Research Article Journal of Orthoptera Research 2023, 32(2): 171-176 A new record of a winged stick insect (Phasmatodea) from Mexico, with a checklist and key to the species of the family Pseudophasmatidae from continental North America MANUEL DE LUNA!, ROBERTO GARCIA-BARRIOS?, GERARDO CUELLAR-RODRIGUEZ!, ULISES LOPEZ-Mora? 1 Facultad de Ciencias Forestales, Universidad Autonoma de Nuevo Leon, San Nicolas de los Garza, Mexico. 2 Facultad de Ciencias Biologicas, Universidad Autonoma de Nuevo Leon, San Nicolas de los Garza, Mexico. 3 Facultad de Ciencias, Universidad Nacional Aut6noma de México, Ciudad de México, Mexico. Corresponding author: Gerardo Cuéllar-Rodriguez (luis.cuellarrd@uanl.edu.mx) Academic editor: Kevin Judge | Received 30 November 2022 | Accepted 31 January 2023 | Published 21 September 2023 https://zoobank. org/7AE85AA9-E81A-4E7B-96CC-C4767D9D7582 Citation: de Luna M, Garcia-Barrios R, Cuéllar-Rodriguez G, L6pez-Mora U (2023) A new record of a winged stick insect (Phasmatodea) from Mexico, with a checklist and key to the species of the family Pseudophasmatidae from continental North America. Journal of Orthoptera Research 32(2): 171-176. https://doi.org/10.3897/jor.32.98203 Abstract The winged stick insect Metriophasma iphicles (Redtenbacher, 1906) (Phasmatodea: Pseudophasmatidae) is recorded for the first time from Mexico (state of Veracruz), making this the northernmost record of both the species and genus. A checklist of species in the family Pseudophasma- tidae from Mexico and the USA is presented, and a key to the species listed is proposed. With the current record, the number of continental North American species of Phasmatodea increases to 108, and the number of genera in the region increases to 23. Keywords Metriophasma iphicles, neotropics, phasmid, Phasmida, Veracruz Introduction The order Phasmatodea (stick and leaf insects) is currently comprised of more than 3500 species worldwide (Brock et al. 2022). In continental North America (Canada, USA, and Mexico), this order is represented by 107 species, grouped into 22 genera and 7 (potentially distantly related) families (L6pez-Mora and Llorentes-Bousquets 2018, 2023, de Luna, in press). Among the North American phasmid fauna, there are records of only 5 winged species: 1 macropterous species of the genus Prisopus Peltier de Saint Fargeau & Audinet-Serville, 1827 (Prisopodidae: Prisopodi- nae: Prisopodini); 1 brachypterous species of the genus Haplopus Burmeister, 1838 (Phasmatidae: Cladomorphinae: Haplopodini); 2 brachypterous species of the genus Hypocyrtus Redtenbacher, 1908 (Phasmatidae: Cladomorphinae: Hesperophasmatini); and 1 macropterous species of the genus Agrostia Redtenbacher, 1906 (Pseudophasmatidae: Stratocleinae: Stratocleini). These winged species are mostly found in neotropical Mexico (Agrostia, Hypocyr- tus, and Prisopus), but one is found in the southernmost state of the United States, Florida (Haplopus) (Arment 2006, L6pez-Mora and Llorente-Bousquets 2018). The family Pseudophasmatidae is of the “Areolatae” group, meaning that its species possess an area apicalis; this is a sunken and usually triangular-shaped areola found in the ventral apex of the middle and hind tibiae (Bradley and Galil 1977, Lopez-Mora and Llorente-Bousquets 2018). This character distinguishes the members of this family from most of the taxa of the region, except the members of the family Timematidae from which they differ most prominently in the number of tarsal segments: 5 in Pseudophasmatidae (and all other families) and 3 in Timematidae. Another exception is the members of the family Prisopodidae, from which they differ in the aspect of the last abdominal segments being laterally expanded into lobes in Prisopodidae (L6pez-Mora and Llorente-Bousquets 2018). This family is currently divided into 3 subfamilies and 7 tribes: Pseudophasmatinae with 3 tribes, Anisomorphini, Paraprisopodini, and Pseudophasmatini; Stratocleinae with 1 tribe, Stratocleini; and Xerosomatinae with 3 tribes, Prexaspini, Setosini, and