Research Article |
Corresponding author: Atilano Contreras-Ramos ( acontreras@ib.unam.mx ) Academic editor: Shaun Winterton
© 2019 Daniel Reynoso-Velasco, Atilano Contreras-Ramos.
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.
Citation:
Reynoso-Velasco D, Contreras-Ramos A (2019) Taxonomic review of the mantidfly genus Nolima Navás (Neuroptera, Mantispidae, Calomantispinae). ZooKeys 853: 131-158. https://doi.org/10.3897/zookeys.853.30317
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The mantidfly genus Nolima Navás, 1914 (Neuroptera, Mantispidae, Calomantispinae) is herein revised. Nolima is endemic to the New World, ranging from the southwestern United States south to Costa Rica. Nolima infensa Navás, N. pinal Rehn, and N. victor Navás are redescribed, while the new species Nolima costaricensis Reynoso & Contreras, sp. nov. is described from Costa Rica. The species N. dine Rehn and N. kantsi Rehn are synonymized with N. pinal. Additionally, the species N. praeliator Navás and N. pugnax Navás are synonymized with N. victor, for which a lectotype is designated. New distribution records are provided from Guatemala and Honduras for Nolima infensa, the state of Nevada in western United States for N. pinal, and the state of Puebla in central Mexico for N. victor. An illustrated key and a distribution map are presented.
Lacewings, mantispids, New World, new species, taxonomy
Mantidflies, mantid lacewings, or mantispids (Mantispidae) are distinctive within the Neuroptera because of their raptorial forelegs (Fig.
Four subfamilies of mantidflies are currently recognized: Calomantispinae, Drepanicinae, Mantispinae, and Symphrasinae (
This study consists of the taxonomic revision of the New World genus Nolima Navás, which previously included seven nominal species and is the sole representative of the subfamily Calomantispinae in this part of the world. The distribution of the species in this genus ranges from southern United States south to Costa Rica in Central America. The original descriptions of the species in Nolima were mainly based on the pigmentation pattern on the head and prothorax. We noticed that those patterns were not consistent and of little help for species identification; for that reason, we decided to study the group and explore other characters (e.g., male genital structures) to better circumscribe the different species.
The specimens examined during this study, including species from other genera of Mantispidae (Table
Comparative taxa examined to establish diagnostic features of the genus Nolima Navás.
Taxon | Distribution | Sex / Repository |
---|---|---|
Calomantispinae | ||
Calomantispa picta Stitz | Australia: Australian Capital Territory: Canberra. | 1♂, 1♀ / |
Calomantispa spectabilis Banks | Australia: Queensland: Herberton. | 1♂ / |
Calomantispa venusta Lambkin | Australia: Australian Capital Territory: Mount Gingera. | 1♀ / |
Australia: Australian Capital Territory: Lee’s Spring. | 1♂ / |
|
Australia: New South Wales: South Black Range. | 1♀ / |
|
Drepanicinae | ||
Drepanicus chrysopinus Brauer | Chile: Los Ríos: Valdivia. | 1♂, 1♀ / |
Gerstaeckerella chilensis (Hagen) | Chile: Metropolitana de Santiago: Til-Til, Santa Maria. | 1♂ / |
Theristria stigma (Esben-Petersen) | Australia: Queensland: West Claudie River. | 1♀ / |
Theristria storeyi Lambkin | Australia: Queensland: Kennedy River. | 1♂ / |
Mantispinae | ||
Climaciella brunnea (Say) | Mexico: Veracruz: San Andrés Tuxtla. | 1♀ / |
Mexico: Veracruz: Santiago Tuxtla. | 1♂ / |
|
Dicromantispa interrupta (Say) | Mexico: Jalisco: Estación de Biología Chamela. | 1♂ / |
Dicromantispa sayi (Banks) | Mexico: Chihuahua: El Jaquex. | 1♀ / |
Zeugomantispa virescens (Rambur) | Mexico: San Luis Potosí: El Limoncito. | 1♂ / |
Symphrasinae | ||
Plega dactylota Rehn | México: Baja California Sur. | 1♂ / |
Trichoscelia sp. 1 | Mexico: Sonora: Cerro Verde. | 1♂ / |
