Stenopelmatus sartorianus Saussure

Stenopelmatus sartorianus Saussure Winged Jerusalem Cricket Figs 143–157, Table 1, 2 1859. Stenopelmatus sartorianus. Revue et Magasin de Zoologie 2(11): 211. 1932. Stenopelmatus sumichrasti. Hebard, Trans. Amer. Entomol. Soc. 58(3): 343 1945. Stenopelmatus sartorianus. Strohecker, Ann. Ent. Soc. Amer. 38(2): 207 1988. Stenopelmatopterus sartorianus Gorochov, Zoologicheskii zhurnal 67(4): 521 Distribution. The most widespread species of Stenopelmatus, known from Veracruz state, Mexico, south into Panama. Recognition characters. Stenopelmatus sartorianus is the only species of New World Jerusalem cricket whose adults have fully developed front and hind wings. In all 15 adults examined, hind wings longer (Fig. 144) than tegmina in situ. Three other species, S. chiapas, S. piceiventris, and S. sanfelipe all have micropterous fore wings as adults, and, at least in the case of the first two, in apparently all early instars. There are no concealed, hind wings, in any instar, under the tegmina of these 3 latter taxa. The dense area of short hairs (see, for example, Fig. 134, p. 82), as seen under the wings of S. chiapas, S. piceiventris and S. sanfelipe, is missing in S. sartorianus. Holotype. Adult female (Fig. 145, with fully formed front and hind wings, right side spread). (1) Anostostoma sartorii Sauss ♀ Tuxtla [Gutierrez, state of Chiapas, Mexico], M. H. S. (2) green label Stenopelmatus sartorianus Sauss. (3) green label 71 (4) red label Holotypus Stenopelmatus sartorianus Saussure. Measurement in mm: Body length 35.9, including ovipositor, hind femur length 14, hind femur width 5.1, tegmina 17.8 and almost reaching tip of abdomen. Left hind leg tibia (Fig. 146) with 4 inner and 4 outer spines. Face with apparent furrow although distorted by damage (Fig. 147). Ovipositor upturned (Fig. 153). Deposited MHNG Geneva. New status: transferred back to Stenopelmatus. Drum. Both sexes from Mexico, Veracruz state, Metlac Canyon (S06-39) drummed similarly. Adult male (R06- 12) had one series (Fig. 148) recorded at 20.5°C, in 2:40 of recording time: 26 fast drums over 1.9s (13.7 d/s) followed by 15 slow drums over 5.8s (2.6 d/s). Adult female (R06-13) had one series (Fig. 149), similar to male R06-12, recorded at 23°C, in 4:15 hours: 39 fast drums over 2.8s (13.9 d/s) followed by 14 slow drums over 4.7s (3.0 d/s). Derivation of name. “sartor” is Latin for tailor, patcher, maybe in reference to the holotype’s long wings and being “well dressed?” Habitat. Riparian forest (Fig. 150) along the Rio Metlac (S06-39). Cloud forests at Monteverde, Costa Rica. Behavior. All instars and adults jumped well (see Discussion below). Heard one mid instar female from Monteverde, Costa Rica (S00-21) femoral-abdominal stridulate. During mating trials (see Discussion below), an adult female femoral-abdominal stridulated while the male tremulated. All instars fed well on wax worm larvae. Is this species an obligate predator? At Metlac Canyon, we put out a nighttime 16 kg long oatmeal trail, especially in the area where we collected the 4 late instars during the daytime. While the oatmeal attracted many other Orthoptera, such as Anabropsis, Glaphyrosoma, Anurogryllus, and Gryllus , no JCs were attracted, including no specimens of a second, sympatric, medium-large, jumping black JC that was also in a log during the daytime and that we discuss on p. 117. Variation. Table 1. Adult female Monteverde (S00-21) with 8 left rear leg inner tibial spines (Fig. 152). This same female had the normal 4 inner rear leg tibial spines on her right side. This left rear tibia was almost 3.5 mm shorter than the right side and indicates, we believe, physical damage of some kind during an earlier instar, with subsequent regeneration (discussed in Weissman 2001a) during succeeding molts. Specimens examined. Costa Rica: Puntarenas , Monteverde, 10° 18’ 01” -84° 47’ 47”, 16-vi-1995 (S95-48) ♀ 2, CAS; 26-vi-1996 (S96-71) Ƌ2, CAS; 14-vii-2000 (S00-21) ♀ 1, CAS. Lo. Palma, P Biolley, no date, 1 adult female. Cangrejal de Asseri, 800m, iv-1906, P. Biolley, 1 last instar female. Guatemala: Huehuetenango Paraiso Municipio La Libertad (~ 5 km E. Hojas Blancas) 1650-1850m (forest, under logs) 29-vi-1966. LC Stuart, 3 late (last?) females with minimal lateral carinae on furrows, UMMZ. Las Mercedes, 3000 ft., Champion, Brit. Mus. 1899-235, BMNH. Other Guatemala nymphs in BMNH from Capetillo and El Tumbador. Mexico: Veracruz , Cualcuapan, between Orizaba & Cordoba, 3800’, 11-vii-1938, E.H. Taylor, adult ♀ 3, adult Ƌ1, ANSP; same data, 18- vii-1938, E.H. Taylor, adult Ƌ1, ANSP. Metlac Canyon, 18° 54’ 30.78” -97° 00’ 44”, 3200’, 20-vi-2006 (S06-39), Ƌ1, ♀ 3, all late instars, DBW, DC Lightfoot, CAS (See Ball & Whitehead (1967) for more information on Metlac Canyon). Portrero Viejo, 13-viii-1941, Mrs. Forbes + E.H. Taylor, adult Ƌ1, ANSP; Chiapas, 8.0 m SE Las Casas (San Cristobal), 11-iv-1941, I. Cantrall, adult ♀ 1 (very small: body length 22.9, hind femur length 8.03, hind femur width 2.53. furrow only with distinct lateral carinae- not connected at top), UMMZ. Two additional Mexico records are cited in Gorochov & Cadena-Castañeda (2016). DNA. Nuclear F0073 (S95-48, Monteverde, Costa Rica) and F1796 (S06-39, Metlac Canyon, Mexico) both recovered together (Fig. 10), and well nested, within Stenopelmatus. The same 2 individuals, along with F1769, also from Metlac Canyon, all recovered together (Fig. 9) for mtDNA. To have left this species in Stenopelmatopterus would have resulted in Stenopelmatus being paraphyletic. Karyotype. Unknown. Discussion. Systematics. Saussure (1859) described 3 Mexican taxa, in one publication, that apparently represent the same species: S. sallei on p. 210, S. sumichastri also on p. 210, and S. sartorianus on p. 211. Elsewhere in this paper, we designate the names of S. sallei (p. 76, a juvenile female) and S. sumichrasti (p. 95, a juvenile male) as nomen dubium because both types are subadults without precise locality data, and such action is consistent with our attempts to modernize the prior taxonomy of this genus. While S. sallei has line priority if we chose to retain that ambiguous name, the type of S. sartorianus is an undisputed adult female with a good type locality, so we follow the suggested action of Strohecker (1945) and retain only that name. Junior synonym S. politus was described from an adult female by Walker, in 1869, from Orizaba, which, like Metlac Canyon (S06-39), is also in the Mexican state of Veracruz. Thus, from Metlac Canyon individuals, we have drums in both sexes, DNA, a small series of adults, mating observations, and habitat information which will now form the basis for further taxonomic decisions and the possibility of new species descriptions in this most widely distributed taxon in Stenopelmatus. Past taxonomy in this species “complex” has been complicated by the surprising difficulty of discerning when females (and males in the case of S. sumichrasti) are adult and for choosing females as holotypes. For instance, in 1859, Saussure illustrated a supposed adult female of S. sallei, an error repeated in 1897, by Saussure & Pictet. Hebard (1932) and Strohecker (1945) both recognized that this female of S. sallei was an immature that would have fully developed hind wings as an adult. When Hubbell (1960) examined this same female of S. sallei, he thought it an adult. In 1988 Gorochov erected his new genus Stenopelmatopterus to include 3 winged species: S. sallei, S. sumichrasti, and S. sartorianus. When DBW examined the holotype of S. sallei in 1999, he also thought that she was an adult female, based on her ovipositor (see Fig. 125, p. 78), and placed an appropriate label. Fortunately, after examination of a good series of S. sartorianus, of many different instars in the collections of ANSP, UMMZ, NHMUK, and MHNG Geneva, we can finally clarify this situation. First, one must distinguish between the 3 taxa (S. piceiventris, S. sanfelipe, and S. chiapas) with micropterous fore wings (and no hind wing) versus subadult specimens of S. sartorianus that have pad-like tegmina (Figs. 124 & 125) in earlier instars, but which also have hind wing pads under those pad-like tegmina (confirmed for S. sallei, see p. 76). Second, once S. sartorianus reaches penultimate and last instar, then these hind wing pads are clearly visible (Fig. 154) without the need to lift the tegmina. Flight attempts of winged adults. Winged adult male (S06-39). Observations made at 14 and 44 days after molt to adult, to ensure not teneral. Made under red and incandescent light. Male moved extremely fast, antennae always in motion, perhaps a wasp mimic? He was able to jump almost 45 cm laterally, without using wings. DBW threw him into air several times and although he clearly spread both his tegmina and hind wings, and maybe fluttered them a little, he still fell straight down onto padded surface. But