Phylogenetic placement of the enigmatic longhorned beetle Vesperoctenus flohri Bates (Vesperidae) and a first description of its female internal structures
expand article infoStephanie Haddad, Nayeli Gutiérrez§, Felipe A. Noguera |, Seunggwan Shin, Petr Svacha#, Duane D. McKenna
‡ University of Memphis, Memphis, United States of America
§ American Museum of Natural History, New York, United States of America
| Universidad Nacional Autónoma de México, Jalisco, Mexico
¶ Seoul National University, Seoul, Republic of Korea
# Institute of Entomology, Czech Academy of Sciences, Ceske Budejovice, Czech Republic
Open Access


Taxonomic placement of the enigmatic monotypic Mexican longhorned beetle genus Vesperoctenus Bates is examined through inclusion in and reanalysis of the dataset of Haddad et al. (2018, Systematic Entomology 43: 68–89). We describe and discuss the phylogenetic significance of the internal structures of a recently collected V. flohri female from the Sierra de la Laguna mountain range in Mexico, the same specimen from which phylogenomic data was generated. Our phylogenomic analyses (469 genes) recovered Vesperoctenus with maximal statistical support within the cerambyciform family Vesperidae, sister to Vesperus Dejean (Vesperinae). Vesperus + Vesperoctenus were recovered sister to Philinae, and collectively form a clade sister to Anoplodermatinae. Thus, we place V. flohri within Vesperidae: Vesperinae: Vesperoctenini based on analyses of large-scale phylogenomic data. Finally, we propose that the conservation status of V. flohri merits assessment.

Key words

Cerambycidae, distribution, Mexico, morphology, taxonomy, Vesperinae, Vesperoctenini

1. Introduction

The taxonomic placement of the longhorned beetle Vesperoctenus flohri Bates has been highly controversial since its initial description by Bates in 1891. It is the only species within the genus Vesperoctenus Bates, is seldom collected, and is only known from Mexico (Baja California Sur, Durango, and Nuevo Leon; Vives 2001). The adults are predominantly nocturnal, females are brachypterous, and the larvae, although undescribed, are presumed to be subterranean root feeders (Bates 1891; Vives 2001). The specimens of the type series collected in Durango were reported in Bates (1891) to have been emerging from the ground and attracted to the light of a fire (the type series contained both sexes and thus the females, although flightless, may also be attracted by light). Besides further specimens attracted to UV light, at least one other was obtained by beating branches of Quercus devia Goldman in Baja California (Hovore 1988).

Svacha and Lawrence (2014) reviewed the various taxonomic placements of V. flohri since its formal description (summarized and updated in Table 1). Recently, Vesperoctenus has been classified either within Cerambycidae: Prioninae following Bousquet et al. 2009 (Bouchard et al. 2011; Tavakilian and Chevillotte 2020; Nearns et al. 2020; Monné 2020) or within Vesperidae (Svacha and Lawrence 2014) as a genus and tribe of uncertain subfamilial placement. Vesperoctenus has not been included in any molecular phylogenetic studies of Coleoptera to date and all attempts to determine its taxonomic affinities have relied on the incompletely known adult morphology (in particular, the female internal structures remain undescribed). We are not aware of any characters supporting the placement of Vesperoctenus in Prioninae. The antennae of both sexes are 12-segmented, unlike any other species in Vesperidae. Further, determining the exact taxonomic placement of V. flohri within Vesperidae has been compounded by its adults sharing individual morphological similarities with each of the three vesperid subfamilies – Anoplodermatinae, Philinae, and Vesperinae – making it difficult to conclusively place the species in one of them (see Svacha and Lawrence 2014 for details). Notable examples include the head of both Vesperus (Vesperinae) and Vesperoctenus constricted posteriorly to form a neck (unlike in Anoplodermatinae and Philinae and in a manner different from the “neck” in Lepturinae), and most Anoplodermatinae and Vesperoctenus sharing a 2-2-1 formula of terminal tibial spurs unlike Vesperinae (2-2-2) and Philinae (2-2-2 or 1-2-2); the 2-2-1 formula is also rare in the Cerambycidae. Saito (1990) described a simplified internal female genital tract in Vesperus (Vesperinae), Philus and Mantitheus Fairmaire (Philinae) with an (almost) completely desclerotized, simple, pouch-like spermatheca lacking a spermathecal gland, a situation apparently not known in any other cerambyciform and very probably apomorphic. A similar pouch-like membranous spermatheca has since been described in the philine genus Heterophilus (Bi and Lin 2015) where it bears a short spermathecal gland, thus confirming Saito’s interpretation of the membranous pouch as a desclerotized spermatheca. Vesperinae and Philinae were also recovered as sister groups in Haddad et al. (2018). However, females of Vesperoctenus are particularly rare in collections, have never been dissected, and their internal genitalia remain undescribed. Moreover, Vesperoctenus is geographically separated from other Vesperidae – it is the only extant Vesperidae known from North/Central America whereas the Anoplodermatinae are Neotropical, and Vesperus and Philinae occur exclusively in the Old World.

Table 1.

Taxonomic placements of Vesperoctenus flohri in major publications since its formal description.

Taxonomic Placement Reference
A longhorned beetle “closely allied” to Vesperus Dejean, the name Vesperoctenus means “related to Vesperus but with comb-like antennae” (of male). Bates (1891) (original description)
Placed in “Rhipiceridae” near Callirhipis Latreille (now Callirhipidae of Byrrhoidea). Horn (1894)
Reconfirmed in the broad Cerambycidae of that time; Horn’s (1894) arguments rejected. Gahan (1895)
In Cerambycidae: Cerambycinae: Lepturini (in Aurivillius next to Vesperus and Paraphilus Gahan = Mantitheus Fairmaire, now in Philinae). Aurivillius (1912)

Blackwelder (1946)
In Cerambycidae: Lepturinae: Vesperini. Boppe (1921)
In Vesperinae placed (together with Disteniinae, Oxypeltinae, and Philinae) as a subfamily of Crowson’s “Disteniidae” (the family should have been named Vesperidae for priority reasons); This is the first modern exclusion of Vesperinae including Vesperoctenus from the family Cerambycidae. Crowson (1981) (he probably never mentioned Vesperoctenus in the book, but table 14 lists Vesperinae as Palaearctic and C. American)
In Cerambycidae: Lepturinae: Lepturini. Catalogs and checklists of Cerambycidae s. l. of the Western Hemisphere and Neotropics beginning with the original versions (Monné and Giesbert 1994; Monné 1995, 2006) up to the publication of Bousquet et al. (2009)
Incertae sedis within the newly defined Vesperidae (containing the Vesperinae and Philinae of Crowson, 1981, plus the newly added Anoplodermatinae). Svacha et al. (1997)
In Vesperidae (with the placement of Anoplodermatinae in that family questioned) and closely related to Vesperus. Vives (2001)
In Vesperidae in its own newly erected tribe Vesperoctenini. Vives (2005)
Vesperoctenini (but not Vesperus) moved without explanation to Cerambycidae: Prioninae. Bousquet et al. (2009)
In Cerambycidae: Prioninae: Vesperoctenini (following Bousquet et al. 2009). Bouchard et al. (2011)
Catalogs and checklists of Cerambycidae s. l. of the Western Hemisphere and Neotropics following the publication of Bousquet et al. (2009), from versions dated 2011 till the present: Bezark (2020), Monné (2020)
Current versions of online cerambycid databases: Tavakilian and Chevillotte (2020), Nearns et al. (2020)
In Vesperidae: Vesperoctenini as a genus and tribe of uncertain subfamilial position; placement in Prioninae rejected. Svacha and Lawrence (2014)

