Eusermyleformia Bradler, 2009: 101, figs 26c–d, 27d–e, 28a–d.

Diapheromerini, Otte & Brock, 2005: 29 (in part).

Diapheromerini: “Diapheromera group”, Zompro, 2001: 208.

Diapheromerini: “Sermyle group”, Zompro, 2001: 215.

Diapheromerini: “Bacteria group”, Zompro, 2001: 202 (in part).

Description.—Small to fairly large (body length ca 30–150 mm), slender to moderately robust, cylindrical and typically stick-like Diapheromerinae. Both sexes apterous. No ocelli. Gula present. Antennae filiform, longer than complete thorax and consisting of > 30 antennomeres. Pronotum without distinct defensive glands. Mesothorax elongate and > 2×longer than head and pronotum combined. Median segment very short, < ¼ the length of metanotum. No praeopercular organ on sternum VII of females. Stigmata of abdominal tergum VIII shifted towards the median portion of corresponding segment. Epiproct small. In females cerci ± (more or less) straight and mostly shorter than anal segment, rarely lanceolate and somewhat longer than anal segment; round to oval in transverse-section. Gonapophyses VIII considerably longer than gonapophyses IX and longer than gonoplacs. Gonoplacs longer than gonapophyses IX. Gonangulum present or absent. Subgenital plate mostly short, scoop-shaped and merely covering bases of gonapophyses, rarely projecting somewhat over apex on anal segment. In males anal segment ventrally closed by sternum X, no paired ventral thorn-pads at posterior margin. Vomer reduced (a rudimentary vomer is present in only one genus). Epiproct very small. Cerci ± prominently enlarged, longer than anal segment, in-curving and specialized in various ways; mostly round to cylindrical in cross-section with an internobasal protuberance and occasionally with the apex laterally flattened and spatulate, bifid, trifid or deeply furcate; in certain genera asymmetrical cerci occur with the dextral cercus smaller than the sinistral one. Poculum small and rarely somewhat larger than basal portion of sternum IX, often specialised. Basal portion of sternum IX occasionally with one finger-like dextrolateral appendix or two lateral appendices, one on the right and one on the left (phalanges). Legs long and slender, the mesofemora (and metafemora) sometimes ± thickened and with single sub-basal lobes on two outer ventral carinae or a subapical spine on medioventral carina. Profemora compressed and curved basally with the anterodorsal carina ± raised, medioventral carina slightly displaced towards anteroventral carina. Four posterior femora and all tibiae trapezoidal in cross-section. No area apicalis on tibiae. Basitarsi longer than following tarsomeres.

Comments.—Bradler (2009) listed the genera Diapheromera Gray, 1835, Megaphasma Caudell, 1903, Manomera Rehn & Hebard, 1907, Pseudosermyle Caudell, 1903, Litosermyle Hebard, 1919 and Sermyle Stål, 1875 (= Hoplolibethra Caudell, 1904) as belonging to Eusermyleformia. Research in the course of an extensive study on the species groups of Diapheromerini suggested by Zompro (2001) has revealed one further genus exhibiting the characteristics of Eusermyleformia: this is Paracalynda Zompro, 2001. Consequently, Paracalynda is here removed from the “Bacteria-group”, where it was originally placed by Zompro (2001), and transferred to Eusermyleformia.

Within Eusermyleformia Paracalynda is characteristic for having the subgenital plate of females slightly longer than in the other genera, reaching as far as to the apex of the anal segment, and for males having a small, hook-like rudimentary vomer at the base of the abdominal sternum X (Figs 10–11). Both features can certainly be interpreted as plesiomorphic and would place Paracalynda as the adelphotaxon of the remaining Eusermyleformia except Litosermyle Hebard, 1919 [rather than Megaphasma as suggested by Bradler (2009)]. This is also supported by females of Paracalynda having the stigmata of tergum VIII just slightly shifted off the anterior margin (Fig. 8), a character state interpreted as apomorphic for the Eusermyleformia by Bradler (2009). The only other Eusermyleformia, in which males have a small rudimentary vomer, is the Colombian Litosermyle Hebard, 1919. However, the large poculum of males, which is noticeably larger than the basal portion of sternum IX, the conspicuous shape of the subgenital plate of females as well as the basal position of the stigmata of abdominal tergum VIII in both sexes distinguish Litosermyle from all other genera of Eusermyleformia (Conle, Hennemann & Gutiérrez 2011). This would suggest Litosermyle as the adelphotaxon of the entire Eusermyleformia including Paracalynda, but the exact systematic position of Litosermyle deserves further evaluation.