Pinned specimens were placed in an airtight chamber with a solution of water and phenol for rehydration for approximately 24 hours. The abdomen of males was dissected and placed in 10% KOH for approximately 10 hours at room temperature, then rinsed in distilled water. The abdomen of each females was treated similarly, except that it was stained with Chlorazol Black E (in ethanol) to enhance contrast of the internal structures. The dye was injected with a syringe into the abdominal cavity for approximately 10 seconds, then the dissected abdomen was transferred to 70% ethanol and the dye was rinsed out. For observation, the abdomen was placed in a Petri dish with glycerin. A Zeiss Stemi SV11 stereomicroscope with 10× eyepieces and 1.0× and 2.5× main objectives (with a zoom magnifying range of 0.6–6.6×) was used for morphological examination. After examination, the dissected abdomens were stored in genitalia microvials with glycerin and pinned under the corresponding specimen. Pencil drawings were elaborated with a camera lucida attached to the stereomicroscope, which were later inked and scanned. Digital images were obtained by use of a Nikon SMZ25 stereomicroscope coupled with the Nikon NIS-Elements Imaging Software. Final figures were prepared with Photoshop CS5 (Adobe Systems Inc., San Jose, California).
This study mainly follows
External and internal structures of males and females where evaluated to serve as potential diagnostic features. The morphology of the female genitalia was found to be conserved and similar among the specimens examined. For that reason, only a generic description of the structures is provided. Attributes related to the presence and position of bristle-bearing chalazae and the pigmentation pattern on the forelegs of both sexes were found to be informative, although the most reliable diagnostic features were related to characteristics of the male abdominal cuticle and genital structures.
The genus Nolima was erected (
As proposed by
Nolima
Navás, 1914: 100–101 (original description, gender: feminine, etymology: anagram of Molina, type species by original designation: Nolima victor);
Bellarminus
Navás, 1914: 102–103 (original description, gender: masculine, etymology: after the Italian cardinal Roberto Bellarmino, type species by original designation: Bellarminus pugnax);
The genus Nolima can be distinguished by the following combination of characters (character states in parentheses are generally exhibited by other mantispid genera): a) Sc comes in contact with C near the middle of costal margin and distal to the base of pterostigma on the forewing (at apex of 2/3 of costal margin and proximal to pterostigma), b) M diverging from R distal to 1m-cu on the forewing (proximal to 1m-cu), c) abdominal terga and sterna or only terga of the male with circular or polygonal structures, respectively, d) male mediuncus apex strongly projecting posteriorly and deeply bifid (shallowly indented), and e) female spermatheca with accessory gland (generally without accessory gland, but if present then associated to copulatory bursa, e.g., species of Calomantispa).
General. Coloration pale yellow, with dark brown pigmentation as stripes or marks in specific areas (detailed in the text below).
Head. Hypognathous. Vertex with a rhomboid protuberance covering nearly its entire area; vertex marking M-shaped, extending behind antennal sockets, where can be bifurcated, if bifurcated then one branch extends posteriorly, parallel to anterior ocular margin, additional branch generally extends anteriorly on frontogenal furrow, or extends on frontogenal and epistomal furrows; vertex with a pair of irregular marks originating posteromedially, extending anteriorly along the coronal suture, then angled at 45° toward anterior ocular margin, reaching the rhomboid protuberance, sometimes converging with upper part of M-shaped mark. Frons generally with pair of semicircular marks. Clypeus and labrum, each sometimes with a medial semicircular mark. Antennal flagellomeres dark brown, as long as wide in basal third of flagellum, twice as long as wide in distal two thirds in frontal view. Mandibles with pigmentation on inner and outer edges.