Recently, phylogenomic studies have greatly improved our understanding of Coleoptera systematics and evolution (e.g., Haddad et al. 2018; Shin et al. 2018; McKenna et al. 2019; Vasilikopoulos et al. 2019). Studies of cerambyciform systematics and evolution have also been gaining momentum (e.g., Haddad and McKenna 2016; Haddad et al. 2018; Lee and Lee 2020; Souza et al. 2020a, b; Nie et al. 2021). In this study, we sought to determine the phylogenetic placement of Vesperoctenus using phylogenomic data, since morphology alone has failed to provide a clear answer to this. We specifically chose to do this within the context of the Haddad et al. (2018) data set since it includes representatives of all cerambyciform families (Cerambycidae sensu stricto, Disteniidae, Oxypeltidae, and Vesperidae) and subfamilies, along with the families Megalopodidae and Orsodacnidae. A female specimen of V. flohri was collected from the Sierra de la Laguna mountain range in Baja California Sur, Mexico. This specimen was used for obtaining phylogenomic data and for morphological study, including the first description of the female genitalia. Our molecular results conclusively establish the placement of this enigmatic genus within the higher-level phylogeny of cerambyciforms. We also argue that the conservation status of V. flohri requires consideration given our poor knowledge of its biology, distribution, genetic differentiation across its populations, and the reality of persistent threats to its habitat.

2. Methods

2.1. Specimen collection

N. Gutiérrez and P. Svacha collected a female specimen of V. flohri in Mexico, Baja California Sur, Sierra de la Laguna (23°32’60”N, 109°58’53”W, 1743 m) on 22 May 2018 at about 9:30 AM (Fig. 1). Vegetation in the area consists of natural grasslands and pine/oak forest, with a temperate subhumid climate and average annual temperatures of 14.7°C (CONANP 2003a). The female was preserved in clean ethyl alcohol and used for both molecular and morphological study. An extensive search for larvae on roots of various trees and herbs at this and another locality was unsuccessful.

Figure 1. 

Collecting site of Vesperoctenus flohri female in Mexico, Baja California Sur, Sierra de la Laguna (margin of pine-oak forest at the dried lake with two small trees of Arbutus in the foreground). Inset: Live female.

2.2. Geographical distribution of Vesperoctenus flohri

Documented localities of V. flohri were obtained from online databases, museum collections, and the literature. This information was stored in an Excel spreadsheet and then exported to the R statistical environment (R Core Team 2019). Once in R, data management was performed using dplyr package tools (Wickham et al. 2020), while the map was produced using the packages sf, ggplot2, gridExtra, OpenStreetMap, ggsn, and patchwork (Wickham 2016; Baptiste 2017; Pebesma 2018; Fellows 2019; Pedersen 2019; Santos-Baquero 2019; Ooms 2020).

2.3. Dissection of female internal structures

The legs, prothorax and right part of pterothorax were preserved for DNA and the remaining body was treated in hot 5% KOH to dissolve soft tissues. The cleaned cuticle was opened by microscissors to expose the internal structures – ovipositor, internal genitalia, metathoracic furca (metendosternite), and tentorium.

2.4. DNA extraction, library preparation, enrichment, and sequencing

We used anchored hybrid enrichment (AHE; Lemmon et al. 2012) to generate phylogenomic data for this study. Our previous studies of beetle phylogeny using AHE have produced well-resolved phylogenies with overall strong statistical support for relationships (Haddad et al. 2018; Shin et al. 2018; Johnson et al. 2018). DNA extraction, library preparation, enrichment and sequencing were performed only for the Vesperoctenus specimen collected for this study and following the methods outlined in Haddad et al. (2018) and Shin et al. (2018) with a few minor changes: an Arbor Biosciences myBaits target capture kit was used for hybridization-based capture and enrichment. Genomic data was extracted from a single leg of the specimen. Sequencing was performed by Novogene using a HiSeq X Ten (Illumina) sequencer and PE150 reads.

2.5. Read processing, assembly, and orthology assessment

Anchored hybrid enrichment data for Vesperoctenus was trimmed with Trimmomatic v 0.36 (Bolger et al. 2014) and then assembled using the program SOAPdenovo2 (Luo et al. 2012). Orthologous genes were identified using Orthograph (Petersen et al. 2017) exactly as in Haddad et al. (2018). Official gene sets (OGS) for three reference taxa in OrthoDB v7 (Waterhouse et al. 2011, 2013) were used: Danaus plexippus Linnaeus (Lepidoptera: Danaidae) (Zhan et al. 2011), Nasonia vitripennis Walker (Hymenoptera: Pteromalidae) (Werren et al. 2010) and Tribolium castaneum (Coleoptera: Tenebrionidae) (Richards et al. 2008). Following the methods in Haddad et al. (2018), 522 COGs (clusters of orthologous groups) were identified based on this pipeline before the filtering stage. We repeated this Orthograph pipeline for all data used in this study. Default parameters were retained in Orthograph throughout this process.

2.6. Multiple sequence alignment (MSA)

A script from Orthograph (Petersen et al. 2017) was used to summarize the fasta files generated from the orthology prediction pipeline for each orthologous gene, and the files were subsequently saved in a folder organized by orthologous gene ID. MSAs were then generated for each orthologous gene using the L-INS-i algorithm in MAFFT v7.130b (Katoh and Standley 2013). FASconCAT v1.0 (Kück and Meusemann 2010) was then used to generate an aligned supermatrix (781,854 NT sites) from the MSAs for the NT data. The resulting supermatrix was partitioned by gene.

2.7. Partitioning and modeling

PartitionFinder v. 2 (Lanfear et al. 2012) identified the best-fit cluster of partitions and models for the nucleotide data (using the rcluster option; Lanfear et al. 2014). The corrected Akaike information criterion (AICc; Hurvich and Tsai 1989) in PartitionFinder was used to determine partition clusters using the following parameters: model_selection = AICC; models = GTR+I+G, GTR+G; branch lengths = linked; search = rcluster, and the options: rate = 1.0, base = 1.0, model = 0.0, alpha = 1.0 with rcluster-percent = 10.0, using the command line script with the –raxml option. This resulted in 274 total partitions (262 with model GTR+I+G and 12 with model GTR+G, so the former model was used for the RAxML run).

2.8. Phylogenetic analyses

We conducted a partitioned maximum likelihood (ML) analysis (1000 rapid bootstrap replicates) for NT data (781,854 bp) in RAxML 8.2.12 (Stamatakis 2014). The analysis was performed on the HPC (high performance computing) cluster at the University of Memphis. Results from the bootstrap analyses were mapped onto the resulting ML tree. We consider nodes with a ML bootstrap support (MLBS) value greater or equal to 95% to be strongly supported.

