Limonius Eschscholtz, 1829: 33

Etzler, Frank E., 2019, Generic Reclassification of Limonius Eschscholtz, 1829 (Elateridae: Dendrometrinae) sensu Candèze 1860 of the World, Zootaxa 4683 (3), pp. 301-335 : 313-319

3. Limonius Eschscholtz, 1829: 33 , 79 species

Type species: Elater minutus Linneaeus, 1758 , designated by Curtis 1838. LeConte 1853: 428; Candèze 1860: 377; Reitter 1905: 16; Schwarz 1907: 193; Leng 1920: 168; Lane 1971: 21; Leseigneur 1972: 180; Sánchez-Ruiz 1996: 108; Johnson 2002: 169; Leseigneur and Mertlik 2007; Cate 2007: 165; Al Dhafer 2009; Han et al 2012: 53.

Cidnopus Thomson, 1859: 106 .

Type species: Elater nigripes Gyllenhal, 1808 (= Elater pilosus Leske, 1785 ), by monotypy. Leseigneur 1972: 175; Kishii 1987: 81; Platia 1994: 102, Sánchez-Ruiz 1996: 105; Cate 2007: 163; Han et al 2012: 49.

Dendrometrus Gistel, 1848 : xi [unjustified replacement name for Limonius ]. Noted as synonym of Limonius ( Sánchez-Ruiz 1996: 108) .

Kibunea Kishii, 1966: 46 .

Type species: Limonius eximius Lewis, 1894: 318 , by original designation. Kishii 1987: 84, Platia 1994: 106. Synonymyzed by Sánchez-Ruiz 1996: 108.

Micrathous Lane, 1971: 24 , new synonymy.

Type species: Athous brevis Van Dyke, 1932: 379 , by original designation. Johnson 2002: 169.

Neoathousius Schimmel and Platia, 1991: 881 , new synonymy.

Type species: Neoathousius loebli Schimmel and Platia, 1991 , by original designation: Schimmel and Platia 1994; Cate 2007: 166.

Nothodes LeConte, 1861: 171 .

Type species: Limonius dubitans LeConte, 1853 , by monotypy: Leng 1920: 168; Kishii 1987: 82; Platia 1994: 112; Sánchez-Ruiz 1996: 108; Cate 2007: 165.

Solskyana Dolin, 1978: 45 , new synonymy.

Type species: Limonius villiger Solsky, 1881 , by original designation. Cate 2007: 166.

The following are suggested species groups. I have included generic names in parentheses near the species groups when a type species is included in the group. For this genus, I have grouped both Nearctic and Palearctic species together under each species-group. This will make it easier for future revisionary work. No consistent character was discovered to be able to separate these groups at the generic-level, with groups blurring together. Species-level synonyms for Palearctic species are not provided, but can be found in Cate (2007). For the Nearctic region, species level synonyms follow Al Dhafer (2009) unless noted.

Minutus species-group ( Limonius , Kibunea )

Nearctic species (4)

Limonius aurifer LeConte, 1853: 429 .

Limonius basilaris (Say, 1823) : 172 ( Elater ).

Limonius lanei Van Dyke, 1932: 350 .

Limonis semianeus LeConte, 1853: 432 removed from synonymy.

Palearctic species (7)

Limonius approximans Lewis, 1894: 198 .

Limonius eximia Lewis, 1894: 318 .

Limonius ignicollis Lewis, 1894: 197 .

Limonius kouichiana (Kishii, 1989) : 63 ( Kibunea ).

Limonius minutus ( Linnaeus, 1758) : 406 ( Elater ).

Limonius narukawai (Kishii, 1993) : 151 ( Kibunea ).

Limonius scutellaris (Dolin, 2003) : 36 ( Cidnopus ).

Parvulus species-group ( Micrathous )

Nearctic species (4)

Limonius aeger LeConte, 1853: 431 .

Limonius brevis ( Van Dyke, 1932) : 379 ( Athous ) new combination.

Limonius quercinus (Say, 1825) : 262 ( Elater ).

Limonius sordidus ( Van Dyke, 1932) : 380 ( Athous ) new combination.

Palearctic species (3)

Limonius marginicollis Lewis, 1894: 196

Limonius parallelus Motschulsky, 1860: 111 .

Limonius parvulus (Panzer, 1799) : 7 ( Elater ).