Eusermyleformia now comprises genera of three species-groups of Diapheromerini suggested by Zompro (2001), the “Diapheromera-group”, the “Sermyle-group” and one genus of the “Bacteria-group”. Since Eusermyleformia contains Diapheromera Gray, 1835, the type- genus of Diapheromerini Kirby, 1904, Bradler's rank-free taxon represents what should in fact be regarded as the tribe Diapheromerini (sensu stricto).

Distribution.—Members of the Eusermyleformia are present in the whole of Central America as far south as the northern portions of Colombia and have northwards inhabited almost the complete United States and dispersed as far as north as SE Canada. The center of its radiation with by far the highest diversity of species and genera is Mexico, which harbors four of the seven genera currently contained.

Genera included.—

Type species.—Bacunculus pictus Brunner v. Wattenwyl, 1907: 333, by original designation.

Paracalynda Zompro, 2001: 207, figs. 19–20, 88–89, 130–131.

Bacunculus, Brunner v. Wattenwyl, 1907: 331 (in part — only species no. 3); Shelford, 1909: 351 (in part).

Heteronemia, Brock, 1993: 19; Brock, 1998: 50.

Pseudobacteria, Zompro, 1998: 215.

Description. — Medium sized to large (body length ♂♂ 70.0–86.0 mm, ♀♀ 111.0–124.0 mm) and very slender members of the Eusermyleformia with slender and entirely unarmed legs. Color of female mostly plain green, more rarely straw; often to a variable degree furnished with small brown and grey speckles. Males multicolorous, generally green or brown, the head often with conspicuous markings on frons and the legs annulated. Body surface of both sexes entirely smooth and very slightly glabrous. Head ovoid (♂♂) or roundly rectangular (♀♀) in dorsal aspect, longer than wide and broadest at the eyes; vertex flattened to very gently rounded and entirely smooth. Eyes of moderate size and projecting hemispherically. Antennae very long and filiform, reaching to abdominal segment III in females and longer than body in males. Scapus dorsoventrally compressed (♀♀ in particular), roundly rectangular in dorsal aspect, and 1.3–1.5× longer than wide. Pedicellus subcylindrical to slightly club shaped and a little shorter than scapus. III slightly longer than pedicellus. Pronotum a little shorter and narrower than head, considerably longer than wide and rectangular. Mesothorax very elongate and >3× longer than head and pronotum combined; in males narrower than prothorax. Metathorax about 2/3 length of mesothorax. Meso- and metasternum simple. Abdomen excluding median segment slightly longer (♀♀) or about equal in length (♂♂) to combined length of head and complete thorax. Median segment very short, scarcely longer than wide in males and slightly transverse in females; less than ¼ the length of metanotum. Abdominal segment II more than 2× longer than median segment. Segments II–VI at least 2.5× longer than wide (relatively more slender and elongate in males) and roughly parallel sided; in males roughly of equal width and length, in females very scarcely increasing in length with VI the longest segment. VII in both sexes shorter than VI. Abdominal sternites II–VII smooth, no praeopercular organ on VII in females. Stigmata of tergum VIII slightly (♀♀) to noticeably (♂♂) displaced and shifted towards the middle of segment (Figs 8–9). Tergites VIII– X in females considerably shorter and just slightly narrower than previous, roughly of uniform width; VIII roughly 1.3× the length of IX or X. Anal segment about as long as IX, longitudinally carinate and narrowed towards the apex, the posterior margin with a shallow median indention. Epiproct very small, rounded and indistinctly projecting over posterior margin of anal segment. Cerci longer than anal segment, slender, lanceolate and gradually narrowed towards a pointed apex; very faintly carinate dorsally (Fig. 12). Gonapophyses VIII slender, elongate and gently up-curving but not reaching to apex of anal segment. Gonoplacs thickened, finger-like and considerably longer than gonapophyses IX, but shorter than gonapophyses VIII (Fig. 12). Subgenital plate slender, longitudinally carinate with the apex strongly narrowed, pointed and ± reaching apex of anal segment (Figs 8, 12). In males tergites VIII–X much shorter than previous and of unequal width. VIII constricted basally but strongly swollen and broadened in posterior 2/3; wider than II–VII. IX shorter and narrower than VIII or X. Anal segment broader than IX and widened toward posterior, the posterior margin with a wide but shallow median indentation; ventrally closed by sternum X; a very small, tooth or hook-like rudimentary vomer present (Figs 10, 11). Epiproct very small and completely hidden under anal segment. Poculum smaller than basal portion of sternum IX, cup-shaped and not reaching to posterior margin of tergum IX. Cerci large, longer than anal segment, distinctly incurving with the basal portion broadened and interobasally bearing a spiniform protuberance (Figs 5, 7). Legs all long, slender and entirely unarmed, the meso- and metafemora ± decidedly thickened. Profemora slightly longer than mesothorax, mesofemora longer than metathorax, and metatibiae almost reaching (♀♀) or prominently projecting over apex of abdomen (♂♂); tibiae slightly longer than corresponding femora. Profemora with anterodorsal carina noticeably raised, the medioventral carina distinct and noticeably displaced towards the anteroventral carina (females in particular). Meso- and metafemora and all tibiae trapezoidal in cross-section with the medioventral carina midways. Two outer ventral carinae of mid and hind legs in females very slightly expanded. Basitarsi slender and roughly equal (♀♀) or longer (♂♂) than remaining tarsomeres combined.

