 with subapical trichobothrium dorsally; continues anterolaterad, ending anterior to eyes; sul- segment 3 shortest; segment 4 acute apically, more cus dividing head into anterior and posterior lobes, heavily setose than preceding segments, length of se- anterior lobe Ϸ0.91 times length of posterior lobe; tae subequal to longer than width of segment; ratio of posterior lobe distinctly elevated above anterior lobe, antennal segment lengths Ϸ10.77:7.24:1.00:7.77. anterior margin occasionally concave medially. Tylus Thorax elongate, narrowest at juncture of pro- and greatly surpassing juga, divided into obscure ante- mesothoraces; ratio of notal lengths to body width clypeus and longer postclypeus; postclypeus with (see deÞnition) Ϸ0.87:0.85:0.69; nota with pair of dor- transverse, brown band anteriorly; juga short, sloping solateral, brown stripes extending from anterior mar- laterad, not reaching distal margins of antenniferous gin of pronotum to posterior margin of metanotum; tubercles. Labrum elongate, digitiform, declivent. dorsomedial white stripe occasionally present, ex- Ventrolateral brown stripe extending each side from tending from anterior margin of mesonotum almost to antenniferous tubercle to almost posterior margin of apex of abdomen. Prothorax subtrapezoidal, narrowed head, stripes interrupted by small, granular, red com- posteriorly; notum weakly overlapping mesonotum pound eyes. Beak three-segmented, brown laterally, posteriorly, posterolateral margin distinctly separated tip Þtting into prosternal, striated groove; segments 1 from prospiracular swelling. Mesothorax subtrapezoi- and 2 subequal, segment 3 longest, narrowing apically. dal, narrowed anteriorly. Metathorax broadly subtrap- Antennae 4-segmented, Þliform, annulated with white ezoidal, narrowed anteriorly. Thoracic spiracles at intersegmental lines; total length Ϸ10.34 times present, directed laterad, peritremes brown; spiracle 1 length of head; segment 1 longest, swollen basally; near posterior margin of proepimeron, spiracle 2 near March 2002 BRADSHAW AND MCPHERSON:LIFE HISTORY OF P. umbrosus 197

surface; spines with tips dark brown; large basal spine with distal two-thirds often brown. Protibiae short, swollen and setose apically; anteroventral longitudinal row of 3Ð5 minute spines apically; setae ventrally en- compassing diagonal, setose, comblike structure; brown basal, subbasal, and apical annuli present. Pro- tarsi 1-segmented, brown apically, with single brown claw. Meso- and metathoracic legs narrow, elongate; coxae globose, often with diagonal brown band; tro- chanters similar in shape and color to protrochanters; femora and tibiae brown with white to yellowish white annuli; tarsi 2-segmented, brown, segments separated by white membranous area; tarsal claws (2) brown, each with medial tooth. Abdomen apparently 9-segmented, generally ap- pearing membranous; intersegmental lines difÞcult to distinguish, that of segments 1 and 2 obscure; segment 1 apparently shorter and segment 2 apparently longer than each of remaining segments. Tergum 1 appar- ently overlapping tergum 2, laterally with brown band reaching anterior margin of spiracle 2; sternum 2 with pair of anterior, submedial brown marks. Sternum 7 with pair of posterior, submedial brown marks. Seg- ment 8 with one lateral and one ventrolateral pair of brown bands. Segment 9 brown ventrally with apical brown annulus to completely brown but if not com- pletely brown, then with pair of lateral brown bands that merge posteriorly with annulus. Dorsal and ven- tral white, discontinuous, longitudinal bands and ir- regular markings occasionally present submedially on dorsal and ventral surfaces. Eight pairs of abdominal spiracles present, peritremes brown; pair 1 dorsolat- eral, pairs 