Odier coll.; NHMUK • 2 ♂♂; Valencia; 39°25'09" N, 0°20'13" W; 17 Apr. 2022; M. Dufaü coll.; male inflorescence of *C. humilis*; JHAR04786; CBGP • 2 ♀♀; Andalusia, Motril; 36°44'24" N, 3°31'45" W; 11 Apr. 1971; J. Péricart coll.; RB • 1 spec. (sex undetermined); Malaga, NW of Otivar, Sierra del Chapparal; 36°49'30" N, 3°42'29" W; 8 May 2013; P. Stüben and A. Schütte coll.; *Quercus ilex* L. sieving; ZFMK (identification by genetic assignation, GenBank acc. MK891322) • 1 ♂; Mallorca Island, Alcúdia; 39°51'07" N, 3°07'38" E; 13 Apr. 1964; W. Leibmann coll.; NHMUK • 1 ♀; same locality as for preceding; 4 May 2000; R. Rober coll.; NHMUK.

UNITED KINGDOM • 1 ♂, 2 ♀♀; Gibraltar; 36°07'55" N, 5°21'03" W; NHMUK • 13 ♂♂, 23 ♀♀; Gibraltar; 36°07'55" N, 5°21'03" W; J.J. Walker coll.; NHMUK.

**Redescription (♂)**

**MEASUREMENTS.** Body length 2.5–3.7 mm.

**COLOR.** Body integument pale brown, scutellar shield, pronotum and head including rostrum generally darker, in some individuals interstriae 1, sometimes 2 darker as well; elytra and pronotum with minute recumbent whitish setae, glabrous in appearance.

**HEAD.** Rostrum slightly longer than pronotum in lateral view (1.1–1.15×), regularly downcurved; in dorsal view 4 to 5.5× as long as wide, integument densely punctate, forming 5 longitudinal carinae; antennae inserted near apical ¼ of length in lateral view; head capsule densely and coarsely punctate in dorsal view, forehead flat or slightly concave; eyes convex, exceeding lateral curve of head capsule in dorsal view; antennal funicle with first segment 2–2.2× as long as wide, slightly longer than segments 2–3 together, 2 longer than wide, 3–7 transverse.

**PRONOTUM.** Wider than long (W:L ratio: 1.34–1.45), widest at basal ⅓ of length, 0.95–1× as wide there as elytra at humeral angles, sides slightly convex, converging regularly from basal ⅓ to apex, lateral carina forming notch near middle of length; apical constriction distinct, about as deep as width of scape in middle of length; integument with punctures rounded, space between punctures dull, micropunctate, wider or narrower than diameter of punctures in middle, narrower anteriorly and laterally.

**METATHORAX.** Metanepisterna with recumbent white setae, non-contiguous.

**ELYTRA.** Longer than wide (W:L ratio: 0.68–0.77); sides slightly convex, widest near middle of length; humeri raised; apex jointly rounded or notched at level of suture; striae with punctures about ⅓–¼ as wide as width of interstriae; interstriae slightly convex, interstriae 5 sometimes raised in basal ⅔, 9 flat; scutellar shield rounded, coated with few small recumbent scales, not concealing integument.

**ABDOMEN.** Underside with ventrites 1–2 darker than 3–5, with minute recumbent whitish setae, not contiguous. Stridulatory plate with lines of 10–11 granules ⅔× as long as median line, slightly convex. Central sclerotized area longer than wide, truncate at base (Fig. 24F).

**LEGS.** Profemora moderately thickened near middle of length; protibiae with external margin straight, internal slightly bisinuate, setae on apical half of internal margin recumbent, at most as long as claws; claws simple.

**Fig. 24** (next page). Habitus and life history of *Derelomus chamaeropis* (Fabricius, 1798). **A–B.** Dorsal (A) and lateral (B) habitus of male, pale variant with 5th interstria not raised (Morocco) (JHAR03220-01). **C.** Dorsal habitus of male, dark variant with 5th interstria costate, pronotum wider (Sicily) (JHAR04886-05, CBGP). **D.** Head and pronotum in lateral view of female, dark variant (Sicily) (JHAR04886-06). **E.** Penis in dorsal (left) and lateral (right) views (JHAR03220-01, CBGP). **F.** Stridulatory plate (JHAR03220-01, CBGP). **G.** Stands of *Chamaerops humilis* L. (Arecaceae) on the western coast of Spain. **H.** Adult of *D. chamaeropis* visiting the male inflorescences of *C. humilis*, note the pollen grains attached to the cuticle. **I.** Male inflorescences of *C. humilis* in anthesis. **J–K.** Larva and pupa of *D. chamaeropis* in dry stipe of male inflorescences of *C. humilis*. Scale bar: A–D=1 mm; E–F not to scale.
<figure><img src="image_62.png" /><figcaption>C</figcaption></figure>