Thorax. Prothorax straight in lateral view, with pigmentation, bristle-bearing chalazae on pronotum and anterolateral and anteroventral areas, a pair of pale spots anterolaterally in dorsal view. Mesothorax with conspicuous mesoscutal and scutoscutellar sutures; scutum generally with two longitudinal stripes anterior to suture and four posterior to suture, two medial and two lateral; scutellum with color pattern variable; pleural area generally with pigmentation. Metathorax with mesoscutal suture obsolete, scutoscutellar suture conspicuous; scutum generally with an M-shaped mark medially, a longitudinal stripe on each side of medial mark. Forecoxa with bristle-bearing chalazae. Forefemur with dorsal margin slightly convex, midsection in dorsal view approximately twice as wide as apex; longitudinal row of spines on ventral side weakly compressed laterally; tibia arched, two thirds as long as femur, with ventral carina; first tarsomere more than twice as long as second. Middle and hindleg not modified, finely and evenly setose. Forewing (Fig.
Abdomen, male (Fig.
Abdomen, female (Fig.
This genus is endemic to the New World, ranging from southwestern United States to Costa Rica (
Little is known about this topic; the available information is related to the cytogenetics and larval diet of Nolima pinal.
The word Nolima is an anagram of Molina, in honor of Luis de Molina (1535–1600), a Jesuit priest who was born in the city of Cuenca, Spain (
Most reliable diagnostic features are related to the external and internal genitalic morphology of males. Other traits (e.g., pigmentation, chalazae) are instructive for males and females but reliance on them alone may lead to misidentification.
1 | Forecoxa with bristle-bearing chalazae on ventral, lateral (basally), and dorsal surfaces; chalazae bases generally surrounded with dark brown pigmentation (Figs |
2 |
– | Forecoxa with bristle-bearing chalazae only on ventral surface; chalazae bases not pigmented (Figs |
3 |
2 | Forefemur dorsal surface with conspicuous dark brown circular marks around bases of chalazae (Fig. |
Nolima victor Navás |
– | Forefemur dorsal surface without dark brown marks around bases of chalazae (Fig. |
Nolima pinal Rehn |
3 | Male ectoprocts with dorsal margin strongly convex in lateral view (Fig. |
Nolima infensa Navás |
– | Male ectoprocts with dorsal margin slightly convex in lateral view (Fig. |
Nolima costaricensis sp. nov. |
It differs from other species in the genus as follows: a) male sterna I–VIII with circular structures only laterally (Fig.
This new species is described based on a single male specimen collected in southeastern Costa Rica, which unfortunately had lost pigmentation; therefore we were not able to specifically evaluate some of the characteristic markings.
Male. Head. Vertex with M-shaped mark with lower arms getting wider towards anterior ocular margin (Fig.
Thorax. Prothorax with pigmentation on entire surface of pronotum (Fig.
Abdomen. Terga and lateral surface of sterna I–VIII with circular structures, not in contact to each other (Fig.
It could not be assessed because only the holotype specimen is known.
Based on the collecting datum from the single specimen examined, adults of the species may be active during spring.
The species name is dedicated to Costa Rica, the only country from which this species is currently known.
The holotype is housed at the
Costa Rica: Puntarenas, Parque Internacional La Amistad, Sector Altamira.
This species is only known from its type locality, which is in the southeastern part of Costa Rica (Fig.
HOLOTYPE ♂ (by present designation): COSTA RICA: Puntarenas: P[arque] I[nternacional] La Amistad, Sector Altamira, Send[ero] Gigantes del Bosque, 1300–1400 m, 13-IV–14-V-2005, R. González, T[ram]p[a] Malaise, LS331300 571500 #83526, INB0004129281, INBIOCRI, Costa Rica (
Nolima infensus
Navás, 1924: 61–62 (original description);
Nolima infensa
Navás:
It differs from other Nolima species as follows: a) male sterna I–VIII with circular structures only laterally (Fig.
This species was described based on a single female specimen collected in Costa Rica;
Male. Head. Vertex with M-shaped mark not bifurcated behind antennal sockets (Fig.
Thorax. Prothorax with pigmentation on pronotum, except anterolateral pale yellow mark on each side of midline (Fig.
Abdomen. Terga and lateral surface of sterna I–VIII with circular structures, not in contact to each other (Fig.
Female. Pigmentation and setation generally same as for male.