3. Results

3.1. Phylogenetic placement of Vesperoctenus flohri

Excluding the placement of Vesperoctenus which we here report for the first time, the recovered relationships and clades in the partitioned maximum likelihood phylogeny for nucleotide data (Fig. 2) are identical to those of Haddad et al. (2018; their fig. 2), with only minor differences in nodal support; also the original cerambycid subfamilial classification of Haddad et al. (2018) is retained (additional support has been in the meantime obtained for placing Necydalinae and Parandrinae as ingroups of Lepturinae and Prioninae, respectively, e.g. in Nie et al. 2021). Vesperoctenus flohri, the focus of our study, is recovered within the cerambyciform family Vesperidae, sister to Vesperus (Vesperinae), with maximal nodal support. Vesperus + Vesperoctenus are recovered sister to the Philinae (represented by Philus pallescens Bates), and these collectively form a clade sister to Anoplodermatinae (represented by Migdolus fryanus Westwood and Mysteria darwini Lameere), all with maximal support. The monophyly of Vesperidae is likewise maximally supported.

Figure 2. 

Partitioned maximum likelihood phylogeny for nucleotide data. Maximum likelihood bootstrap support (MLBS) is shown only for nodes with MLBS ≤ 95%. Information regarding the systematics of the sampled exemplars is indicated on the right of the tree. Photograph of male Vesperoctenus flohri from Zacatecas (Sierra de Organos) by Juan Carrillo Contreras.

3.2. Female morphology of Vesperoctenus flohri

For terminology, see Svacha and Lawrence (2014). Vesperoctenus flohri was redescribed by Vives (2001) and the male also by Svacha and Lawrence (2014). Since Vives redescribed the female based on two old dry specimens, we include photographs of our fresh specimen (Fig. 3), the cleared head with open mandibles (Fig. 4a), the cleared pterothorax in lateral view (Fig. 4b), and the right wing (Fig. 4c). As in the male, the anterior margin of the sclerotized postclypeus slightly projects laterally above the membranous anteclypeus (Fig. 4a, arrow), but the paired projections are smaller and less prominent. Fig. 4b shows the unusually broad triangular metanepisternum (broadest of all studied vesperids). Vesperoctenus flohri females are brachypterous, but the wing of our specimen (Fig. 4c) is relatively long (its length comprises 90 % of the elytron length), has relatively complete venation with a minute rudiment of the wedge cell, and is more fully developed than that of the female lectotype from Durango depicted in Vives (2001: Fig. 4).

Figure 3. 

Vesperoctenus flohri, female: dorsal, lateral and ventral view (length 21.5 mm).

Figure 4. 

Vesperoctenus flohri, female: (a) Head, dorsal view (cleared specimen); (b) pterothorax, left lateral view (cleared specimen); (c) right wing. Lettering: mep – metanepisternum; wc – rudimentary wedge cell; arrow – lateral postclypeal projection.

The tentorium agrees with other Vesperidae (Svacha and Lawrence 2014) in having a broad roof-like tentorial bridge. The metatentorial arms bear long, flat and distally broadened dorsal arms. An extremely narrow connection exists between the pre- and metatentorial arms (it was torn during dissection).

Unlike the male hind coxae which are close together (Fig. 5a), those of the brachypterous flightless female are broadly separate (Figs 3, 6a) and the metendosternite lacks the common basal stalk and both arms abruptly diverge from their origins and run laterodorsally close to the thoracic walls, leaving the central metathoracic space free. The left and right laminae and anterior tendons thus become broadly separate (Fig. 6b).

Figure 5. 

(a) Vesperoctenus flohri, male, hind coxae, ventral view (the dense ventral pilosity was largely removed); (b) Vesperus strepens, male (cleared specimen), dtto; (c) V. strepens, male (cleared specimen), metendosternite, dorsal view. Lettering: arm – metendosternal arm; at – anterior tendon; lam – metendosternal lamina; st – basal stem.

Figure 6. 

(all cleared specimens). (a) Vesperoctenus flohri, female, hind coxae, ventral view; (b) V. flohri, female, metendosternite, dorsal and slightly anterior view (tip of right arm missing); (c) Vesperus strepens, female, hind coxae, ventral view; (d) V. strepens, female, metendosternite, dorsal view; (e) Mantitheus pekinensis, female, hind coxae, ventral view; (f) M. pekinensis, female, metendosternite, dorsal view (tip of left arm missing). Lettering: arm – metendosternal arm; at – anterior tendon; lam – metendosternal lamina; st – basal stem (missing in females of Vesperoctenus and Vesperus).

Female terminalia and internal genitalia. Sternum VIII has a V-shaped sclerotization and long spiculum gastrale (Fig. 7c). The ovipositor (Figs 7a, b) is long, flexible, and without any modifications for digging. The proctiger (the dorsal part between anus and ovipositor base in Fig. 7b) is extremely long, exceeding half of the ovipositor length and bearing two pairs of baculi; the medial pair is long and reaches the ovipositor base while the lateral pair is short and its ends approach the proximal ends of the paraproctal baculi. The valvifer is not distinguished from the coxite, the coxite lobes are long and slender and bear terminal gonostyli. The dorsal baculi are distinctly shorter than the medial pair of proctigeral baculi. The base of the membrane ensheathing the ovipositor is produced into small “intersegmental pouches” (see Discussion). The vagina forms a simple membranous tube, broadened bursa copulatrix is small, without vaginal plates at base, spermathecal duct arising apically (not sidewise as in Anoplodermatinae and most Cerambycidae), short and thin, terminated by a finely sclerotized C-shaped spermatheca with a spermathecal gland at its base (Fig. 7d).

Figure 7. 

Vesperoctenus flohri, female genitalia and sternum VIII. (a) Ovipositor and internal genital tract, ventral view; (b) ovipositor, dorsal view; (c) abdominal sternum VIII and membranous tube ensheathing the ovipositor, ventral view; (d) detail of internal genital tract. Lettering: a – anus; bc – bursa copulatrix; cb – coxital baculum; cl – coxite lobe; co – common oviduct; db – dorsal baculum; gpo – gonopore; hg – hindgut; ip – intersegmental pouches; lo – lateral oviduct; lptb – lateral proctigeral baculum; m – membranous tube ensheathing ovipositor; mptb – medial proctigeral baculum; ppb – paraproctal baculum; sp – spermatheca; spd – spermathecal duct; spgl – spermathecal gland; sty – stylus; sv – spiculum ventrale (apodeme of sVIII); sVIII – sternum VIII; vg – vagina.

4. Discussion

4.1. Molecular phylogeny and systematic placement

The lack of V. flohri larvae and molecular data (until now), the scarcity of available adult specimens, and the disjunct distribution of vesperids (not many Old World taxonomists had specimens available for comparison) has rendered the evaluation of the relationships and systematic placement of the genus Vesperoctenus difficult and variable (see Introduction and Table 1). Our molecular phylogenetic analysis reconfirms the monophyly of the family Vesperidae in the sense of Svacha et al. (1997) and Haddad et al. (2018) and firmly recovers Vesperoctenus within Vesperidae as the sister taxon of Vesperus. We therefore place V. flohri within Vesperidae: Vesperinae: Vesperoctenini as sister to the Vesperini containing a single genus Vesperus with about 20 species (status of some species-level taxa is not fully agreed upon). This phylogenetic placement is interesting, particularly from a biogeographic point of view. Vesperoctenus is endemic to Mexico, whereas the monogeneric Vesperini is strictly Mediterranean. Moreover, the Philinae (5 genera, ~19 spp.), which is sister to Vesperinae, is similarly an Old World taxon occurring predominantly in the Oriental Region (one species of Doesus reaches subsaharan Africa). Only the Anoplodermatinae is Neotropical in distribution (Svacha and Lawrence 2014). Thus, V. flohri is the only extant vesperid in North/Central America and the only non-anoplodermatine vesperid in the New World. Crowson (1981: table 14 on p. 626) listed Vesperinae (occurring in the Mediterranean region of the Palearctic and in Mexico) among taxa with a relict distribution.