<figure><img src="image_63.png" /><figcaption>E</figcaption></figure>

<figure><img src="image_64.png" /><figcaption>F</figcaption></figure>

<figure><img src="image_65.png" /><figcaption>B</figcaption></figure>

<figure><img src="image_66.png" /><figcaption>D</figcaption></figure>

<figure><img src="image_67.png" /><figcaption>G</figcaption></figure>

<figure><img src="image_68.png" /><figcaption>H</figcaption></figure>

<figure><img src="image_69.png" /><figcaption>I</figcaption></figure>

<figure><img src="image_70.png" /><figcaption>J</figcaption></figure>

<figure><img src="image_71.png" /><figcaption>K</figcaption></figure>

<figure><img src="image_72.png" /></figure>
**TERMINALIA.** Body of penis elongate (W : L ratio: 0.25), about 1.3 × as long as apodemes; sides subparallel in dorsal view, widest in basal 2/3, converging apicad from apical 1/3, apex acuminate; in lateral view curvature stronger in basal 1/3 of length, width narrowing regularly from near 1/2 of length, apex slightly curved upward (Fig. 24E).

**Sexual dimorphism**

Females can be distinguished from males by the rostrum which is narrower and longer in dorsal view. In lateral view, the rostrum is distinctly longer than the pronotum in females (slightly longer than pronotum in males; Fig. 24B–D). The antennae are inserted at apical 1/3 in females (1/4 in males).

**Variation**

This species is quite polymorphic. Morphological variation occurs at two levels. First, *D. chamaeropis* consists of genetically isolated lineages distributed around the western part of the Mediterranean region (see Remarks section below). These populations show slight morphological divergences. For example, the rostrum of males is on average slightly longer and narrower in populations from Spain (mean W : L ratio: 0.24 (0.21 – 0.25); width measured at antennal insertion) than those from Sicily (mean W : L ratio: 0.18 (0.15 – 0.22)). Within the eastern clade, rostrum length and width also vary slightly between the French, Italian and Moroccan specimens examined. In addition to these geographic variations, the phenotype of adults in a given population is also variable. Variation includes ‘major’ males, with a comparatively wide pronotum and 5th elytral interstriae raised and ‘minor’ males with a comparatively narrow pronotum and 5th elytral interstriae flat. Color variation ranges from specimens uniformly pale brown, to specimens with brown elytra and a dark brown pronotum, head and rostrum (Fig. 24A–D), more rarely with the body (head, pronotum and elytra) uniformly dark brown.

**Life history**

*Derelomus chamaeropis* is the specific brood-site pollinator of the Mediterranean dwarf palm: *Chamaerops humilis* (Arecaceae; Anstett 1999). Adults fly and reach inflorescences of both male and female plants, attracted by scents emitted by leaves (Dufaü *et al.* 2002), thus transferring pollen to the stigma of female inflorescences. They mate and oviposit in the rachis of the inflorescences (Anstett 1999; Fig. 24I–K) and larvae develop in the stem of inflorescences, mainly on male plants. Successful larval development has also been described in female plants, but with markedly lower frequency and densities than on male plants (Dufaü & Anstett 2004; Jácome-Flores *et al.* 2018). Adults are active during the flowering season of *C. humilis*, from March to June. Larval development takes place in the decaying woody tissues of rachises from early summer to late winter. Pupation takes place within the rachises. As for many brood-site pollination systems involving weevils, *C. humilis* is co-pollinated by the sap beetle *Meligethinus pallidulus* (Erichson, 1843) also developing on male inflorescences (JH obs.) and visiting females ones (M. Dufaü pers. com.). Both species are efficient pollinators, even for isolated populations of palms resprouting after fire (García *et al.* 2018). The report of *D. chamaeropis* on *P. canariensis* (Lepseme 1941) is likely accidental.

**Distribution**

Coastal regions of the Western Mediterranean area: Algeria, Croatia, France, Italy, Morocco, Spain, including islands (Corsica, Sardinia, Sicily, Malta; Caldara 2013).