outstretched wings might have slowed the rate of his descent. Blowing onto his face had no effect nor did he raise his wings when provoked. If he was up in a tree (postulated by Weissman & Lightfoot [2007] and confirmed by Gutiérrez-Rodríguez & Riverón [2018]), jumping would be a good escape mechanism. Winged adult female (S06-39): Jumped readily when disturbed or reached edge of desk. Could see her “cock” her rear legs and then spring. On flat surface, jumps exceeded 15 cm. When jumped off edge of desk, she extended wings as “parachutes” but still fell straight down. No flapping seen. Similar behavior if thrown into air. Another time, jumped from 30 cm above surface and landed a good 30 cm away. We conclude that while neither sex can fly, their wings could slow descent if they jump from a high elevation to escape a predator. Wings apparently not used to startle a predator. In any case, there must be a good reason to have such wings when residing deep inside logs during the daytime. We also never saw any fanning during courtship (see below) as might occur should the adults be releasing pheromones. Mating trials. Virgin adults, both individuals having molted to adult shortly after capture, from Metlac Canyon (S06-39), Veracruz, were enclosed together on 19-vii-2006, initially under incandescent light. While the basic mating details are the same as discussed and documented, for Ammopelmatus sp., in Weissman (2001b), Weissman et al. (2008), and filmed in 4K video (https://www.youtube.com/watch?v=mHbwC-AIyTE), there are similarities and differences worth noting because this is the first time we have documented mating in Stenopelmatus. Four minutes after being placed together (Fig. 155), the male approached the female and touched her antennae with his antennae. One minute later, the female tremulated (moved her abdomen up and down without striking the substrate) and moved closer to the male. We then switched from incandescent to red light. At 29 minutes, the female tremulated and performed audible abdominal-femoral stridulation. The male tremulated but no stridulation was heard. At 33 minutes, the male quickly got into a “bite-back” position, biting the female’s left rear leg tibia (Fig. 155), curling his abdomen toward the female’s genitalia while extruding his internal genitalia (Fig. 156), until he could anchor his hooks and quickly pass a spermatophore (Fig. 156). Note absence of any obvious, clear spermatophylax, which is normally easily visible in a mating involving a virgin male Ammopelmatus (see Fig. 1j in Weissman et al. 2008). They stayed adjacent (Fig. 157) for 2 minutes and then separated. Yet within 30 seconds, the male was again “onto” the female, a situation never seen in some 90 matings involving various species of Ammopelmatus. Over the next 10 minutes, this male repeatedly showed interest in the female by approaching her, rolling onto his side, biting her, and trying to physically get her to engage with him. She showed no interest, kicking him off several times. Such a refractory period is typically seen in just-mated female Ammopelmatus. At 11 minutes post coitus, the male ceased his sexual aggressiveness. Around that time, the female curled her abdomen 3 times, in 5 minutes, as if trying to get to the spermatophore. She never succeeded and the sticky spermatophore quickly accumulated a layer of substrate sand (Fig. 157). Four months later, this now once mated male was enclosed with a second virgin adult female from S06-39. While there was no consummation, the male made 2 novel motions not seen in his previous mating or in any previous Stenopelmatinae pairing: (1) a silent, oscillating back and forth motion, along his long body axis, and (2) a rapid “shivering” motion that was faster than (1) and with less total body movement than (1). Whether these unique body movements are widespread in this species is unknown given our small sample size. We also don’t know if such motions are ancient, or recently derived, within this group, although we subsequently saw oscillations, as described in (1) above, in S. perote (p. 69). Weissman et al. (2008) noted how “the [bite-back] positions assumed during copulation are so characteristic and unique [in the Stenopelmatinae] that they may be used, perhaps for the first time, to delimit an entire subfamily of insects.” It will be interesting to see if members of the other two extant Jerusalem cricket subfamilies use such a bite-back mechanism.