Consimilis species-group ( Solskyana )

Nearctic species (5)

Limonius beutenmuelleri Van Dyke, 1932: 351 .

Limonius consimilis Walker, 1866: 325 . [see note below]

Limonius nitidulus Horn, 1871: 315 .

Limonius jonesi Lane, 1965: 182 .

Limonius seibertae Etzler, 2018: 863

Limonius pappi Becker, 1976: 689 .

Palearctic species (6)

Limonius crassipes Schwarz, 1900: 105 .

Limonius hirtus ( Dolin, 1978) : 48 ( Solskyana ) new combination.

Limonius koltzei Reitter, 1895: 256 .

Limonius montivagus Lewis, 1894: 194 . [see notes below]

Limonius nigronitidus ( Han and Lee, 2012) : 50 ( Cidnopus ) new combination.

Limonius villiger Solsky, 1881: 237 .

Pilosus species-group ( Cidnopus )

Nearctic species (3)

Limonius auripilis (Say, 1823) : 172. ( Elater )

Limonius kuschei Van Dyke, 1932: 349 .

Limonius pectoralis LeConte, 1853: 391 .

Palearctic species (6)

Limonius marginellus brusteli (Leseigneur, 2004) : 231 ( Cidnopus ) new combination.

Limonius marginellus marginellus Perris, 1864: 284 .

Limonius pilosus (Leske, 1785) : 11 ( Elater ).

Limonius pseudopilosus (Platia and Gudenzi 1985) : 21 ( Cidnopus ) new combination.

Limonius schurmanni (Platia and Gudenzi, 1998) : 53. ( Cidnopus ) new combination.

Limonius turcicus (Platia, 2004) : 40. ( Cidnopus ) new combination.

Canus species-group ( Nothodes , Neoathousius )

Nearctic species (13)

Limonius agonus (Say, 1839) : 171 ( Elater ).

Limonius anceps LeConte, 1853: 433 removed from synonymy.

Limonius californicus (Mannerheim, 1843) : 238 ( Cardiophorus ).

Limonius canus LeConte, 1853: 433 .

Limonius dubitans LeConte, 1853: 433 removed from synonymy.

Limonius ectypus (Say, 1839) : 167 ( Elater ).

Limonius infuscatus Motschulsky, 1859: 371 removed from synonymy.

Limonius longulus Champion, 1896: 455 .

Limonius mandibularis Motschulsky, 1859: 370 .

Limonius pilosulus Candèze, 1891: 149 removed from synonymy.

Limonius quadraticollis Candèze, 1860: 412 .

Limonius rectangularis Fall, 1934: 30 .

Limonius subauratus LeConte, 1853: 433 .

Palearctic species (26)

Limonius aeruginosus (Olivier, 1790) : 33 ( Elater ).

Limonius baconi Candèze, 1860: 412 .

Limonius brancuccii ( Schimmel and Platia, 1991) : 884 ( Neoathousius ) new combination.

Limonius decorus (Gurjeva, 1975) : 169 ( Athousius ) new combination.

Limonius exiguus ( Schimmel and Platia, 1991) : 885 ( Neoathousius ) new combination.

Limonius hartmanni (Schimmel, 1998) : 151 ( Neoathousius ) new combination.

Limonius hiermeieri ( Schimmel and Platia, 1991) : 888 ( Neoathousius ) new combination.

Limonius hubeiensis (Kishii and Jiang, 1996) : 131. ( Cidnopus ) new combination.

Limonius kubani (Schimmel, 1996) : 172 ( Neoathousius ) new combination.

Limonius loebli ( Schimmel and Platia, 1991) : 890 ( Neoathousius ) new combination.

Limonius longicornis ( Schimmel and Platia, 1991) : 890 ( Neoathousius ) new combination.

Limonius macedonicus ( Cate and Platia, 1989) : 185 ( Cidnopus ) new combination.

Limonius manaliensis ( Schimmel and Platia, 1991) : 892 ( Neoathousius ) new combination.

Limonius marginipennis Lewis, 1894: 195 . [see note below]

Limonius miandamensis ( Schimmel and Platia, 1991) : 892 ( Neoathousius ) new combination.

Limonius minusculus ( Schimmel and Platia, 1991) : 101 ( Neoathousius ) new combination.

Limonius nigrinus Candèze, 1900: 94 .

Limonius platiai Mertlik, 1996: 203 (Cidnopus) new combination.