4.2. Morphology

Refer to Svacha and Lawrence (2014) for morphological summaries of individual vesperid subfamilies and for some figures referenced herein. When interpreting morphological characters in light of our obtained molecular phylogeny (monophyletic Philinae + (Vesperus + Vesperoctenus) as a sister group to the Anoplodermatinae), we find that the similarities of the “non-anoplodermatine” vesperid clade are either difficult to polarize or probable symplesiomorphies.

Vesperoctenus (as well as Vesperus and the Philinae) does not share most of the probable synapomorphies of the Neotropical Anoplodermatinae: procoxal cavities closed externally (open in other Vesperidae); wing invariably without a wedge cell (wedge cell present in macropterous specimens of Philinae, Vesperoctenus and some Vesperus) and with MP3+4 attached to MP1+2 at about half length of the latter (at about one-third length or less in all other Vesperidae and in most other non-chrysomelid chrysomeloids); metendosternite in dissected species (both sexes of Pathocerus and Migdolus, male of Hypocephalus) without laminae (with laminae in males and most females of other dissected Vesperidae including the modified metendosternites of some flightless females, although strongly reduced in females of some species of Vesperus, see below and Fig. 6); short sclerotized “digging” ovipositor (long flexible non-digging ovipositor with terminal gonostyli in all other Vesperidae, Figs 7a, b and Saito 1990, although the ovipositor is slightly shorter and more robust in females of Heterophilus whose oviposition mode is unknown: Bi and Lin 2015).

Svacha (in Svacha et al. 1997) established the family Vesperidae in the present extent and included Vesperoctenus as a genus incertae sedis, pointing out some potential synapomorphies with the Neotropical Anoplodermatinae. In light of the present results, the shared and undoubtedly apomorphic 2-2-1 pattern of tibial spurs in Vesperoctenus and the anoplodermatine groundplan, and the extreme hairiness of males of Vesperoctenus and of some genera of Anoplodermatinae (Sypilus Guérin-Méneville, Cherrocrius Berg) should be considered parallelisms. The mandibles of both sexes of Vesperoctenus with asymmetrically dentate incisor edges and an outer tubercle (present Fig. 4a for female, fig. 2.1.4 C in Svacha and Lawrence 2014 for male) resemble some Anoplodermatinae and differ from Vesperus and Philinae with more or less simple incisor edges and no distinct outer tubercle, but polarity of these characters is uncertain.

No reliable synapomorphies could be found for the Philinae + Vesperinae clade because some adults of Philinae are morphologically very generalized cerambyciforms or even chrysomeloids and all similarities are probably plesiomorphic or at least difficult to interpret. The interpretation of larval characters is also difficult (Svacha and Lawrence 2014: 35), particularly in light of the molecular results (Haddad et al. 2018), and since the larvae of Vesperoctenus are unknown. Saito (1990) described the female ovipositor and internal genital tract in one Vesperus and two Philinae (Philus and Mantitheus) and found them unique and very probably apomorphic in several characters: the ovipositor has an extremely long proctiger with two pairs of proctigeral baculi instead of one, the bursa copulatrix is small or absent and the spermathecal duct arises apically instead of sidewise as in many other cerambyciforms, the vaginal plates are virtually absent, and the spermatheca is very lightly sclerotized in Vesperus strepens (Fabricius) (“fairly stiff”: Saito 1990: 250; confirmed by us on another dissected specimen) or membranous in the two philines (and also in Vesperus luridus Rossi dissected by us). In all those taxa, the spermatheca lacks a spermathecal gland. The ovipositor of Vesperoctenus (Figs 7a, b) is very similar to that of Vesperus and the two Philinae in Saito (1990), including the extremely long proctiger bearing two pairs of baculi, but the short sclerotized “digging” ovipositor of the Anoplodermatinae (Svacha and Lawrence 2014: fig. 2.1.5 K) is so derived that it would be difficult to make speculations regarding ovipositor morphology in a vesperid ancestor and claim the ovipositor synapomorphic for Vesperinae + Philinae. The intersegmental pouches (Figs 7a, b) also occur in Vesperus and Philinae and have no clear homologues in the derived anoplodermatine ovipositors (Svacha and Lawrence 2014), but they are relatively widespread in various cerambycoids and thus cannot be labelled as a synapomorphy of the non-anoplodermatine vesperid clade. In some Cerambycidae, they serve for storing yeast-like larval gut symbionts which are transferred onto the egg at oviposition. However, Schomann (1937) did not find any such symbionts in those pouches in Vesperus and their function in vesperids remains unknown. Also, the desclerotized spermatheca lacking a gland can no longer be considered a synapomorphy of the non-anoplodermatine clade. The spermatheca of another philine (Heterophilus), though also membranous, has a small spermathecal gland (Bi and Lin 2015), and Vesperoctenus possesses a relatively typical C-shaped moderately sclerotized spermatheca with a distinct gland (Fig. 7d). The reduced bursa copulatrix with missing vaginal plates and with the spermathecal duct arising apically is not entirely unique in the cerambyciforms, but it is worth noting that the only two dissected females of Anoplodermatinae (Pathocerus and Migdolus) have a large sac-like bursa copulatrix with the spermathecal duct arising sidewise near to its base (Svacha and Lawrence 2014).

Vesperoctenus, Vesperus, and known Philinae share the large thin flattened dorsal tentorial arms; in dissected Anoplodermatinae these arms are present but are short and tapering or rod-like, not broad and flattened. Large dorsal arms should be plesiomorphic and although the broad flat almost foliar dorsal ends are unusual, we cannot decide whether such dorsal arms were present in the vesperid ancestor and reduced in the Anoplodermatinae, or whether the broad flat dorsal ends may represent a synapomorphy of the Philinae and Vesperinae.

It would be difficult to list any synapomorphies of the Philinae because of the above mentioned generalized plesiomorphic morphology of some philine taxa. However, Philinae were represented by three relatively different genera (Philus, Spiniphilus Lin & Bi and Mantitheus, the latter with brachelytrous micropterous females resembling females of some Vesperus) in the mitochondrial phylogeny of Nie et al. (2021) and formed a strongly supported monophyletic clade sister to Vesperus. Potential synapomorphies of the Vesperinae (Vesperoctenus and Vesperus) include several characters:

1. Head with large inflated occipital region and then abruptly constricted into a narrow neck not including posterior gular region and metatentorial slits. This head type (similar to, but in details, different from that of some lepturine or lepturine-like Cerambycidae in which the neck does include metatentorial slits) is virtually unknown in other groups of the non-chrysomelid Chrysomeloidea.