Eggs.— (Figs 13–14) Moderately sized (capsule length < 4.0 mm), capsule ovoid, laterally compressed, distinctly oval in cross-section and slightly bullet-like; >1.8× longer than wide. Capsule surface entirely smooth and strongly shiny. Micropylar plate very slightly displaced towards anterior and roughly ½ the length of capsule and 3× longer than wide; the posterior portion gently widened and rounded. Interior surface smooth and outer margin roundly swollen. Micropylar cup represented by a rounded median swelling at polar end of plate. Median line distinct, almost reaching to polar-area. Operculum oval with the anterior portion gently down-curving; opercular angle roughly 10°. Capitulum an irregular, dorsally flattened swelling deeply pitted throughout, which covers most of the surface of the operculum. Color of capsule pale amber or straw with lateral surfaces irregularly furnished with brown speckles and markings. The micropylar plate surrounded by a broad whitish or pale cream colored area. Micropylar plate very pale cream to ivory. Capitulum dull orange.

Differentiation.—Within Eusermyleformia, females of Paracalynda are characterized by an elongate and apically narrowed subgenital plate, which ± reaches to the apex of the abdomen, and by having the stigmata of abdominal tergum VIII just slightly displaced off the anterolateral angles of the segment (Figs 8, 9). Males are characterized by the presence of a small, rudimentary vomer. The latter is of particular interest and, since represented by a small tooth or hook-like protuberance at the base of sternum X (Figs 10, 11), is here interpreted as the remains of a vomer. A similar structure, which may be regarded a rudimentary vomer, is seen amongst the Eusermyleformia only in Litosermyle Hebard, 1919 (Conle, Hennemann & Guttiérrez, 2011) and certain species of Pseudosermyle Caudell, 1903 (Conle, Hennemann & Fontana 2007). The small poculum of males, which is smaller than the basal portion of abdominal sternum IX (Figs 4, 6, 9) is shared with Pseudosermyle, but while it is typically cup-shaped in Paracalynda, conspicuous specializations of the poculum are seen in the latter genus.