Limonius recticornis ( Schimmel and Platia, 1991) : 893 ( Neoathousius ) new combination.

Limonius riesei Platia, 1988: 9 (Athousius) new combination.

Limonius rusticus ( Schimmel and Platia, 1991) : 894 ( Neoathousius ) new combination.

Limonius ruzenae Laibner, 1977: 388 .

Limonius schimmeli new replacement name.

Neoathousius ferrugineus Schimmel and Platia, 1991: 88 [HN], not Limonius ferrugineus Buysson, 1902 . Limonius sinensis ( Schimmel and Platia, 1994) : 233 ( Neoathousius ) new combination.

Limonius singularis ( Schimmel and Platia, 1991) : 894 ( Neoathousius ) new combination.

Limonius stapfi (Schimmel, 2007) : 183 ( Neoathousius ) new combination.

Limonius wittmeri ( Schimmel and Platia, 1991) : 895 ( Neoathousius ) new combination.

Species incertae sedis

Limonius obienesis Cherepanov, 1966: 23 , Western Siberia.

Limonius substriatus (Gebler, 1833) : 279 ( Elater ) “Siberia”.

Nomina dubia in Cate 2007 Limonius araratensis Faldermann, 1835: 168 .

Diagnosis. This genus can be distinguished from other genera in the Limonius genera-group by the non-emarginate posterior edge of the hypomeron and the procoxal cavity narrowly open posteriorly. Other characters are quite variable among species.

Generic Description. Small to medium sized, 5.5 mm to 13 mm in length, sub-cylindrical. Head with simple or umbilicate punctures; frons weakly depressed or rather flattened; frontal margin variable, entirely carinate to absent medially, straight to strongly undulate in speciemens with margin present. Antennae variable and sexually dimorphic, usually longer in males, extending past posterior edge of pronotum by one or two antennomeres, and reaching posterior edge in females, although exceptions are common; 2 nd and 3 rd antennomeres usually subcylindrical, ratio of 2 nd and 3 rd variable, ratio of 2 nd and 3 rd to 4 th variable, usually serrated from 4th to 10th antennomeres, although some species have a weakly serrate 3 rd antennomere, 11 th ovoid. Pronotum variable and sexually dimorphic, usually appearing longer than wide but actually as wide as long, although some species clearly longer than wide or wider than long; anterior angles variable, sometimes expanded; hind angles short, broad at base, often with short single carina; posterior margin lacking incisures at base of hind angles and usually lacking notches anterior of scutellum; prosternal sutures double, anterior end variable, distinctly grooved and strongly excavate anteriorly to nearly closed with barest hint of groove; posterior edge of hypomeron not emarginate posteriorly, straight and “ear-like” or slightly sinuate; prosternal process expanded laterally; procoxal cavities narrowly opened. Scutellum variable, simple, smoothly convex, or with medial longitudinal ridge, often an oblong oval shape (Fig. 14) or a broad sub-quadrate shape (Fig. 15). Mesepisternum sometimes not reaching mesocoxal cavity (Fig. 24). Legs slender, moderately long; tarsomeres simple, often with dense setal pads ventrally; claws simple, lacking basal setae. Male genitlia variable, but always with apical expansions on the parameres (Figs. 17–23). Female internal genitalia variable, but all bearing two distinct colleterial glands off of weakly sclerotized uterus; uterus never with distinct sclerotizations (Figs. 25–34).

Discussion. This concept of Limonius returns to the older, broader concept of the genus. The biggest issue with the concept of this genus was the confusion about the type species during the mid-20 th century ( Sánchez-Ruiz 1996). Due to this confusion, many authors tried to move some species to other genera without looking at broader patterns, leaving this group essentially unworkable.

Species that are listed as incertae sedis were treated as Limonius by Cate (2007), but I have not been able to examine specimens of these names, nor have I been able to look at the original descriptions. As Cate spent considerable time on the Palearctic Catalogue, I accept his placement, but am unable to place these species into a speciesgroup. The nomina dubia is a name that Cate (2007) did not place in any genus. I have included the name here to have a full catalog of species names for the genus.

The order of species groups above is important, as the order is a hypothesis of which species groups have closer relationships, which are discussed below. No attempt at a phylogeny has been attempted for species groups or species within each species group.