Xerosomatini (Brock et al. 2022). Pseudophasmatidae is represented in continental North America by 6 species: the macropterous Agrostia rugicollis (Gray, 1835); 3 apterous species of the genus Anisomorpha Gray, 1835 (Pseudophasmatinae: Anisomorphini), Anisomorpha buprestoides (Houttuyn, 1813), Anisomorpha ferruginea (Palisot de Beauvois, 1805), and Anisomorpha paromalus Westwood, 1859; and 2 apterous species of the genus Autolyca Stal, 1875 (Pseudophasmatinae: Anisomorphini), Autolyca elena Gorochov & Berezin, 2008 and Autolyca pallidicornis Stal, 1875. Agrostia rugicollis, Anisomorpha paromalus, and both species of Autolyca are found in neotropical Mexico; the remaining 2 species of Anisomorpha are found in southeastern USA (Arment 2006, Lopez- Mora and Llorente-Bousquets 2018, de Luna in press). Until now, the genus Agrostia was the only recorded winged genus of this family in the region (Arment 2006, L6pez-Mora and Llorente-Bousquets 2018). The genus Metriophasma Uvarov, 1940 (Pseudophasmatidae: Xerosomatinae: Prexaspini) contains 11 macropterous species Copyright Manuel de Luna et al. This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited. JOURNAL OF ORTHOPTERA RESEARCH 2023, 32(2) 172 that are distributed in the neotropical region. They are divided further into 2 subgenera: Acanthometriotes Hebard, 1924, which comprises 3 stocky species native of South America; and Metrio- phasma Uvarov, 1940, which comprises 8 elongated species na- tive mostly to South America. At least 2 species, Metriophasma dio- cles (Westwood, 1859) and Metriophasma iphicles (Redtenbacher, 1906), have been recorded in Central America (Brock et al. 2022). In the present publication, Metriophasma iphicles is recorded for the first time in Mexico, being found in the state of Veracruz; this is currently the northernmost distribution record of any species of the genus. A checklist of the species of the family Pseudophasma- tidae from continental North America is presented, and a key to all listed species is proposed. Methods During a visit (July 2022) to the Estacion de Biologia Tropical “Los Tuxtlas”, in the municipality of San Andres Tuxtla, state of Veracruz, Mexico, 8 specimens (744, 12) of a macropterous species of stick insect were collected at night (Fig. 1A). They were found perched on branches and vines, but there was no evidence indicating that the insects were feeding on these plants. The specimens were preserved individually in 70% ethanol and are kept at the Entomology Lab of the Facultad de Ciencias Forestales (FCF) of the Universidad Auton- oma de Nuevo Leon (UANL) under the vouchers PHASM054-061, with 1 specimen being dry-mounted (PHASM056 - ¢ [Fig. 1B]) to obtain a better view of the pattern of the hindwings (Fig. 1B). Addi- tionally, 3 other specimens from the same locality were examined; these are deposited in the Entomological Collection of the Estacién de Biologia Tropical “Los Tuxtlas” (EBTX45-47). M. DE LUNA, R. GARCIA-BARRIOS, G. CUELLAR-RODRIGUEZ AND U. LOPEZ-MORA The keys of L6pez-Mora and Llorente-Bousquets (2018) were employed in an attempt to identify the genus, the specimens key- ing to Perliodes (now a synonym of Agrostia). However, it was not- ed that the hindwings reached tergite IX (Figs 1B, 2A-C, 3A, C), while in Agrostia, it is known that the hindwings do not reach ter- gite VII (Fig. 4A) (Aquino-Heleodoro et al. 2017). The presence of carinae in the middle and the hind femora was also noted, which are absent in all Stratocleinae including Agrostia (Zom- pro 2005). When the keys of Redtenbacher (1906) and Shelford (1909) were used instead, the specimens keyed to Metriotes (now a synonym of Metriophasma Uvarov, 1940), finding the same re- sults when employing the key of Zompro (2005). After the genus was established, the keys to species present in the works of Red- tenbacher (1906) and Shelford (1909) were employed, with the specimens keying to Metriophasma iphicles. Finally, the specimens were compared to photographs of the type material of Metriophas- ma iphicles from the