2Ð8 lateral. Second Instar (Fig. 5). Length, 5.41 Ϯ 0.05; width, 0.42 Ϯ 0.01. Body with sparse setigerous pustules on head laterally, thoracic pleura, and abdomen. Head with anterior lobe Ϸ1.07 times length of posterior lobe; posterior lobe less elevated above anterior lobe. Fig. 4. First instar (general dorsal view) of P. umbrosus Antennae Ϸ11.41 times length of head. Ratio of an- (scale bar ϭ 1 mm). tennal segment lengths Ϸ13.53:10.05:1.00:7.84. Thorax more elongate, ratio of notal lengths to body Ϸ posterior margin of mesoepimeron. Pleura with lateral width 1.33:1.24:0.95. Pronotum less ßattened, more and ventrolateral brown stripes extending from ante- noticeably overlapping mesonotum, posterolateral rior margin of propleuron to posterior margin of meta- margin closer to prospiracular swelling; sternum with pleuron; lateral stripe touching dorsolateral stripe at pair of submedial, irregular longitudinal rows of 5Ð12 anterior and posterior margins of propleuron and pos- setigerous, brown pustules. Meso- and metanota with terior margin of mesopleuron, sometimes also touch- wing pads evident, weakly developed. Prothoracic ing near middle of propleuron and anterior margin of legs (Fig. 14) more spinose; femora with antero- and mesopleuron; ventrolateral stripe discontinuous, ex- posteroventral longitudinal rows of 10Ð13 and 14Ð19 tending from anterior margin of propleuron to poste- spines, respectively; tibiae with anteroventral longi- rior margin of metapleuron. Prothoracic legs raptorial, tudinal row of 9Ð13 spines, now extending length of spinose (Figs. 4 and 13). Procoxae elongate with lat- tibia. eral longitudinal brown stripe. Protrochanters el- Abdomen with pair of discontinuous, longitudinal bowed with short brown bands proximally and dis- brown bands dorsolaterally, ventrally, and, often, on tally. Profemora with antero- and posteroventral terga 5 and 6. Medial brown stripe present on terga longitudinal rows of 8Ð11 and 12Ð14 spines, respec- 7Ð8, interrupted at intersegmental line of 7 and 8. tively; anteroventral row interrupted basally, contin- Tergum 9 mostly brown. Dorso- and ventromedial, ued as single spine basad of interruption; posteroven- white, continuous, longitudinal stripes occasionally tral row continuous, terminating basally in large spine, present; dorsomedial stripe extending from posterior basal spine of anteroventral row not located basad of margin of metanotum (occasionally continuous with this large spine; irregular brown markings on posterior dorsomedial stripe of thorax) to, at most, tergum 7; 198 ANNALS OF THE ENTOMOLOGICAL SOCIETY OF AMERICA Vol. 95, no. 2

Figs. 5–12. SecondÐÞfth instars of P. umbrosus (scale bars ϭ 1 mm). General dorsal view (5Ð8). 5. Second instar. 6. Third instar. 7. Fourth instar. 8. Fifth instar. Terminalia, Þfth instar (9Ð12). 9. Male, ventral view. 10. Female, ventral view. 11. Male, lateral view. 12. Female, lateral view. March 2002 BRADSHAW AND MCPHERSON:LIFE HISTORY OF P. umbrosus 199

Figs. 13–17. Prothoracic legs of P. umbrosus (scale bars ϭ 1 mm). 13. First instar. 14. Second instar. 15. Third instar. 16. Fourth instar. 17. Fifth instar. ventromedial stripe extending from posterior margin Abdomen with pair of discontinuous, longitudinal of metasternum to sternum 8. brown bands on terga 4Ð6, frequently on tergum 3. Third Instar (Fig. 6). Length, 7.49 Ϯ 0.05; width, Medial brown stripe on tergum 7 bifurcated anteri- 0.46 Ϯ 0.01. Body with more setigerous pustules. Head orly. Sternum 9 with midtransverse sulcus. with middorsal transverse sulcus more evident later- Fourth Instar (Fig. 7). Length, 10.51 Ϯ 0.16; width, ally; anterior lobe Ϸ1.21 times length of posterior lobe; 0.58 Ϯ 0.01. Body with more setigerous pustules. Head posterior lobe