In general Paracalynda resembles Manomera Rehn & Hebard, 1907 from the eastern United States; but it differs by the noticeably less elongate and oval head (> 2× longer than wide in Manomera) by the entirely unarmed legs of both sexes, by the much longer subgenital plate of females, which roughly reaches to the apex of the abdomen (Fig. 12), as well as by the presence of a rudimentary vomer and slender mesofemora of males; the last lacking the conspicuous sub-apical spine on the medioventral carina seen in Manomera. The eggs differ from those of Manomera mainly in the pale, smooth and strongly shiny capsule surface (Figs 13, 14).

Males also resemble some of the larger and more slender representatives of Pseudosermyle, but frequently differ by the typically incurving cerci (specialized in Pseudosermyle), which bear a interobasal spine, and larger unspecialized poculum.

Comments.—Although this genus is very close to Pseudosermyle and the nearctic Manomera Rehn & Hebard, 1907 and, based on striking genital features, apparently belongs in the Eusermyleformia, Zompro (2001) placed this genus in the “Bacteria-group”; then in contrast to this placement, he postulated close relation to the Central American Calynda Stål, 1875, a member of the “Phanocles-group”. The genital morphology of Paracalynda however differs fundamentally from that of both those species-groups. Males lack a well-developed vomer and paired ventral thorn pads on the posterior margin of the anal segment, have the anal segment ventrally closed by sternum X and the long incurving cerci have an interobasal spine. Both sexes have the stigmata of tergum VIII displaced and shifted towards the middle of the segment and females lack a praeopercular organ. Consequently, the generic name Paracalynda, which errorneously postulates relation to Calynda, is an unfortunate choice.

The Brazilian Pseudobacteria longipes Toledo Piza, 1938 was errorneously placed in Paracalynda by Zompro & Domenico (2005), although this species has a much longer median segment, dorsally raised basitarsi, a well-developed vomer, paired ventral thornpads on the posterior margin of the anal segment, short and simple cerci and the stigmata of tergum VIII positioned in the anterolateral angles of the corresponding segment. Hence, this species was removed from Paracalynda and provisionally transferred to Bacteria Latreille et al., 1825 by Otte & Brock (2005). The exact generic position of this species will be clarified by Hennemann & Conle (in press). Research on the type-series of P. picta (Brunner v. Wattenwyl, 1907), the only species that has previously been included in the genus, has revealed inaccuracies concerning its recognition as presented by Zompro (2001).

Distribution.—Northern Central America (Belize, Honduras and Guatemala). Although not yet recorded from Mexico the genus is also likely to be found in the southeastern provinces, Chiapas, Tabasco and Campeche. Principally Paracalynda is a neotropical taxon which is restricted to the Mesoamerican Dominion and three biogeographical provinces termed Mexican Gulf, Chiapas and Eastern Central America (Morrone 2006).

Species included.—

Bacunculus pictus Brunner v. Wattenwyl, 1907: 333. LT [by present designation], ♂: Coll. Br. v. W., Vera Paz, Godman; det. Br. v. W., Bacunculus pictus; 23.856 [NHMW, No. 684]. (Not: PLT's in NHMW and BMNH. These are P. utilaensis (Zompro, 1998), see below); Shelford, 1909: 351.

Heteronemia picta, Brock, 1993: 19; Brock, 1998: 50.

Pseudobacteria picta, Zompro, 1998: 215.

Paracalynda picta, Zompro, 2001: 207.

Paracalynda picta picta, Otte & Brock, 2005: 245.

Diagnosis.—Males (the only sex known) differ from those of P. utilaensis (Zompro, 1998) by: the larger size; more slender body; green general coloration (Fig. 3); considerably more elongate and slender head; relatively more slender and elongate abdominal tergites VIII–X (Fig. 4); somewhat smaller poculum; more prominent basal protuberance of sternum X and relatively longer and more slender cerci, the cerci having the interobasal spine slightly shorter but broader than in utilaensis (Fig. 5).