Phasmida Species File website (Fig. 3A-C) (Brock et al. 2022), corroborating the identity of the collected material (Figs 1A, B, 2A-F). Results and discussion Checklist of species of PSeudophasmatidae from continen- tal North America The checklist includes records to state level, including those made or compiled by Redtenbacher (1906), Shelford (1909), Marifio and Marquez (1983), Conle and Hennemann (2002), Ar- ment (2006), Gorochov and Berezin (2008), and L6pez-Mora and Llorente-Bousquets (2018). Fig. 1. Metriophasma iphicles, males. A. Live specimen found at night; wings spread; photo by Roberto Garcia-Barrios; B. Dry-mounted specimen (PHASMO056) with one tegmina and hindwing extended; photo by Manuel de Luna. JOURNAL OF ORTHOPTERA RESEARCH 2023, 32(2) M. DE LUNA, R. GARCIA-BARRIOS, G. CUELLAR-RODRIGUEZ AND U. LOPEZ-MORA Family PPEUDOPHASMATIDAE Rehn, 1904 Subfamily Pseudophasmatinae Rehn, 1904 Tribe Anisomorphini Redtenbacher, 1906 Genus Anisomorpha Gray, 1835 1. Anisomorpha buprestoides (Houttuyn, 1813) USA (Alabama, Florida, Georgia, Louisiana, Mississippi, North Carolina, South Carolina, Texas). 2. Anisomorpha ferruginea (Palisot de Beauvois, 1805) USA (Del- aware, Florida, Georgia, Illinois, Indiana, Kentucky, Louisiana, Mississippi, North Carolina, Nebraska [dubious record accord- ing to Arment 2006], Oklahoma, Pennsylvania, South Carolina, Texas, and Virginia). 3. Anisomorpha paromalus Westwood, 1859 MEXICO (Yucatan). Genus Autolyca Stal, 1875 4. Autolyca elena Gorochov & Berezin, 2008 MEXICO (Chiapas). 5. Autolyca pallidicornis Stal, 1875 MEXICO (Chiapas). Subfamily Stratocleinae Gtinther, 1953 Tribe Stratocleini Gtinther, 1953 Genus Agrostia Redtenbacher, 1906 6. Agrostia rugicollis (Gray, 1835) MEXICO (Colima). 173 Subfamily Xerosomatinae Bradley & Galil, 1977 Tribe Prexaspini Zompro, 2004 Genus Metriophasma Uvarov, 1940 7. Metriophasma iphicles (Redtenbacher, 1906) MEXICO (Veracruz). New record. Material examined.—MEXICO ¢ 1 9; Estaci6n de Biologia Tropical “Los Tuxtlas”, municipality of San Andres Tuxtla, state of Veracruz; 18.5848°N, -95.0741 °W, 147 maz.s.l.; 25 July 2022; on branches and vines; Roberto Garcia-Barrios and Manuel de Luna leg.; wet speci- men (70% ethanol); collected under permit SGPA/DGVS/04352/22; voucher PHASM054 (FCF-UANL) ¢ 1 3; same data; voucher PHASM055 (FCF-UANL) ¢ 1 3; same data; dry-mounted specimen (Figs 2B, 3A-C); voucher PHASM056 (FCF-UANL) ¢ 1 3; same data; voucher PHASM057 (FCF-UANL) ¢ 1 4; same data; voucher PHASMO058 (FCF-UANL) ¢ 1 3; same data; voucher PHASM059 (FCF-UANL) ¢ 1 3; same data; voucher PHASMOG60 (FCF-UANL) ¢ 1 4; same data; voucher PHASMO61 (FCF-UANL) ¢ 19; same lo- cality; 18.5831°N, -95.0741°W, 154 m a.s.l.; 24 September 2017; on Araceae; Ulises Lopez Mora and Luis Rai Ruiz-Sanchez leg.; dry mounted; collected under permit SGPA/DGVS/03316/17; voucher EBTX45 (UNAM) ¢ 19; same locality; 18.5847°N, -95.0735°W, 125 m as.l.; 25 September 2017; on Araceae; Ulises L6pez Mora and Luis Rai Ruiz-Sanchez leg.; dry mounted; collected under per- mit SGPA/DGVS/03316/17; voucher EBTX46 (UNAM) ¢ 3; same locality; 18.5862°N, -95.0768°W, 170 ma.s.l.; 18 August 2018; on Araceae; Ulises L6pez Mora leg.; dry mounted; collected under per- mit SGPA/DGVS/002646/18; voucher EBTX47 (UNAM). DY Fig. 2. Metriophasma iphicles, abdominal segments VIII, IX, and X. A. Dorsal aspect, male; B. Lateral aspect, male; C. Ventral aspect, male; D. Dorsal aspect, female; E. Lateral aspect, female; FE. Ventral aspect, female. Photos by Manuel de Luna. JOURNAL OF ORTHOPTERA RESEARCH 2023, 32(2) 174 a ™ ’ - a ; r M. DE LUNA, R. GARCIA-BARRIOS, G. CUELLAR-RODRIGUEZ AND U. LOPEZ-MORA A 3 2 —_ —, —— ——$$—$—$__——_ oe oo EE — a _———— — 1 =! a Fig. 3. Metriophasma iphicles; photographs by Dr. Paul D. Brock, copyright Natural History Museum of London. A. Dorsal aspect, male paralectotype; B. Lateral aspect, male paralectotype; C. Dorsal aspect, female lectotype. New diagnosis.