**Remarks**

In Fabricius’ collection housed at ZMK, a unique female specimen under the identification name “*Curculio* (=*Rhynchaenus*) *chamaeropsis* F., 1798” and fitted with a red type label was located. This specimen is the holotype of *Curculio chamaeropis* Fabricius, 1798 and was labelled accordingly. In Shoenherr’s collection, housed at NHRS, the holotype of *Derelomus chamaeropis* var. *β* Gyllenhal,
1836 was located and labelled accordingly. In the same collection, a male specimen bearing a red type label and corresponding in all aspects to the description of *Derelomus subcostatus* Boheman, 1844 was located. This specimen is the holotype of *Derelomus subcostatus* Fåhraeus, 1844 and was labelled accordingly. Based on a detailed examination of these types and a molecular analysis of specimens newly sampled in the region of the type localities, it is stated here that the specimens used to describe *Derelomus chamaeropis* var. α and β and *D. subcostatus* all belong to the same species. As a result, the names *Derelomus subcostatus* Fåhraeus, 1844 and *Derelomus chamaeropis* var. β Gyllenhal, 1836 are proposed as junior synonyms [new synonymy] of *Derelomus chamaeropis* (Fabricius, 1798). The elements used to reach this conclusion are detailed hereafter.

*Derelomus chamaeropis* was described on a female from N Morocco. This species is specific to *Chamaerops humilis* (see Life history section below) and follows the distribution range of this palm in the western Mediterranean region (Caldara *et al.* 2013). *Derelomus subcostatus* was described from a male specimen from Sardinia Island (Italy), the specimens related to this species being also associated with *C. humilis* (Veyret 1940). The limited details in the original descriptions and the substantial intraspecific variation found in *D. chamaeropis* have resulted in confusions in the interpretation of both species. For some authors, *D. subcostatus* is a valid species occurring in sympatry with *D. chamaeropis* (Tempère & Péricart 1989; Caldara *et al.* 2013) while others treat it as a morphological variant of *D. chamaeropis* (Hoffmann 1958). In order to clarify this situation, fourteen populations (1-4 specimens per population) were sampled across the distribution range of *C. humilis* (France, mainland and Corsica Island; Italy, Sicily and Pantelleria Island; Morocco and Spain; see detail in Other material section) and sequenced for the standard barcode fragment (see Material and methods section for protocol). This sampling included specimens identified by direct comparison with the holotypes: a female specimen of *D. chamaeropis* from Morocco (JHAR03220-02) and a male specimen of *D. subcostatus* from Italy (Sicily; JHAR04886-08). The populations from Spain also included specimens related to the varieties α and β of *D. chamaeropis* (male with or without costate elytra, with or without dark head and pronotum; Fig. 24A-C). *Derelomus chamaeropis* was recovered as a single species for all the species delimitation methods used (Fig. 34), despite intraspecific distances ranking up to 3.4% between specimens from Morocco (KC783895) and Spain (JHAR07851-02). Also, the variations observed at population level in the male are not associated with genetic divergences in mitochondrial sequences (Fig. 33). As such, all specimens previously referred to as *D. chamaeropis* and *D. subcostatus* are in fact a unique species forming a ring of more or less isolated populations distributed in the western mediterranean area. This phylogeographic pattern is typical of Mediterranean species and results from the isolation of populations in southern European refugia during glacial maxima (Hewitt 2000). This pattern is also in agreement to some extent with the genetic structure of *Chamaerops humilis* that shows substantial differentiation among populations across its distribution range (De Cauwer *et al.* 2025). This may suggest that both this palm and its associated pollinator experienced similar fragmentation of distribution range during glacial maxima. At population level, specimens show substantial morphological variation, in particular in males, a condition widely encountered in weevils engaged in brood-site pollination (Haran *et al.* 2020, 2023a) and not associated with genetic divergence.

The Sardinian specimen used by Boheman to describe *Derelomus subcostatus* combines the two factors of morphological variability in *D. chamaeropis*: it is slightly morphologically divergent from populations from Morocco used to describe *D. chamaeropis* (rostrum comparatively short) and exhibits the costate elytra of a 'major' male, thus showing a substantial apparent divergence with the female holotype of *D. chamaeropis*. But as detailed above, all these specimens belong to the same species. The species name *Ochninulus antigae* Reitter, 1887 was set in synonymy with *Derelomus chamaeropis* (Fabricius 1798) by Reitter himself (Reitter 1887), the specimen used to describe this species was collected in Barcelona (Spain) [not verified in the context of this study].
*Derelomus piriformis* (Hoffmann, 1938)
Fig. 25

*Pseudoderelomus piriformis* Hoffmann, 1938a: 107.