Unlike the other genera presented here, there is quite a bit of variation in the internal female genital tracts. Based on my observations, there are at least six types present within the genus. These genitalic groups likely represent true species-groups, however, I am unable to find consistent external morphological characters to readily distinguish genera. In many instances, there is more variation within a species group than between species groups, increasing the difficulties in choosing informative characters. I have chosen the conservative approach and recognized a large, variable genus until all species can be examined in detail and a phylogenetic analysis conducted.

Despite the variation, all female internal genitalia do appear to follow a similar pattern. All examined species bear two distinct colleterial glands off the uterus. The uterus is also often slightly more sclerotized that the bursa copulatrix, which is indicated on the drawings by brown lines. Unlike the genera Pheletes and Tetralimonius , there are no distinct sclerotized sections of the uterus.

The Minutus species-group is equivalent to Limonius sensu stricto. It is based on members sharing the characters of flared anterior margins of the pronotum, and a narrow aedeagus (all similar to Fig. 17). The female internal genitalia belong to what I call Type A (Fig. 25). The bursa copulatrix is often weakly sclerotized basally, bears at least two accessory glands, a smaller one basally and a moderate one medially, and a tubular extension at the end. Some species (not figured) also bear another gland near the tubular extension (diverticulum of Becker 1956). Some authors (e.g. Platia 1994) have used the flared anterior margins to separate this group at the genus level. However, this character also repeatedly appears in the genus Gambrinus . An unreported character is that L. minutus , the type species of Limonius , possesses a mesocoxal cavity where the mesepisternum does not reach the mesocoxal cavity (Fig. 24). This character is shared by all North American species in this species group. The modern concept of L. minutus was established by Leseigneur and Mertlik (2007). Larvae have only been associated with a few Palearctic species (e.g. Dolin 1978), and are of the typical Limonius type, with the ninth abdominal segment similar to Fig. 72, but with urogomphal prongs reduced to tubercles.

The Parvulus species-group is equivalent of Nothodes auct . The misuse of the generic name for this group stems from confusion of the Japanese species L. marginicollis and L. marginipennis , both described by Lewis (1894) (C. M. F. von Hayek, unpublished BMNH collection notes). I have examined both types at the BMNH and only the later species is closely related to the type species of Nothodes . Members of this group are intermediate between the preceding group and the following group (Consimilis species-group). Members also share characters with the other two species groups (Pilosus species-group and the Canus species-group). Species in both regions look very similar to species in the Minutus species-group, but lack the flaired anterior margins of the pronotum. The male genitalia of species in this group are also intermediate, combining characteristics of both groups (e.g. Figs. 18 and 19). For the mesocoxal cavity, the mesepisternum just reaches the cavity in L. parvulus , and narrowly reaches to appearing closed in L. aeger , suggesting this character is variable within the species group.

Larvae are associated with species from both regions, with the larva of L. aeger described for the Nearctic ( Glen 1950), and L. parvulus for the Palearctic ( Dolin 1978). These larvae belong to the same group as the known larvae in the Minutus species-group. I fully expect this group to be artificial when a phylogenetic study of the genus is conducted.

To the Parvulus species-group, I also add two species from California formerly placed in the genus Micrathous . This genus was defined by small, setose lobes on the tarsi, and grooved prosternal sutures, which separated these species from Athous , the genus the two included species were described in. However, careful examination revealed that the setose lobes are better classified as dense setose pads, a common character in Limonius species. No other characters were discovered to recognize these two species as belonging to a distinct genus.

The female internal genitalia of species I place into the Parvulus species-group are quite variable relative to other groups. There are four main genitalia types known, all possessing a curved bursa copulatrix. Type B (Fig. 26), found in L. aeger , is similar to Type A, but lacks the small accessory gland, a more curved bursa copulatrix and a corkscrew-shaped tubular extension. It appears similar to the drawing of the genitalia for L. pseudopilosus in Fig. 6d of Han et al. (2012). Type C (Fig. 27), found in L. quercinus , is strongly modified, possess a tubular bursa copulatrix without a clear separation from the uterus and lacking distinct accessory glands or a tubular extension. The bursa copulatrix also bears a heavily sclerotized area at the inner curve. However, the aedeagus of L. quercinus is very similar to that of L. aeger (Fig. 18). While the male genitalia of L. parvulus appear similar to those found in the Consimilis species-group (Fig. 19), the female internal genitalia link this species to others within the speciesgroup (Fig. 28). The female internal genitalia of L. parvulus share many characteristics with other groups. It shares the basal accessory gland and basal sclerotization of the bursa copulatrix with Type A genitalia. It also has the inner heavy sclerotization of the bursa copulatrix as in Type C (insert, Fig. 28). Finally, it has a medial accessory gland and general bursa copulatrix shape as in Type F (below). Due to this combination of many characters, I consider this a Type C/ F combination. Finally, L. brevis (formerly in Micrathous ) has Type D genitalia (Fig. 30), with a long, curved bursa copulatrix, lacking basal or medial accessory glands. This shares some characteristics with species in the Pilosus species-group (see below).