—Metriophasma has an area apicalis in the middle and hind tibiae, unlike all Diapheromeridae, Parabacillus Caudell, 1903, and Phasmatidae. It possesses 5-segmented tarsi, unlike the Timematidae. Metriophasma differs from all the North American genera, except 4 others, in having wings: the wings of Metriophasma are well-developed, unlike in the brachypterous species of Haplo- pus and Hypocyrtus (Phasmatidae). The abdomen of Metriophasma is not strongly lobed distally, as seen in macropterous species of Prisopus (Prisopodidae). Finally, Metriophasma has carinae on the ventral side of the middle and hind femora; these are lacking in all Stratocleinae, including Agrostia (Pseudophasmatidae) (Bradley and Galil 1977, Zompro 2005). Metriophasma iphicles differs from the 3 species included in the subgenus Acanthometriotes for its elongated body and in having mesonotal carinae (Hebard 1924); from Metriophasma armatum (Redtenbacher, 1906), Metriophasma baculus (De Geer, 1773), and Metriophasma stollii (Gray, 1835) in having tubercles instead of spines in the dorsal aspect of the mesonotum (Redtenbacher JOURNAL OF ORTHOPTERA RESEARCH 2023, 32(2) M. DE LUNA, R. GARCIA-BARRIOS, G. CUELLAR-RODRIGUEZ AND U. LOPEZ-MORA 175 BMNH(E) 4845248 Fig. 4. Other North American representatives of the family Pseudophasmatidae; photographs by Dr. Paul D. Brock. A. Agrostia rugicollis, male syntype (type material of Perliodes nigrogranulosus, a synonym), copyright Naturhistorisches Museum Wien; B. Autolyca pallidicornis, male lec- totype, copyright Naturhistorisches Museum Wien; C. Anisomorpha paromalus, male lectotype, copyright Natural History Museum of London. 1906); from Metriophasma agathocles (Stal, 1875), Metriophasma baculus, and Metriophasma diocles in having concolorous hindwings (Redtenbacher 1906, Shelford 1909); and from Metriophasma pericles (Redtenbacher, 1906) in not having a median carina in the mesonotum (Redtenbacher 1906). This is the first time a species of the genus Metriophasma has been recorded for Mexico. The current record increases the num- ber of species in continental North America to 108, the number of species of North American Pseudophasmatidae to 7, and the number of genera of the region to 23. Metriophasma iphicles had been previously recorded in the Central American countries of Honduras and Panama (Redtenbacher 1906, Shelford 1909), be- ing the northernmost record of the species and genus. It is likely that this genus originated in South America, as this is where the majority of its species are found. At least 2 species reached Cen- tral America: Metiophasma diocles and Metriophasma iphicles. Fol- lowing the tropical and subtropical areas found in the Atlantic versant, a population of Metriophasma iphicles reached Veracruz (Fig. 5). This distribution and tropical migration pathway has been observed in other animals, even those with low vagility, such as pit vipers (Saldarriaga-Cordoba et al. 2017), so it is not surprising that a winged species could have easily followed it. The presence of this species is expected in more southern states, such as Chiapas or Oaxaca, as well as in northern Central Amer- ican countries, such as Guatemala and Belize. Recently, there have been several instances of cryptic diversity in Phasmatodea; therefore, further molecular and morphological (internal gen- italia) studies should follow to confirm whether this disjunct population is indeed Metriophasma iphicles or a closely related but undescribed species of the same species complex. Fig. 5. Map of Mexico; pink circle points to the new record of Metriophasma iphicles. Key to the North American Pseudophasmatidae The following key works in adults of any sex from either the USA or Mexico. It follows keys and descriptions present in the works of Redtenbacher (1906), Shelford (1909), Conle and Henne- mann (2002), Zompro (2005), Gorochov and Berezin (2008), and Lépez-Mora and Llorente-Bousquets (2018). Care must be taken when examining specimens from neotropical Mexico, as some taxa might still be unreported or undescribed for this region. 