somewhat swollen posterolaterally; with middorsal transverse sulcus well developed; an- juga more evident, more horizontal in position. An- terior lobe Ϸ1.11 times length of posterior lobe; pos- tennae Ϸ11.98 times length of head. Ratio of antennal terior lobe more swollen posterolaterally. Antennae segment lengths Ϸ15.52:12.57:1.00:7.43. Ϸ13.19 times length of head. Ratio of antennal seg- Thorax more elongate, ratio of notal lengths to body ment lengths Ϸ18.71:15.71:1.00:7.45. width Ϸ1.74:1.48:1.20. Pronotum more developed pos- Thorax more elongate; ratio of notal lengths to body terolaterally, margin touching prospiracular swelling; width Ϸ2.04:1.57:1.35. Pronotum more developed pos- sternum with pair of submedial, irregular longitudinal terolaterally, spiracles directed ventrolaterad; ster- rows of 9Ð18 setigerous, brown pustules. Meso- and num with pair of submedial, irregular longitudinal metanotal wing pads more developed. Prothoracic rows of 25Ð35 setigerous, brown pustules. Meso- and legs (Fig. 15) more spinose; femora with antero- and metanotal wing pads more developed. Mesosternum posteroventral longitudinal rows of 12Ð19 and 17Ð25 with few anterior white to yellowish white setigerous spines, respectively; tibiae with anteroventral longi- pustules. Metasternum usually with pair of posterior, tudinal row of 11Ð16 spines. submedial brown marks; posteriorly with few setiger- 200 ANNALS OF THE ENTOMOLOGICAL SOCIETY OF AMERICA Vol. 95, no. 2 ous pustules. Prothoracic legs (Fig. 16) more spinose; and Hilary “Dee” Gates (Integrated Microscopy and Graph- femora with antero- and posteroventral longitudinal ics Expertise) for use of their facilities, technical expertise, rows of 19Ð25 and 26Ð32 spines, respectively; tibiae and assistance in producing electron micrographs; J. R. Bray, with anteroventral longitudinal row of 16Ð19 spines. Jr. (Department of Plant Biology) for the identiÞcation of Abdomen with pair of discontinuous, longitudinal bryophytes and Mike Mibb (Department of Plant Biology) for the identiÞcation of all other ßora; T. R. Anthony (De- brown bands present on terga 2Ð6. Sternum 9 notice- partment of Zoology) for his statistical expertise; and Beth ably constricted posterior to ventral midtransverse Burke (Department of Zoology) for supplying the fruit ßy sulcus. cultures. We also thank D. W. Webb (Illinois Natural History Fifth Instar (Fig. 8). Length, 15.35 Ϯ 0.32; width, Survey, Champaign); E. D. DeWalt (Illinois Natural History 0.77 Ϯ 0.02. Body with more setigerous pustules. Head Survey, Champaign) and Terry L. Harrison (Department of with anteclypeus appearing as small white to yellowish Entomology, University of Illinois, Urbana); and R. J. Snider white band anterior to brown band on postclypeus; (Department of Zoology, Michigan State University, East anterior lobe Ϸ1.17 times length of posterior lobe; Lansing) for their identiÞcations of dipteran, lepidopteran, posterior lobe distinctly swollen. Antennae Ϸ13.36 and collembolan prey, respectively. We also are grateful to A. M. Haggerty (Clemson University, Clemson, SC) for his times length of head. Ratio of antennal segment Ϸ discovery of P. umbrosus at Little Grand Canyon and pre- lengths 19.84:17.03:1.00:6.94. liminary observations on this species, and R. W. Sites (De- Thorax more elongate; ratio of notal lengths to body partment of Entomology, University of Missouri, Columbia) width Ϸ2.31:1.71:1.56. Prosternum with pair of sub- for his input on illustration techniques. Finally, we thank the medial, irregular longitudinal rows of 44Ð58 setiger- U.S. Forest Service for granting permission to collect in the ous, brown pustules. Meso- and metanotal wing pads Shawnee National Forest, and R. L. Johnson (U.S. Forest similar to those of fourth