The Consimilis species-group contains species previously placed in Cidnopus and Solskyana and may also be an artificial grouping. These species are grouped together by the slightly sinuate posterior edge of the hypomeron (Fig. 13), and the widened parameres of the aedeagus (Fig. 20). The majority of specimens have the prosternal sutures strongly excavate anteriorly, with the antennae able to fit in a short groove between the hypomeron and the prosternum. In all examined specimens the mesepisternum does not reach the mesocoxal cavity, similar to the Minutus species-group. The female internal genitalia are also quite distinctive (Fig. 29), bearing a cork-screw shaped, elongate bursa copulatrix ending in many accessory glands and a long tubular extension. I consider this as Type E genitalia. Figure 29 should be compared to Figs. 3b and 3c in Han et al. (2012).

Some of the species in the Consimilis species-group require explanation. Until recently, considerable confusion existed about L. consimilis . These issues are discussed in detail in Etzler (2018), where a key to the North American species of this species group is also provided.

Limonius montivagus is also an interesting case. Many authors treat this species as a Gambrinus (i.e. Cate 2007). However, I have examined the type and it seems better placed in the Consimilis species-group. The slightly sinuate posterior edge of the hypomeron may have led to this placement in Gambrinus .

Finally, the two species formerly in Solskyana ( L. hirtu s and L. villiger ) deserve comment. The genus was described by Dolin (1978) based on larval characters. He stated that they had a double pointed mandible (Fig. 67) compared to typical Limonius (Figs. 65, 69). I examined two specimens identified as L. villiger , one from the NMPC and one at the BMNH. I am unable to find any characters to distinguish this species at the genus level. While Dolin’s (1978) larval key is a foundational work, he doesn’t state how he made the associations or numbers of individuals he examined. Due to this, I have taken the conservative approach and included these species here until a larger phylogenetic study can be conducted to determine their relationship and the strength of the mandibular character in the larvae.

The Pilosus species-group encompasses species that are apparently closely related to the type species of Cidnopus . These species all share excavated anterior ends of the prosternal suture, and strongly hooked parameres of the aedeagus (Fig. 21). Species are intermediate between the previous group and the following (Canus species-group). The mesepisternum does not reach the mesocoxal cavity in L. auripilis , barely reaches it in L. pectoralis , and clearly reaches it in L. pilosus , again showing variability of this character within a species group.

The understanding of internal female genitalic characters for this group is hindered by a lack of availability of specimens for dissection. Illustrations of L. pseudopilosus ( Han et al. 2012, Fig. 6d) share a general shape with that of L. aeger (Fig. 26). I also studied the internal genitalia of L. auripilis , and they are quite similar to L. brevis (Fig. 30), lending additional support in synonymizing Micrathous with Limonius . Of the approximately 20 specimens of L. pilosus I have been able to examine in detail, all have been males, leaving the female internal genitalia of that species unknown.

Of the species group, only L. pilosus has an associated larva, which is unique for the genus. While the adults are typical Limonius , the questionably associated larvae are not. The larvae possess stemmata and have a singlepointed nasale (Fig. 64), neither of which is known in other Limonius species ( Glen 1950, Dolin 1978).Additionally, the mandible (Fig. 66) is subtly different than typical Limonius (Fig. 65), bearing tubercles on the main tooth. This association needs to be re-confirmed, since I predict that the original match is based on miss-associated specimens and the error has been simply repeated in subsequent literature. It is unclear what these unknown larvae are, but it is likely a common northern European species.