Published as part of Weissman, David B., Vandergast, Amy G., Song, Hojun, Shin, Seunggwan, Mckenna, Duane D. & Ueshima, Norihiro, 2021, Generic relationships of New World Jerusalem crickets (Orthoptera: Stenopelmatoidea: Stenopelmatinae), including all known species of Stenopelmatus, pp. 1-122 in Zootaxa 4917 (1) on pages 86-94, DOI: 10.11646/zootaxa.4917.1.1, http://zenodo.org/record/4472033
{"references":["Weissman, D. B. (2001 a) North and Central America Jerusalem crickets (Orthoptera: Stenopelmatidae): taxonomy, distribution, life cycle, ecology, and related biology of the American species. In Field, L. H. (Ed.), The Biology of Wetas, King Crickets and Their Allies. CAB International, New York, pp. 57 - 72. https: // doi. org / 10.1079 / 9780851994086.0057","Gorochov, A. V. & Cadena-Castaneda, O. J. (2016) New and little known Stenopelmatoidea (Orthoptera: Ensifera) from America. Zoosystematica Rossica, 25, 98 - 143. https: // doi. org / 10.31610 / zsr / 2016.25.1.98","Saussure, H. de (1859) Orthoptera nova America. Revue et Magasin de Zoologie, 2 (11), 201 - 212.","Strohecker, H. F. (1945) Notes on and descriptions of Mexican Orthoptera. Annals of the Entomological Society of America, 38, 207 - 215. https: // doi. org / 10.1093 / aesa / 38.2.207","Hebard, M. (1932) New species and records of Mexican Orthoptera. Transactions of the American Entomological Society, 58, 201 - 371.","Hubbell, T. H. (1960) Notes on Orthoptera in European Museums. Unpublished notes deposited at University of Michigan, Museum of Zoology, Ann Arbor, Michigan, 28 pp.","Weissman, D. B. & Lightfoot, D. C. (2007) Techniques for the field capture and captive rearing of North American Jerusalem crickets (Orthoptera: Stenopelmatinae). Sonoran Arthropod Studies Institute 2007 Invertebrates in Captivity Conference, 2007, pp. 22 - 29.","Gutierrez-Rodriguez, J. & Riveron, A. Z. (2018) Los cara de nino (Orthoptera, Stenopelmatidae, Stenopelmatinae): insectos inofensivos pero con muy mala reputacion. Boletin de la Asociacion Mexicana de Sistematica de Artropodos, 2, 5 - 6.","Weissman, D. B. (2001 b) Communication and reproductive behaviour in North American Jerusalem crickets (Stenopelmatus) (Orthoptera: Stenopelmatidae). In: Field, L. H. (Ed.), The Biology of Wetas, King Crickets, and Their Allies. CAB International, New York, pp. 351 - 373. https: // doi. org / 10.1079 / 9780851994086.0351","Weissman, D. B., Judge, K. A., Williams, S. C., Whitman, D. W. & Lee, V. F. (2008) Small-male mating advantage in a species of Jerusalem cricket (Orthoptera: Stenopelmatinae: Stenopelmatus). Journal of Orthoptera Research, 17, 321 - 332. https: // doi. org / 10.1665 / 1082 - 6467 - 17.2.321"]}