2. Modified female metendosternite. Unlike the metendosternites in the flying males of the Anoplodermatinae which have the common basal stem but lack the laminae, those of the Philinae + Vesperinae clade are “hylecoetoid” (sensu Crowson 1938) in males (all are capable of flight), with a long basal stem and large laminae. Fig. 5c shows the male metendosternite of Vesperus strepens, that of male Vesperoctenus could not be studied in detail but should be similar because the males share the closely adjacent hind coxae (Fig. 5a) and the laminae are preserved even in the modified female metendosternite (see below). This hylecoetoid type also occurs in macropterous females (Philus dissected), though the basal stem may be shorter, and is plesiomorphic beyond doubt. Metendosternites of the brachy- or micropterous and (when bearing full egg load) often more or less physogastric females of some genera (Vesperoctenus and Vesperus of Vesperinae and Mantitheus and Heterophilus of Philinae) have apparently never been studied. Svacha and Lawrence (2014: 22) noted that the hind coxae are “widely separated in some flightless females” and depicted that character in Vesperus (their fig. 2.1.3 A), but none of these females were dissected. Females of Vesperus (V. strepens, V. luridus and V. conicicollis Fairmaire and Coquerel dissected) and Vesperoctenus have very broadly separated hind coxae and their metendosternite is strongly different from that of the males: The basal stem is completely missing and the arms abruptly diverge from the very base, making the anterior tendons broadly separate; the laminae may be preserved (they are strongly reduced in Vesperus luridus and V. conicicollis), but likewise widely apart (Figs 6b, d). However, this modification is obviously strongly adaptive and may develop parallelly. While in the female of Mantitheus a short basal stem is retained (Fig. 6f), in Heterophilus it is extremely reduced, approaching the situation in Vesperus strepens (Meiying Lin, pers. comm.). All females of Anoplodermatinae are flightless and the dissected females of Pathocerus and Migdolus also lack the basal stem and the arms abruptly diverge from the very base (but they lack the laminae like the males). This modification opens the space between abdomen and metathorax and enables protrusion of the egg mass into the posterior part of the metathoracic cavity (confirmed in the dissected female of Mantitheus which was collected from its soil pupal chamber with full egg load).

3. Wing, even in macropterous specimens, with at most, a narrow reduced wedge cell (missing in some Vesperus); that cell is always large in macropterous Philinae (figures in Lin and Bi 2011, or Svacha and Lawrence 2014). The invariable absence of that cell in all Anoplodermatinae should be a parallel reduction independent from that in some species of Vesperus.

4. Vesperus and Vesperoctenus share the prominent procoxae and reduced narrow intercoxal process. In Philinae the intercoxal process is high and broad and the procoxae, although relatively prominent, are inserted under the process and articulating on it, occasionally by a distinct tubercle (Svacha and Lawrence 2014: fig. 2.1.5 H). With the exception of the extremely derived Hypocephalus, the situation is virtually identical in the Anoplodermatinae including the occasional presence of the articulating tubercle (e.g., in Anoploderma). Thus, the free prominent coxae of Vesperus and Vesperoctenus could be synapomorphic.

Autapomorphies of Vesperoctenus include: the 12-segmented antennae (the terminal flagellomere may be appendiculate but never completely divided in the other Vesperidae), possibly the very broad (high) metanepisternum (Fig. 4b, narrower in all other studied vesperids), and the extremely long and dense male pilosity if we consider the similarly hairy males in a few Anoplodermatinae a parallel development (see above). The more or less projecting lateral postclypeus should also be autapomorphic because various postclypeal modifications in some Anoplodermatinae are different in details.

No reliable synapomorphies could be found for the sister genus Vesperus. Females often have shortened elytra, but in some species, they are about as well developed as in females of Vesperoctenus and in Vesperus macropterus Sama even the female wings are fully developed and can be used (together with the well-developed elytra) for parachuting from the tall dry herbs whose inflorescences the females oviposit in (Sechi 2011). The desclerotized spermatheca lacking the spermathecal gland (which can no longer be considered synapomorphic for the Philinae + Vesperinae clade) would be a synapomorphy of Vesperus if its presence is universal within the genus, but so far it can be confirmed for only two species (V. strepens and V. luridus).

Other characters occasionally employed in the classification of this group are not helpful in evaluating the relationships and taxonomic position of Vesperoctenus. The lateral pronotal margin or carina is present in the Anoplodermatinae and virtually absent in Vesperinae, but it may be present or absent with various transitions in Philinae. The most complete pronotal margin occurs in the flightless females of Heterophilus (Lin and Bi 2011; Bi and Lin 2015) which are almost certainly far from the philine groundplan. The mesoscutal stridulatory file is absent in Anoplodermatinae and Vesperoctenus, but present to absent in Philinae, and distinct rudiments were also found in a species of Vesperus (Svacha and Lawrence 2014: fig. 2.1.4 G).

4.3. Distribution and conservation status

Vesperoctenus flohri has rarely been collected and historically appears to have a very narrow geographic distribution which raises concerns about its conservation status. Few specimens of V. flohri are available in museum collections and only a handful of collection records exist in the published literature which have so far limited the distribution of this species to only three Mexican states (Baja California Sur, Durango, and Nuevo Leon). Interestingly, we found a few online observations of the species (on Facebook and iNaturalist) that expand its known geographic distribution to include the Mexican states of Tlaxcala, Veracruz, and Zacatecas (Fig. 8). The new state records in Veracruz and Tlaxcala confirm the presence of V. flohri in the Trans-Mexican Volcanic Belt, whereas the ones in Zacatecas might be a possible connection between previous records of the species in Nuevo Leon and Durango. These new records also expand the known habitat of V. flohri which is oak/pine forests and grasslands in northern Mexico. The locality in Veracruz (at 2520 m) is the highest altitude where Vesperoctenus has been recorded (considering only those records with altitude data), is characterized by dry desert scrub with Opuntia cacti and Yucca, and its annual temperature varies between 6°C and 11°C (Santiago Jaume-Schinkel, pers. comm.; Lara-Lagunes et al. 2017). On the other hand, vegetation in Nanacamilpa, Tlaxcala is composed of coniferous and latifoliate forest, with subhumid climate and average annual temperature between 12°C and 18°C (INEGI 2009). Finally, V. flohri recorded in Sierra de Organos, Zacatecas were attracted to light from a cabin surrounded by grassland and Pinus-Quercus forests, with a temperate subhumid climate and an average annual temperature between 12°C and 18°C (Juan Carrillo, pers. comm.; CONANP 2003b).

Figure 8. 

Documented localities of Vesperoctenus flohri based on data from online databases, museum collections, and the literature. Tlaxcala, Veracruz, and Zacatecas are new state records; they originate from observations of V. flohri available online.

Sixty percent of Vesperoctenus records are located in or near national and state protected natural areas with high biological diversity (CONANP 2020). These areas also host countless other endemic species and have historically been under critical threat of mismanagement due to various political and socioeconomic factors (Valdez et al. 2006). For example, the Sierra de la Laguna (the collection site of this study’s V. flohri specimen) is an isolated mountain range at the southern end of the Mexican State of Baja California Sur. It was declared a Biosphere Reserve by a Mexican presidential decree on June 6, 1994 (DOF 1994), and subsequently by UNESCO (United Nations Educational, Scientific, and Cultural Organization) in 2003 (UNESCO 2011). This geographically isolated mountain complex is comprised of tropical dry deciduous forest and semi-desert ecosystems with high levels of diversity and endemism and is home to several endangered species and species new to science (e.g., Riemann and Ezcurra 2005; León de la Luz and Breceda 2006; Vandeplank et al. 2016). This protected natural area has recently been threatened by an open-pit gold mining project that would negatively impact habitat and water quality in addition to having numerous other harmful outcomes that would impact the reserve (Vandeplank et al. 2016). Mining could cause irrevocable detrimental effects to the diverse species inhabiting the mountain range, including V. flohri. Moreover, morphological differences such as that between the female wings of the specimens from Baja California Sur and Durango (see above) may indicate a substantial degree of genetic differentiation between populations. Thus, we argue that the conservation status of V. flohri necessitates assessment considering how little we know of its biology and ecology, our growing knowledge of its distribution and habitat, and given the persistent threats to known populations posed by human-induced habitat loss and degradation.