In both sexes, the pair of marks on the frons may be absent. An irregular mark may be present on the clypeus. The pigmentation on the forefemur may be absent. Specimens may also present pigmentation on the mesopleural katepisternum and anepimeron, on the metapleural anepisternum and katepisternum, or lack pigmentation on the pteropleural area. The dorsal margin of male ectoprocts may be only slightly convex in lateral view.
Based on collecting data, adults of this species may be found active from May through August.
The holotype is housed at the
Costa Rica.
This species is distributed from central Mexico (Chiapas, Morelos, Oaxaca, Veracruz) south to Costa Rica (Puntarenas), including Guatemala (Zacapa) and Honduras (Comayagua, Yoro) (Fig.
Costa Rica; México: Morelos, Oaxaca (
HOLOTYPE ♀ (by monotypy): COSTA RICA: Nolima infensus Nav. P. Navás S.J. det [1st label], TYPE [2nd label], Museum Paris, Costa Rica, Paul Serre 192 [3rd label]. Microvial with last abdominal segments of the holotype in glycerine, pinned next to specimen: HOLOTYPE Nolima infensus Navás ♀, Genitalia in Glycerin BEARD [single label] (
COSTA RICA: Puntarenas: Las Alturas, 1500 m, 22-V-1992, F. Andrews & A. Gilbert, Nolima infensus det. N. Penny (1♀, 1 adult without abdomen
Nolima pinal
Rehn, 1939: 256–259, 263 (key, original description);
Nolima dine
Rehn, 1939: 256–257, 261–263 (key, original description);
Nolima kantsi
Rehn, 1939: 256–257, 260–262 (key, original description);
It differs from other species in the genus as follows: a) male sterna I–VIII with circular structures on nearly the entire surface (Fig.
Nolima pinal was described based on a single female specimen collected in Arizona, United States. In the original description the holotype was erroneously reported as a male specimen.
Male. Head. Vertex with M-shaped mark bifurcated behind antennal sockets, one branch extending posteriorly parallel to anterior ocular margin, additional branch extending anteriorly on frontogenal furrow (Fig.
Thorax. Prothorax with pigmentation on pronotum, except narrow pale yellow longitudinal stripe along midline and anterolateral pale yellow mark on each side of midline (Fig.
Abdomen. Terga and sterna I–VIII with circular structures, not in contact to each other, microsetae in space between circular structures (Fig.
Female. Pigmentation and setation generally same as for male, except the antennal scape, which presents pigmentation on entire posterior surface.
The mark located on the frontogenal furrow sometimes extends ventrally onto the epistomal furrow, a feature more common in females. The clypeus may present a single irregular mark medially. The anterolateral pale yellow mark of the pronotum sometimes exhibits pigmentation medially, giving an appearance of two marks. The forefemur may present two marks on the lateral surface, a trait more common in females. Some females from Texas exhibited three marks on the lateral surface of the forefemur. Also, the forefemur may present an elongate mark on the first half of the mesal surface. Sometimes the foretibia presents three dorsal marks. The membrane between apexes of ectoprocts may be slightly sclerotized.
The cytogenetics of 15 species of mantispids from 11 genera and three subfamilies has been studied to date (
.
The holotype is housed at the
United States: Arizona: Gila Co., Pinal Mountains.
This species is distributed in the southwestern United States (Fig.
United States: Arizona, New Mexico, Texas (
HOLOTYPE ♀ (by original designation): UNITED STATES: Arizona: [Gila Co.], base of Pinal M[oun]t[ain]s, Ariz. [1st label, with antennal flagellum glued], Sep[tember], D.K. Duncan [2nd label], Oak [3rd label], M.C.Z. type 23645 [4th label], Nolima pinal Rehn TYPE [5th label],
UNITED STATES: Arizona: [Pinal Co.], Peppersauce C[a]n[yon], Aug. 16, 1924 [1st label], Santa Catalina Mts. [2nd label], J.O. Martin Collector [3rd label], Nolima dine Rehn TYPE [4th label], California Academy of Sciences Type No. 4927 [5th label] (♀
UNITED STATES: Arizona: Cochise Co., Cave Creek Canyon, 3 mi W Portal, 31°53.023'N, 109°10.715'W, 5120 ft, 9-VIII-2000, A. Gilbert & N. Smith (1♀
Nolima victor
Navás, 1914: 101 (original description);
Nolima praeliator
Navás, 1914: 101–102 (original description);
Nolima pugnax
(Navás), 1914: 103 (original description);
It differs from other Nolima species as follows: a) male terga I–VIII with polygonal structures (Fig.