The final group is the Canus species-group that encompasses species formerly in the genus Neoathousius , includes the type species of Nothodes , and part of Cidnopus . The group name was first suggested by Lanchester (1946) for some North American species, and I have broadened the definition here. The Canus species-group encompasses a wide variety of characters, with some species having excavated anterior ends of the prosternal suture, while others are nearly closed anteriorly. The variability of characters is best demonstrated with the Nearctic species of the Canus species-group. There is a wide range of variation between species with frontal margins (Figs. 35 and 36), antennomere ratios, the anterior ends of prosternal sutures (Figs. 37 and 38), types of ovipositors (Figs. 39–42), and two main types of male genitalia (Figs. 22 and 23). However, they all share similar internal female internal genitalia (Figs. 31–34, all considered Type F). It also appears that the mesepisternum reaches the mesocoxal cavity for species in this group. This group contains the species of Limonius traditionally considered agricultural pests and requires a tremendous amount of study to understand the diversity. Additionally, L. mandibularis is returned to the species list in this group, after it has been absent from most Nearctic catalogs for nearly 100 years (e.g. Woodworth 1913, Leng 1920).

The examined internal female internal genitalia of this group show some variation, but all share common characteristics. The majority of North American species in the group are similar to Fig. 33 (this is the genitalia type depicted in Becker 1956, for his third group). This type has a large sclerotized area in the bursa copulatrix near the junction of the bursa copulatrix and the uterus. The bursa copulatrix also has a medial offshoot bearing two accessory glands and a terminal tubular extension, shared by all variations (Figs. 31–34). This type is heavily modified in L. infuscatus (Fig. 34) where the accessory glands appear to be expanded into a large sac. The genitalia of L. aeruginosus (Fig. 31) has a small sclerotized area at the base of the bursa copulatrix near the junction with the uterus. The genitalia of L. longicornis (formerly in Neoathousius ) (Fig. 32) are very similar to that of L. aeruginosus , but apparently lack any sclerotized sections of the bursa copulatrix. It should be noted that the genitalia depicted in Fig. 32 are shrunken compared to the inflated genitalia in Fig. 31.

In many collections, species in Sylvanelater Johnson, 2008 (formerly part of Ctenicera ) are misidentified as Nothodes dubitans (= Limonius dubitans here). These specimens can be readily differentiated by their long pronotal hind angles and posteriorly widely open procoxal cavities (similar to Fig. 43). This confusion resulted in the overemphasis on the weak frontal margin in Limonius dubitans , which often led to this species being placed in other tribes, such as the Steatoderini (= modern Elaterini and Agriotini ) ( Schwarz 1906).

The synonymy of Neoathousius may seem controversial, but the relationship of these species to Limonius has been noted since its description ( Schimmel and Platia 1991, Schimmel and Platia 1994). There is no character that reliably separates these species at the generic level, and they are remarkably close to species from Northern Mexico and Texas. Additionally, as stated above, associated females share similar genitalia to Limonius species, confirming the synonymy. A species level revision is needed, as L. baconi and L. nigrinus are related to the species formerly in Neoathousius . The Palearctic Limonius baconi is also removed from the Nearctic species list, first appearing there without comment in Schwarz 1907. With the synonymy of Neoathousius with Limonius , Neoathousius ferrugineus Schimmel and Platia becomes a secondary homynym of Limonius ferrugineus Buysson (= L. pilosus ). I have renamed this species Limonius schimmeli in honor of one of the original describers, the late Rainier Schimmel.

The species L. rectangularis from west Texas is similar to the two Mexican species L. longulus and L. quadraticollis . This species is known from a single male holotype which I have recently (2017) examined (MCZC), and it was undissected. The genitalia figured in Al Dhafer (2009, Fig. 56) belongs to a different species, as the photograph is identical to that of a specimen of L. auripilis also at the MCZC. The actual genitalia of L. rectangularis remain unknown. As stated above, these three species are quite close to Palearctic species formerly in Neoathousius , and there are likely more species to be discovered in Northern Mexico.

Larvae have been associated with many species in the Canus species-group from all regions ( Lanchester 1946, Ôhira 1962, Dolin 1978, Etzler et al. 2014). Species in this group are considered to be economically important in the Nearctic, particularly the Pacific Northwest ( Lanchester 1946, Milosavljević et al. 2017, among many others). The larvae can be diagnosed by the frontal clypeal area with a tridentate nasale (Figs. 62, 71) and lacking setemata on the head (Fig. 69). The ninth abdominal segment is also characteristic (e. g. Fig. 72), with all species possessing urogomphi forming a caudal notch. The dorsal tubercles (Fig. 72, arrow) are helpful for species-level diagnoses ( Lanchester 1946).