5. Acknowledgements

We would like to thank Drs. Paulina Cifuentes Ruiz and Santiago Zaragoza Caballero (UNAM) for their assistance with the logistics of our collecting trip in Sierra de la Laguna and support in obtaining the required permits. We are grateful to Martin Cota, Forest Keeper of the Sierra de la Laguna Biosphere Reserve, who kindly helped us during the collecting trip, and to Juan Carrillo and Santiago Jaume-Schinkel for information on the habitat where they found Vesperoctenus. We would also like to thank Alejandra Barrios Sifuentes, Lazaro Rangel Castaneda, and Jesús Quiñonez Gómez (CONANP, Sierra de La Laguna Biosphere Reserve) for approving and facilitating our stay on the reserve. Finally, we thank Meiying Lin (Institute of Zoology, Chinese Academy of Sciences, Beijing) for information on the female metendosternite of Heterophilus. This research was funded in part by NSF grant DEB-1355169 to DDM.

6. References

  • Aurivillius C (1912) Coleopterorum Catalogus. Pars 39. Cerambycidae: Cerambycinae. 574 pp. W. Junk, Berlin.
  • Bates HW (1891) New longicorn Coleoptera, chiefly from Mexico. Entomologist’s Monthly Magazine 27: 158–161.
  • Bi W, Lin M (2015) Discovery of second new species of the genus Spiniphilus Lin & Bi, and female of Heterophilus scabricollis Pu with its biological notes (Coleoptera: Vesperidae: Philinae: Philini). Zootaxa 3949(4): 575–583.
  • Blackwelder RE (1946) Checklist of the coleopterous insects of Mexico, Central America, the West Indies, and South America. Part 4, Cerambycidae. Bulletin of the United States National Museum 185: 551–627.
  • Boppe P (1921) Coleoptera Longicornia Fam. Cerambycidae Subfam. DisteniinaeLepturinae. In Wytsman, P. (Ed). Genera Insectorum Fascicule 178: 1–121, 8 plates. L. Desmet-Verteneuil, Brussels.
  • Bolger AM., Lohse M, Usadel B (2014) Trimmomatic: a flexible trimmer for Illumina sequence data. Bioinformatics 30: 2114–2120.
  • Bouchard P, Bousquet Y, Davies AE, Alonso-Zarazaga MA, Lawrence JF, Lyal CHC, Newton AF, Reid CAM, Schmitt M, Slipinski SA, Smith ABT (2011) Family group names in Coleoptera (Insecta). ZooKeys 88: 1–972.
  • Bousquet Y, Heffern DJ, Bouchard P, Nearns E (2009) Catalogue of family-group names in Cerambycidae (Coleoptera). Zootaxa 2321: 1–80.
  • CONANP (2003a) Programa de manejo Reserva de la Biosfera Sierra de la laguna. 212 pp.
  • CONANP (2003b) Programa de Manejo Parque Nacional Sierra de Órganos. 136 pp.
  • Crowson RA (1938) The metendosternite of the Coleoptera: A comparative study. Transactions of the Royal Entomological Society London 87: 397–415, pls. 1–13.
  • Crowson RA (1981) The Biology of the Coleoptera. Academic Press, London, 802 pp.
  • Diario Oficial (DOF) (1994) Decreto por el que se declara como Reserva de la Biosfera a La Sierra La Laguna. Pp. 52–58 en: Diario Oficial de la Federación, Estados Unidos Mexicanos, Presidencía de la República.
  • Gahan CJ (1895) Vesperoctenus, Bates, and its systematic position. Entomologist’s Monthly Magazine 31: 22–24.
  • Haddad S, McKenna DD (2016) Phylogeny and evolution of the superfamily Chrysomeloidea (Coleoptera: Cucujiformia). Systematic Entomology 41: 697–716.
  • Haddad S, Shin S, Lemmon AR, Lemmon EM, Svacha P, Farrell BD, Ślipiński A, Windsor D, McKenna DD (2018) Phylogenomics resolves the enigmatic higher-level phylogeny of longhorned beetles (Cerambycidae). Systematic Entomology 43: 68–89.
  • Horn GH (1894) The Coleoptera of Baja California. Proceedings of the California Academy of Sciences (Series 2) 4: 302–449, pls. VII and VIII.
  • Hovore FT (1988) Additions to the cerambycid beetle fauna of Baja California, Mexico: Records and descriptions (Coleoptera: Cerambycidae). Wasmann Journal of Biology 46(1–2): 1–29.
  • Hurvich CM, Tsai CL (1989) Regression and time series model selection in small samples. Biometrika 76: 297–307.
  • Johnson AJ, McKenna DD, Jordal BH, Cognato AI, Smith-Cognato SM, Lemmon AR, Lemmon EL, Hulcr J (2018) Phylogenomics clarifies repeated evolutionary origins of inbreeding and fungus farming in bark beetles (Curculionidae, Scolytinae). Molecular Phylogenetics and Evolution 127: 229–238.
  • Katoh K, Standley DM (2013) MAFFT Multiple sequence alignment software Version 7: Improvements in performance and usability. Molecular Biology and Evolution 30(4): 772–780.
  • Kück P, Meusemann K (2010) FASconCAT: Convenient handling of data matrices. Molecular Phylogenetics and Evolution 56(3): 1115–1118.
  • Lanfear R, Calcott B, Ho SY, Guindon S (2012) Partitionfinder: combined selection of partitioning schemes and substitution models for phylogenetic analyses. Molecular Biology and Evolution 29(6): 1695–1701.
  • Lanfear R, Calcott B, Kainer D, Mayer C, Stamatakis A (2014) Selecting optimal partitioning schemes for phylogenomic datasets. BMC Evolutionary Biology 14: 82.
  • Lara-Lagunes N, Jaume-Schinkel S, Ibáñez-Bernal S (2017) An incidental case of gastric pseudomyiasis in Canis latrans (Carnivora: Canidae) by a rabbit bot fly Cuterebra sp. (Diptera: Oestridae). Revista Mexicana de Biodiversidad 88(2): 410–414.
  • Lee S, Lee S (2020) Multigene phylogeny uncovers oviposition-related evolutionary history of Cerambycinae (Coleoptera: Cerambycidae). Molecular Phylogenetics and Evolution 106707.
  • Lemmon AR, Emme SA, Lemmon EM (2012) Anchored hybrid enrichment for massively high-throughput phylogenomics. Systematic Biology 61: 727–744.
  • León de la Luz JL, Breceda A (2006) Using endemic plant species to establish critical habitats in the Sierra La Laguna Biosphere Reserve, Baja California Sur, Mexico. Biodiversity and Conservation 15(3): 1043–1055.