*Derelomus kocheri* Hoffmann, 1957: 83. **Syn. nov.**

*Neoderelomus piriformis* — Hoffmann 1938b (genus name preoccupied). — Abbazi & Osella 1992 (distribution). — Piry & Gompel 2002 (description of larva, biology, distribution). — Alonso-Zarazaga & Lyal 1999 (distribution). — Meekijjaroenroj & Anstett 2003 (biology).

*Derelomus piriformis* — Franz 2006: 275 (new combination). — Haran *et al.* 2022a (phylogenetic relationship).

**Diagnosis**

In the *D. ephippiger* group, this species can be distinguished by the combination of a uniformly pale brown integument, and males with the elytra widest beyond the middle of the length, protibiae with an internal comb of elongate setae and a comparatively moderately long rostrum (4.5–5 × as long as wide in dorsal view). Females can be distinguished by the distinct narrowing at the base of the rostrum. *Derelomus piriformis* is closely related and sometimes sympatric with *D. chamaeropis*, but in the latter species, males have the elytra slightly convex in dorsal view and the protibiae lacking an internal comb (very convex elytra and protibiae with a comb of setae in *D. piriformis*) and females have a shorter and wider rostrum (Fig. 24D). Uncorrected *p*-distances between these species range from 8.0 to 8.8%. GenBank accession number for the corresponding DNA barcode: OK188814.

**Material examined**

**Lectotype** of *Pseudoderelomus piriformis* (here designated)

MADEIRA • ♂; “Funchal; Madère;15.8.1936; Balachowsky [Alferd Balachowsky, french entomologist]” “*Pseudoderelomus*; *piriformis*; Hoffm.” “genotype; TYPE” “TYPE[red label]” “Museum Paris; 1968; Col. A. Hoffmann” “Lectotype ♂; *Derelomus piriformis*; (Hoffmann, 1938); J. Haran des. 2025”; MNHN.

**Holotype** of *Derelomus kocheri* Hoffmann, 1957

MOROCCO • ♀; “Rabat (Maroc); (Kocher) 10. 52 (Oct. 1952)” “*Derelomus*; *kocheri* m.[me, handwritten by Hoffmann]; A. Hoffmann det.” “Typus [red label]” “Museum Paris; 1968; Col. A. Hoffmann” “Holotype ♀; *Derelomus*; *kocheri*; Hoffmann, 1957; J. Haran 2025” “*Derelomus*; *piriformis* (Hoff.); Haran det. 2024”; MNHN.

**Paralectotype**

MOROCCO • 1 ♂; Rabat; 34°02′47.0″ N, 6°50′25.0″ W; 1 Feb. 1930; Bremond coll.; MNHN.

**Other material**

FRANCE • 4 ♂♂, 12 ♀♀; Hérault, Aéroport de Maugio; 43°33′57″ N, 3°57′00″ E; 5 Nov. 2000; S. Piry coll.; male inflorescences of *Phoenix canariensis*; MNHN • 2 ♂♂, 1 ♀; Montpellier, Hopital Gui de Chauliac; 43°37′56″ N, 3°51′18″ E; 1 Nov. 2000; S. Piry coll.; male inflorescences of *Phoenix canariensis*; CBGP • 2 ♂♂, 1 spec. (preserved in ethanol); Corsica, Macinaggio; 42°57′32″ N, 9°27′10″ E; 8 Oct. 2015; J. Haran coll.; male inflorescences of *P. canariensis*; CBGP • 1 ♀; Le Cannet; 43°34′41″ N, 7°00′26″ E; 8 Aug. 1954; P. Bonadona coll.; PW • 1 ♂; La Garde; 43°07′30″ N, 6°01′19″ E; 9 Jul. 1953; P. Veyret; PW • 1 ♀; La Garde; 43°07′30″ N, 6°01′19″ E; 7 Nov. 1952; P. Veyret; PW • 1 ♂, 2 ♀♀; Toulon; 43°08′13″ N, 5°55′26″ E; Dec. 1936; P. Veyret; PW • 1 ♂; Toulon; 43°08′13″ N, 5°55′26″ E; Nov. 1936; P. Veyret; PW • 1 ♀; Nice, Jardin Alsace Lorraine; 43°41′52″ N, 7°15′26″ E; 28 Feb. 1999; S. Piry coll.; ex. flowers of *Phoenix* sp.; CBGP.
ITALY • 3 ♂♂, 3 ♀♀; Western Liguria, Varigotti; 44°10′52″ N, 8°23′54″ E; 4 Oct. 1966; G. Bartoli coll.; *Phoenix* sp.; JHAR07418; CBGP • 1 ♂; Roma, Tuscolano; 41°52′12″ N, 12°32′18″ E; 17 Sep. 1946; L. Magnano coll.; at light; NHMUK.