Description.— Male (Fig. 3): Fairly large (body length 86.0 mm) and slender for the genus. General color dull yellowish to brownish green, meso- and metanotum in the HT with a dark greyish wash dorsally. Head with a bold, oval ivory-colored spot between the eyes and in front with a black transverse line. Eyes and cerci reddish mid brown. Meso- and metafemora and all tibiae with three faint yellow annulations; all femora with a brown wash apically. Antennae dark brown with the ventral surface black, scapus and pedicellus greenish middle brown.

Head: Elongate oval, about 2× longer than wide, vertex flattened. Eyes prominent and contained almost 3× in length of cheeks. Scapus roughly 2× longer than wide, pedicellus only ½ the length of scapus. Segment III considerably longer than pedicellus.

Thorax: pronotum slightly shorter but as broad as head, rectangular. Transverse median sulcus faint, gently curved and very slightly displaced towards the posterior. Mesothorax 4.3× longer than head and pronotum combined. Metanotum a little more than half as long as mesonotum.

Abdomen: median segment about 1.7× longer than wide and gently widened towards the posterior; <1/6 the length of metanotum. Segment II about 3× longer than median segment. II–VI about equal in length and width, all gently constricted medially; on average 5.3× longer than wide. VII noticeably shorter than previous. VIII hardly more than half the length of VII, constricted in the basal 1/4, the posterior portion widened, roughly parallel-sided and somewhat broader than II–VII (Fig. 5). Stigmata of tergum VIII small and considerably displaced, being positioned some 1/3 off the anterior angle (Fig. 4). IX shorter and slightly narrower than VII, rectangular. Anal segment scarcely longer than IX and gently widened in posterior portion, the posterior margin with a shallow median indentation. Rudimentary vomer is an acute back-curving, hook-like protuberance (Fig. 4). Cerci about 1.3× longer than anal segment, slender, incurving and with the apex slightly club-like; intero-basal spine short but acute (Fig. 5). Poculum strongly convex, cup-shaped and reaching roughly halfway along tergum IX.

Legs: all very long and slender. Profemora almost equal in length to head, pro- and mesonotum combined, mesofemora reaching 2/3 the way along abdominal segment II and metafemora reaching to posterior margin of V. Basitarsi very elongate and almost 1. 3× longer than remaining tarsomeres combined.

Comments.—The here designated and redescribed LT of Bacunculus pictus Brunner v. Wattenwyl, 1907 in NHMW is the only specimen of the type-series that fully matches the original description and key features used by Brunner v. Wattenwyl (1907). The PLT's in NHMW and BMNH are not conspecific and both represent P. utilaensis (Zompro, 1998 n. stat.). Hence, P. picta is so far known only from a unique type specimen.

Distribution.—Only known from the type-locality Vera Paz in northern Central Guatemala.

Pseudobacteria picta utilaensis Zompro, 1998: 215, figs. 6–8. HT, ♂: Honduras, Isl. Utila, III. 1997, T. Kujawski leg. [coll. OZ]; PT, 1 ♂, 1 ♀, 1 ♀ (nymph n5): Honduras, Isl. Utila, III. 1997, T. Kujawski leg. [coll. OZ]; PT, 2 ♂♂, 1 ♀: Captive reared by O. Zompro, F1-generation [coll. OZ]. n. stat.

Paracalynda picta utilaensis, Zompro, 2001: 207, figs. 19–20, 88–89, 130–131; Otte & Brock, 2005: 245.

Bacunculus pictus Brunner v. Wattenwyl, 1907: 333 (in part — only PLT's). PLT, ♂: Coll. Br. v. W., Guatemala, Mus. Stuttgart; det. Br. v. W., Bacunculus pictus; 23.376 [NHMW, No. 684; specimen retained from SMNS]; PLT, ♂: Syntype; Senahu, Vera Paz, Champion; 32.; Godman-Salvin Coll. 1908.-168.; B.C.A. Orth. II, Bacunculus pictus Redt.; Bacunculus pictus Brunner, 1907, det. John Huxley, 1964, BMNH(E) #844537 [BMNH]. (Not: PLT, ♂: Syntype; Atoyac, Vera Cruz, Schumann; 29.; Godman-Salvin Coll. 1908.-168.; B.C.A. Orth. II, Bacunculus pictus Redt.; Bacunculus pictus Brunner, 1907, det. John Huxley, 1964, BMNH(E) #844536 [BMNH]. à This is notcongeneric and belongs in the genus Pseudosermyle Caudell, 1903).