1 Macropterous (tegmina reduced, hindwings well-developed, capable Oftheht hie seA AB) Sot Cee arc Moulen al entencdde Manel snss Muenselas ales 2 - Apterous (completely wingless and flightless) [Fig. 4B, C]............... 3 JOURNAL OF ORTHOPTERA RESEARCH 2023, 32(2) 176 2 Hindwings reaching the ninth abdominal tergite [Figs 1B, 3A-—C]; mid- dle and hind femora with a ventral carina. In the region, it has only been recorded in southeastern Mexico (Veracruz)........... Metriophasma iphicles - Hindwings shorter, not reaching the eighth abdominal tergite [Fig. 4A]; middle and hind femora without ventral carina. In the re- gion, it has only been recorded in southwestern Mexico (Colima) .... iAP aisaoen esha AsRtisaiva tan taaaneqleRiroariengh, amen hen iaReestu Renee bewsilters Agrostia rugicollis 3. Body concolorous [Fig. 4B] (juveniles can be speckled); forefemur most- ly straight [Fig. 4B]. Jd with ninth abdominal tergite with forcep-like lobes; 22 subgenital plate large, reaching the tip of the anal segment or surpassing it. Found in southernmost Mexic0..........::ceeee Autolyca*: 4 - Body striped (sometimes faded, but nonetheless visible) [Fig. 4C]; forefemur depressed and curved basally [Fig. 4C]. d¢ with small, non-convex poculum; 99 subgenital plate short, reaching, at max, halfway the anal segment. Found in southeastern USA as well as the Yucatan/Penninsulaiin Mexico. ween iadiatetdten tien Anisomorpha: 5 4 9 operculum with rounded apex; 3'¢ genital plate apex bifurcated with 2 tubercle-like projections ...........ccccccsceeesseeeeeeeesees Autolyca elena - 9 operculum with angular apex; ¢ genital plate apex en- LIT Ce onl pe oen aunt aM meatal ie ante wey Sect eRe ua: Autolyca pallidicornis 5 Pronotum slightly wider than long; 4 mesonotum 1.8 times larger than wide, at max; 99 mesonotum 1.4 times larger than wide, at max. Found in the Mexican Yucatan Penninsula.......... Anisomorpha paromalus - Pronotum slightly longer than wide; 34 mesonotum 2.4 times larger than wide, at minimum; 29 mesonotum 1.6 times larger than wide, at minimum. Found in southeastern USA\......c::..:4..seecciteeteetle 6 6 Middle mesonotal suture clearly visible; with a distinctive black stripe that runs from the head to abdominal tergite X; 4 tergite X with a concave prolateral incision, 3d total body length 38-50 mm, 99 total body length 58-85 mm..............eeee Anisomorpha buprestoides - Middle mesonotal suture not clearly defined; with a discontinuous and not clearly defined black stripe that usually runs from the head to abdominal tergite IX, although it can appear as a faint line in tergite X; do tergite X without incisions; 33 total body length 22-35 mm; 2 total body length 40-55 mm... Anisomorpha ferruginea Remarks on Autolyca: Some authors have stated that “the known representatives of Autolyca are invariably apterous” (Conle et al. 2009), which holds true for the majority of the described species. However, Autolyca albifrons Redtenbacher, 1906 was described to possess brachypterous hindwings but not tegmina (Redtenbacher 1906). The origin of the lone male holotype is unknown but is unlikely to be from New Caledonia (Redtenbacher even marked this dubious locality with “(?)”). Much more recently, Bank and Bradler (2022) mentioned an undescribed brachypterous species of Autolyca from Panama. The revision and redescription of the holotype of Autolyca albifrons, the collection of more material, and the description of the undescribed Panamanian Autolyca are need- ed to begin resolving this matter. Acknowledgements We want to thank Dr. Paul D. Brock and the Phasmida Spe- cies File Website as well as the Naturhistorisches Museum Wien and the Natural History Museum of London for providing us with some of the photographs used in the current publication. We also want to thank Biol. Rosamond Iones Coates and Biol. Martha Madora-Astudillo for their help at the station and our friends and colleagues David Ortega, Biol. Nora Nino, Edilia Lopez, Victor M. DE LUNA, R. GARCIA-BARRIOS, G. CUELLAR-RODRIGUEZ AND U. 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