  • Lin M-Y, Bi W-X (2011) A new genus and species of the subfamily Philinae (Coleoptera: Vesperidae). Zootaxa 2777: 54–60.
  • Luo R, Liu B, Xie Y, Li Z, Huang W, Yuan J, He G, Chen Y, Pan Q, Liu Y, Tang J, Wu G, Zhang H, Shi Y, Liu Y, Yu C, Wang B, Lu Y, Han C, Cheung DW, Yiu SM, Peng S, Xiaoqian Z, Liu G, Liao X, Li Y, Yang H, Wang J, Lam TW, Wang J (2012) SOAPdenovo2: an empirically improved memory-efficient short-read de novo assembler. Gigascience 1(1): 18.
  • McKenna DD, Shin S, Ahrens D, Balke M, Beza-Beza C, Clarke DJ, Donath A, Escalona HE, Friedrich F, Letsch H, Liu S, Maddison D, Mayer C, Misof B, Murin PJ, Niehuis O, Peters RS, Podsiadlowski L, Pohl H, Scully ED, Yan EV, Zhou X, Ślipiński A, Beutel RG (2019) The evolution and genomic basis of beetle diversity. Proceedings of the National Academy of Sciences 49: 24729–24737.
  • Monné MA (1995) Catalogue of the Cerambycidae (Coleoptera) of the Western Hemisphere. Part XXI. Subfamily Lepturinae. Sociedade Brasileira de Entomologia, São Paulo, 159 pp.
  • Monné MA (2006) Catalogue of the Cerambycidae (Coleoptera) of the Neotropical Region. Part III. Subfamilies Parandrinae, Prioninae, Anoplodermatinae, Aseminae, Spondylidinae, Lepturinae, Oxypeltinae, and addenda to the Cerambycinae and Lamiinae. Zootaxa 1212: 1–244.
  • Monné MA (2020) Catalogue of the Cerambycidae (Coleoptera) of the Neotropical Region. Part III. Subfamilies Lepturinae, Necydalinae, Parandrinae, Prioninae, Spondylidinae and Families Oxypeltidae, Vesperidae and Disteniidae. Available at:
  • Monné MA, Giesbert EF (1994) Checklist of the Cerambycidae and Disteniidae (Coleoptera) of the Western Hemisphere. Wolfsgarden Books, Burbank, CA, xiv+410 pp.
  • Nearns EH, Lord NP, Lingafelter SW, Santos-Silva A, Miller KB, Zaspel JM (2020) Longicorn ID: Tool for Diagnosing Cerambycoid Families, Subfamilies, and Tribes. The University of New Mexico, Purdue University, and USDA APHIS PPQ Identification Technology Program (ITP). Available from: (last accessed on December 17, 2020)
  • Nie R, Vogler AP, Yang X-K, Lin M (2021) Higher-level phylogeny of longhorn beetles (Coleoptera: Chrysomeloidea) inferred from mitochondrial genomes. Systematic Entomology 46: 56–70.
  • Petersen M, Meusemann K, Donath A, Dowling D, Liu S, Peters RS, Podsiadlowski L, Vasilikopoulos A, Zhou X, Misof B, Niehuis O (2017) Orthograph: a versatile tool for mapping coding nucleotide sequences to clusters of orthologous genes. BMC Bioinformatics 18: 111.
  • R Core Team (2019) R: A language and environment for statistical computing. R Foundation for Statistical Computing, Vienna, Austria.
  • Richards S, Gibbs RA, Weinstock GM, Brown SJ, Denell R, Beeman RW, Gibbs R, Beeman RW, Brown SJ, Bucher G, Friedrich M, Grimmelikhuijzen CJ, Klingler M, Lorenzen M, Richards S, Roth S, Schröder R, Tautz D, Zdobnov EM, Muzny D, Gibbs RA, Weinstock GM, Attaway T, Bell S, Buhay CJ, Chandrabose MN, Chavez D, Clerk-Blankenburg KP, Cree A, Dao M, Davis C, Chacko J, Dinh H, Dugan-Rocha S, Fowler G, Garner TT, Garnes J, Gnirke A, Hawes A, Hernandez J, Hines S, Holder M, Hume J, Jhangiani SN, Joshi V, Khan ZM, Jackson L, Kovar C, Kowis A, Lee S, Lewis LR, Margolis J, Morgan M, Nazareth LV, Nguyen N, Okwuonu G, Parker D, Richards S, Ruiz SJ, Santibanez J, Savard J, Scherer SE, Schneider B, Sodergren E, Tautz D, Vattahil S, Villasana D, White CS, Wright R, Park Y, Beeman RW, Lord J, Oppert B, Lorenzen M, Brown S, Wang L, Savard J, Tautz D, Richards S, Weinstock G, Gibbs RA, Liu Y, Worley K, Weinstock G, Elsik CG, Reese JT, Elhaik E, Landan G, Graur D, Arensburger P, Atkinson P, Beeman RW, Beidler J, Brown SJ, Demuth JP, Drury DW, Du YZ, Fujiwara H, Lorenzen M, Maselli V, Osanai M, Park Y, Robertson HM, Tu Z, Wang JJ, Wang S, Richards S, Song H, Zhang L, Sodergren E, Werner D, Stanke M, Morgenstern B, Solovyev V, Kosarev P, Brown G, Chen HC, Ermolaeva O, Hlavina W, Kapustin Y, Kiryutin B, Kitts P, Maglott D, Pruitt K, Sapojnikov V, Souvorov A, Mackey AJ, Waterhouse RM, Wyder S, Zdobnov EM, Zdobnov EM, Wyder S, Kriventseva EV, Kadowaki T, Bork P, Aranda M, Bao R, Beermann A, Berns N, Bolognesi R, Bonneton F, Bopp D, Brown SJ, Bucher G, Butts T, Chaumot A, Denell RE, Ferrier DE, Friedrich M, Gordon CM, Jindra M, Klingler M, Lan Q, Lattorff HM, Laudet V, von Levetsow C, Liu Z, Lutz R, Lynch JA, da Fonseca RN, Posnien N, Reuter R, Roth S, Savard J, Schinko JB, Schmitt C, Schoppmeier M, Schröder R, Shippy TD, Simonnet F, Marques-Souza H, Tautz D, Tomoyasu Y, Trauner J, Van der Zee M, Vervoort M, Wittkopp N, Wimmer EA, Yang X, Jones AK, Sattelle DB, Ebert PR, Nelson D, Scott JG, Beeman RW, Muthukrishnan S, Kramer KJ, Arakane Y, Beeman RW, Zhu Q, Hogenkamp D, Dixit R, Oppert B, Jiang H, Zou Z, Marshall J, Elpidina E, Vinokurov K, Oppert C, Zou Z, Evans J, Lu Z, Zhao P, Sumathipala N, Altincicek B, Vilcinskas A, Williams M, Hultmark D, Hetru C, Jiang H, Grimmelikhuijzen CJ, Hauser F, Cazzamali G, Williamson M, Park Y, Li B, Tanaka Y, Predel R, Neupert S, Schachtner J, Verleyen P, Raible F, Bork P, Friedrich M, Walden KK, Robertson HM, Angeli S, Forêt S, Bucher G, Schuetz S, Maleszka R, Wimmer EA, Beeman RW, Lorenzen M, Tomoyasu Y, Miller SC, Grossmann D, Bucher G (2008) The genome of the model beetle and pest Tribolium castaneum. Nature 452(7190): 949–955.
  • Riemann H, Ezcurra E (2005) Plant endemism and natural protected areas in the peninsula of Baja California, Mexico. Biological Conservation 122(1): 141–150.
  • Saito A (1990) Female reproductive organs of cerambycid beetles from Japan and the neighboring areas. I. Philini through Atimiini. Elytra (Tokyo) 18(2): 231–260.
  • Schomann H (1937) Die Symbiose der Bockkäfer. Zeitschrift für Morphologie und Ökologie der Tiere 32: 542–611.