Further material.— (16 ♂♂, 10 ♀♀, eggs): BELIZE: 1 ♀, 2 ♂♂, eggs: ex Zucht: F. Hennemann, 2007, Herkunft: Belize, Green Hills, Cayo, 2005, leg. J. Meerman [coll. FH, No's 0613-1 to 3 & E]; 3 ♀♀: ex Zucht: F. Hennemann, 2008/09, Herkunft: Belize, Green Hills, Cayo, 2005, leg. J. Meerman [coll. FH, No's 0613-4 to 6]; 6 ♀♀, 14 ♂♂: ex Zucht B. Kneubühler 2006, F1 gen., Zuchtstamm ex Belize, Green Hills, Cayo, leg. J. Meerman, 2005 [coll. OC].

Diagnosis.— Males differ from those of P. picta (Brunner v. Wattenwyl, 1907) by: the smaller size; less slender body; brown general coloration (Fig. 2); shorter and more ovate head; relatively shorter and broader abdominal tergites VIII–X (Fig. 5); somewhat larger poculum; less prominent and acute rudimentary vomer (Figs 10–11) as well as by the relatively shorter and more robust cerci, which have the interobasal spine longer and narrower than in picta (Fig. 6).

Comments.—Zompro (1998) originally described Pseudobacteria picta utilaensis based on material originating from Utila Island (Honduras) some kilometres off the south coast of Belize and regarded this as a subspecies of P. picta (Brunner v. Wattenwyl, 1907), a species described from Guatemala. Subsequently, Zompro (2001) placed both the nominate form and subspecies in the newly described genus Paracalynda. However, Zompro was apparently not aware that P. picta was described from as many as four syntypes, two each in NHMW and BMNH, since the author only quoted a holotype in NHMW and provided the very unprecise data “Coll. Br. v. W., Guatemala, Godman” (Zompro 2001). Examination has proven these four syntypes to represent as many as three distinct species, one of which is not even congeneric and belongs in Pseudosermyle Caudell, 1903. Only the male from Vera Paz in NHMW fully matches with the original description and is here designated as the LT of P. picta (see above). The remaining two PLTs, one each in NHMW and BMNH, are conspecific and represent P. picta utilaensis. Since these two specimens clearly represent a separate species, Zompro's subspecies is here raised to species level (n. stat.).

One adult pair of P. utilaensis was collected by Jan Meerman in Central Belize near Green Hills in the Cayo District during 2005. The collector stated Piper amalgo (Piperaceae) to be the preferred host plant and Paulina sp. (Sapindaceae) to have been accepted as an alternative food-plant in captivity in Belize (Jan Meerman personal communication). Eggs laid by this couple were sent to Bruno Kneubühler (Switzerland), who first reared this species in captivity in Europe, mostly using ferns as an alternative food plant. Eggs of the F1-generation were kindly passed on to the authors, who were able to culture from these on Prunus laureocerasus (Rosaceae) and ferns in fairly dry and well-ventilated conditions. Unfortunately, the F3-generation has become parthenogenetic, but further breeding has shown this species to be capable of parthenogenetic reproduction. Some variation is seen in the coloration of females which range from pale over mid-green to straw, have the body, to a variable degree, furnished with small pale brown to grey speckles, and may sometimes have the bases of all femora mid to dark brown. Males show slight variation concerning the coloration, with specimens from Utila Island having the pale bands of the legs more distinct than from other localities. Zompro (1998) cited a body length of 83.2 mm for the male HT, which is slightly larger than the captive reared specimens at hand.

Distribution.—Utila Island (politically a part of Honduras), Belize (Cayo District: Green Hills) and Guatemala (Vera Paz: Senahu).