The original description of Nolima victor apparently was based on at least two specimens because in that work,
Male. Head. Vertex with M-shaped mark bifurcated behind antennal sockets, one branch extending posteriorly parallel to anterior ocular margin, additional branch not extending anteriorly (Fig.
Thorax. Prothorax with pigmentation on pronotum, except narrow pale yellow longitudinal stripe along midline and anterolateral pale yellow oval mark on each side of midline (Fig.
Abdomen. Terga I–VIII with polygonal structures, in close contact to each other (Fig.
Female. Pigmentation and setation generally same as for male.
Both sexes may exhibit a circular mark on clypeus and one on labrum. The pronotum may be yellowish, with pigmentation only on the chalazae. In females, the bifurcated M-shaped mark on the vertex may present the branch that extends anteriorly, on the frontogenal and epistomal furrows. The irregular marks that originate posteromedially on vertex may be fused to bifurcation of M-shaped mark that extends posteriorly. Also, the antennal scape may exhibit a small mark on the mesal surface and the pedicel may be pigmented on the posterior surface. Sometimes with small circular marks on entire surface of forefemur.
Based on collecting data from material examined, adults of this species are active from February through October.
Navás did not specify the etymology of the species name. However, the specific epithet victor is a Latin adjective meaning victorious. According to this, the name could be read as “Molina victorious,” a phrase acclaimed by Father Molina’s adherents when in 1607 Pope Paul V decided not to condemn the ideas of Molinism.
The lectotype is housed at the
México: Guerrero: Mpio. Chilpancingo de los Bravo, Xocomanatlán.
This species is distributed in Mexico (Chiapas, Guerrero, Hidalgo, Jalisco, Morelos, Oaxaca, Puebla, Querétaro) and Guatemala (Baja Verapaz) (Fig.
Guatemala: Baja Verapaz; México: Chiapas, Guerrero, Hidalgo, Jalisco, Morelos, Oaxaca, Querétaro (
LECTOTYPE ♀ (by present designation): MEXICO: Guerrero: [Mpio Chilpancingo de los Bravo], Xucumanatlan [Xocomanatlán], 7000 ft, July [no year], H.H. Smith [1st label, with antennal flagellum glued], Godman-Salvin Collection 1913-214 [2nd label], Typus [3rd label], Nolima victor ♀ Nav. Navás S.J. det. [4th label], Genitalia prep. in vial on other pin made 20-V-1969, R.G. Beard # 1008 [5th label], Type H.T. [6th label]. Microvial with last abdominal segments of the holotype in glycerine, pinned next to specimen: HOLOTYPE ♀ Nolima victor Navás 1909, ♀ Genitalia in glycerine [1st label], R.G. Beard # 1008 Genitalia prep. of Nolima victor Navás 1909 ♀ Holotype in glycerine [2nd label] (
[MEXICO]: Guerrero: [Mpio Chilpancingo de los Bravo], Omilteme [Omiltemi], 8000 ft, Aug., H.H. Smith [1st label], Godman-Salvin Collection 1913–214 [2nd label], Nolima praeliator Nav. Navás S.J. det. [3rd label], Typus [4th label], Nolima victor Navás ♀ D. Reynoso-Velasco det. 2008 [5th label],
MÉXICO: Chiapas: Hwy 199, 11 km NE San Cristóbal, 8000 ft, 25-V-1987, D.A. Rider et al. (1♀
Thanks are due to the curators and supporting personnel of museums that kindly provided specimens for this study, including: Wendy A Whitby (