  • Sechi D. (2012) Osservazioni sulla biologia di Vesperus macropterus (Coleoptera, Cerambycidae). Fragmenta Entomologica 43(1): 75–87.
  • Shin S, Clarke DJ, Lemmon AR, Moriarty Lemmon E, Aitken AL, Haddad S, Farrell BD, Marvaldi AE, Oberprieler RG, McKenna DD (2018) Phylogenomic data yield new and robust insights into the phylogeny and evolution of weevils. Molecular Biology and Evolution 35: 823–836.
  • Slater GS, Birney E (2005) Automated generation of heuristics for biological sequence comparison. BMC Bioinformatics 6: 31.
  • Souza DS, Marinoni L, Monné ML, Gómez-Zurita J (2020a) Molecular phylogenetic assessment of the tribal classification of Lamiinae (Coleoptera: Cerambycidae). Molecular Phylogenetics and Evolution 145: 106736.
  • Souza DS, Sepúlveda TA, Marinoni L, Monné ML (2020b) Phylogenetic analyses provide new insights into systematics of the longhorned beetle tribe Acrocinini (Coleoptera: Cerambycidae: Lamiinae). Arthropod Systematics and Phylogeny 78. 17–27.
  • Stamatakis A (2014) RAxML version 8: a tool for phylogenetic analysis and post-analysis of large phylogenies. Bioinformatics 30: 1312–1313.
  • Svacha P, Lawrence JF (2014) 2.1. Vesperidae Mulsant, 1839. In: Leschen RAB and Beutel RG (Eds) Handbook of Zoology, Band 4: Arthropoda: Insecta, Teilband/part 40 Coleoptera, Beetles, Vol. 3: Morphology and Systematics (Phytophaga). Walter de Gruyter, Berlin, 16–177.
  • Svacha P, Wang JJ, Chen SC (1997) Larval morphology and biology of Philus antennatus and Heterophilus punctulatus, and systematic position of the Philinae (Coleoptera: Cerambycidae and Vesperidae). Annales de la Société Entomologique de France 33: 323–369.
  • Tavakilian G, Chevillotte H (2020) TITAN: base de données internationales sur les Cerambycidae ou Longicornes. Available at: (updated December 10, 2020)
  • Valdez R, Guzmán-Aranda JC, Abarca FJ, Tarango-Arámbula LA, Sánchez FC (2006) Wildlife conservation and management in Mexico. Wildlife Society Bulletin 34(2): 270–282.
  • Vanderplank S, Wilder BT, Ezcurra E (2016) Botanical Research Institute of Texas, Next Generation Sonoran Desert Researchers, and UC MEXUS [(accessed on 11 February 2020)]. Arroyo la Junta: Una Joya de Biodiversidad en la Reserva de la Biosfera Sierra la Laguna/a Biodiversity Jewel in the Sierra la Laguna Biosphere Reserve. 159 pp.
  • Vasilikopoulos A, Balke M, Beutel R, Donath A, Podsiadlowski L, Pflug J, Waterhouse R, Meusemann K, Peters R, Escalona H, Mayer C, Liu S, Hendrich L, Alarie Y, Bilton D, Jia F, Zhou X, Maddison D, Niehuis O, Misof B (2019) Phylogenomics of the superfamily Dytiscoidea (Coleoptera: Adephaga) with an evaluation of phylogenetic conflict and systematic error. Molecular Phylogenetics and Evolution 135: 270–285.
  • Vives E (2001) The systematic position of Vesperoctenus flohri Bates, 1891 and the taxonomic status of the Vesperidae (Coleoptera). Occasional Papers of the Consortium Coleopterorum 4(1): 35–44.
  • Vives E (2005) Revision du genre Vesperus Dejean 1821 (Coleoptera: Cerambycidae). Annales de la Société Entomologique de France 40(3–4): 437–457.
  • Waterhouse RM, Zdobnov EM, Tegenfeldt F, Li J, Kriventseva EV (2011) OrthoDB: the hierarchical catalog of eukaryotic orthologs in 2011. Nucleic Acids Research 39 (Database issue): D283–288.
  • Waterhouse RM, Tegenfeldt F, Li J, Zdobnov EM, Kriventseva EV (2013) OrthoDB: a hierarchical catalog of animal, fungal and bacterial orthologs. Nucleic Acids Research 41 (Database issue): D358–365.
  • Werren JH, Richards S, Desjardins CA, Niehuis O, Gadau J, Colbourne JK, Beukeboom LW, Desplan C, Elsik CG, Grimmelikhuijzen CJ, Kitts P, Lynch JA, Murphy T, Oliveira DC, Smith CD, van de Zande L, Worley KC, Zdobnov EM, Aerts M, Albert S, Anaya VH, Anzola JM, Barchuk AR, Behura SK, Bera AN, Berenbaum MR, Bertossa RC, Bitondi MM, Bordenstein SR, Bork P, Bornberg-Bauer E, Brunain M, Cazzamali G, Chaboub L, Chacko J, Chavez D, Childers CP, Choi JH, Clark ME, Claudianos C, Clinton RA, Cree AG, Cristino AS, Dang PM, Darby AC, de Graaf DC, Devreese B, Dinh HH, Edwards R, Elango N, Elhaik E, Ermolaeva O, Evans JD, Foret S, Fowler GR, Gerlach D, Gibson JD, Gilbert DG, Graur D, Gründer S, Hagen DE, Han Y, Hauser F, Hultmark D, Hunter HC 4th, Hurst GD, Jhangian SN, Jiang H, Johnson RM, Jones AK, Junier T, Kadowaki T, Kamping A, Kapustin Y, Kechavarzi B, Kim J, Kim J, Kiryutin B, Koevoets T, Kovar CL, Kriventseva EV, Kucharski R, Lee H, Lee SL, Lees K, Lewis LR, Loehlin DW, Logsdon JM Jr, Lopez JA, Lozado RJ, Maglott D, Maleszka R, Mayampurath A, Mazur DJ, McClure MA, Moore AD, Morgan MB, Muller J, Munoz-Torres MC, Muzny DM, Nazareth LV, Neupert S, Nguyen NB, Nunes FM, Oakeshott JG, Okwuonu GO, Pannebakker BA, Pejaver VR, Peng Z, Pratt SC, Predel R, Pu LL, Ranson H, Raychoudhury R, Rechtsteiner A, Reese JT, Reid JG, Riddle M, Robertson HM, Romero-Severson J, Rosenberg M, Sackton TB, Sattelle DB, Schlüns H, Schmitt T, Schneider M, Schüler A, Schurko AM, Shuker DM, Simões ZL, Sinha S, Smith Z, Solovyev V, Souvorov A, Springauf A, Stafflinger E, Stage DE, Stanke M, Tanaka Y, Telschow A, Trent C, Vattathil S, Verhulst EC, Viljakainen L, Wanner KW, Waterhouse RM, Whitfield JB, Wilkes TE, Williamson M, Willis JH, Wolschin F, Wyder S, Yamada T, Yi SV, Zecher CN, Zhang L, Gibbs RA (2010) Functional and evolutionary insights from the genomes of three parasitoid Nasonia species. Science 327(5963): 343–348.
  • Wickham H (2016) ggplot2: Elegant Graphics for Data Analysis. Springer-Verlag New York.
  • Zhan S, Merlin C, Boore JL, Reppert SM (2011) The monarch butterfly genome yields insights into long-distance migration. Cell 147(5): 1171–1185.