instar. Mesosternum with Service, Vienna Ranger Station) for expediting processing of dense, anterior, white to yellowish white setigerous the permit application form. This article is part of a thesis pustules. Metasternum always with pair of posterior, submitted to Southern Illinois University at Carbondale by J.D.B. in partial fulÞllment of the requirements for the M.S. submedial brown marks; posteriorly with more seti- degree in zoology. gerous pustules. Prothoracic legs (Fig. 17) more spi- nose; femora with antero- and posteroventral longi- tudinal rows of 24Ð30 and 35Ð39 spines, respectively, References Cited usually with small spine of variable size and length just basad of large basal spine of posteroventral series; Blatchley, W. S. 1926. Heteroptera or true bugs of eastern tibiae with anteroventral longitudinal row of 17Ð22 North America with special reference to the faunas of spines. Indiana and Florida. Nature Publ. Co., Indianapolis. 1116 pp. Abdomen with posterior margin of sternum 8 and Froeschner, R. C. 1988. Family Reduviidae Latreille, 1807. basal half of 9 showing developing external genitalia The assassin bugs, pp. 616Ð651. In T. J. Henry and R. C. (Figs. 9Ð12). Sternum 9 more noticeably constricted Froeschner [eds.], Catalog of the Heteroptera, or true posterior to midtransverse sulcus. Terminal segments bugs, of Canada and the continental United States. E. J. further modiÞed, showing distinct sexual dimorphism, Brill, New York. 958 pp. males with posterior margin of tergum 7 slightly ex- Hagerty, A. M., and J. E. McPherson. 1999. Survey of the tended posteriorly (Fig. 11), females with tergum 8 Reduviidae (Heteroptera) of southern Illinois, excluding somewhat extending posteriorly and laterally (Fig. the Phymatinae, with notes on biology. Great Lakes En- 12). tomol. 32: 133Ð160. Hopkins, M. E., and J. A. Simon. 1975. Pennsylvanian sys- Diagnosis. The Þve instars are distinguished readily tem, pp. 163Ð201. In H. B. Willman, E. Atherton, T. C. by characters other than body size and ratios of var- Buschbach, C. Collinson, J. C. Frye, M. E. Hopkins, J. A. ious measurements. The Þrst instar differs from the Lineback, and J. A. Simon, Handbook of Illinois stratig- secondÐÞfth instars by the lack of wing pads; these raphy. Illinois State Geol. Surv. Bull. 95: 1Ð261. later instars show progressive development of wing McPherson, J. E. 1991a. Noteworthy range extensions of pads up to the fourth instar, which is similar to the Þfth. three emesine species (Heteroptera: Reduviidae). Great The thirdÐÞfth instars can be distinguished by pro- Lakes Entomol. 24: 99Ð101. gressive genital development, which Þrst appears in McPherson, J. E. 1991b. Range extensions of three emesine the third instar as a midtransverse sulcus on abdominal species in North America (Heteroptera: Reduviidae). Great Lakes Entomol. 24: 263Ð264. sternum 9 and is followed by noticeable constriction Mohlenbrock, R. H. 1977. Evaluation of Little Grand Can- of the segment posterior to the sulcus in the fourth and yon Jackson County, Illinois for eligibility of Registered distinct sexual dimorphism in the Þfth. Natural Landmark designation. For. Serv. public Þle 2360, Spec. Interest Areas, Little Grand Canyon. United States Forest Service. Murphysboro, IL. 7 pp. Acknowledgments Wygodzinsky, P. W. 1966. A monograph of the Emesinae We thank the following faculty and staff members of (Reduviidae, Hemiptera). Bull. Am. Mus. Nat. Hist. 133: Southern Illinois University at Carbondale: B. M. Burr and 1Ð614. W. G. Dyer (Department of Zoology) for their critical re- views of the manuscript; J. A. Beatty (Department of Zool- Received for publication 26 July 2001; accepted 10 October ogy) for identiÞcation of spiders; J. J. Bozzola, S. J. Schmitt, 2001.