MOROCCO • 1 ♀; Oujda; 34°40′23″ N, 1°54′18″ W; 13 Dec. 1998; G. Chavanon coll.; on the ground; JHAR7419; CBGP.

PORTUGAL • 1 ♂, 1 ♀; Porto Santo Island [Madeira archipelago]; 33°03′50″ N, 16°19′26″ W; 3 Apr. 2015; J-D. Chapelin Viscardi coll.; male inflorescences of *P. canariensis*; JHAR2983; CBGP.

SPAIN • 1 ♀; Las Palmas de Gran Canaria, Santa Catalina area; 28°08′21″ N, 15°26′11″ W; 10 Mar. 1903; P. Lesne coll.; emerged from plant debris collected at base of *Tamarix* L. sp.; MNHN • 1 ♂; Melilla [located on the northern coast of Morocco]; 35°18′04″ N, 2°56′13″ W; May 1951; F. Codina coll.; MNHN • 1 ♀; Melilla; 35°18′04″ N, 2°56′13″ W; Dec. 1951; Prado Alcaide coll.; MNHN.

UNITED STATES OF AMERICA • 1 ♂; California, Roseville; 38°45′41.3″ N, 121°17′22.2″ W; 31 Aug. 2020; G. Forister coll.; (https://bugguide.net/).

**Redescription (♂)**

**MEASUREMENTS.** Body length 3.0–3.5 mm.

**COLOR.** Body integument uniformly pale brown, apex of rostrum slightly darker; elytra and pronotum with minute recumbent whitish setae, glabrous in appearance.

**HEAD.** Rostrum slightly longer than pronotum in lateral view (1.1×), regularly and moderately downcurved; in dorsal view 4.5–5× as long as wide, integument densely punctate, forming 5 longitudinal carinae; base of rostrum slightly constricted laterally; antennae inserted near apical ¼ of length in lateral view; head capsule densely punctate in dorsal view, forehead flat; eyes convex, exceeding lateral curve of head capsule in dorsal view; antennal funicle with first segment 2.8× as long as wide, slightly longer than segments 2–3 together, 2 about 2× as long as wide, 3–7 transverse.

**PRONOTUM.** Wider than long (W:L ratio: 1.50–1.63), widest at base, 0.95× as wide there as elytra at humeral angles, sides slightly convex, converging regularly from base to apex; apical constriction distinct, about as deep as width of scape at base; integument dull, punctures superficial, hardly visible among granules.

**METATHORAX.** Metanepisterna with recumbent white setae, non-contiguous.

**ELYTRA.** Longer than wide (W:L ratio: 0.85); sides strongly convex, widest beyond middle of length; humeri raised; apex jointly rounded or notched at level of suture; striae with punctures about ¼–½ as wide as width of interstriae; interstriae flat, interstriae 5 raised, more so between basal and apical 1/6 of length, 9 flat, enlarged and merged with 10th at base; scutellar shield rounded, coated with few small recumbent scales, not concealing integument.

**ABDOMEN.** Underside with ventrites uniformly pale brown, with overlapping whitish setae, usually longer in middle of each ventrite. Stridulatory plate with lines of 8 granules ½× as long as median line, slightly convex. Central sclerotized area longer than wide, rounded at base (Fig. 25F).

**LEGS.** Profemora moderately thickened near middle of length; protibiae with external margin straight, internal slightly bisinuate, bearing a comb of setae on apical ¾ of length, setae about as long as 4th tarsomere; claws simple.
<figure><img src="image_73.png" /><figcaption>**Fig. 25.** Habitus and life history of *Derelomus piriformis* (Hoffmann, 1938). **A.** Lectotype, ♂, habitus in dorsal view (Madeira, MNHN). **B.** ♀, habitus in dorsal view (Italy) (JHAR07418-02, CBGP). **C.** Same as A, ♂, habitus in lateral view. **D.** Same as B, ♀, habitus in lateral view. **E.** Same as A, penis in dorsal (left) and lateral (right) views. **F.** Same as A, stridulatory plate. **G.** The Canary Island date palm (*Phoenix canariensis* H.Wildpret; Arecaceae) in natural habitat (credit: Gertjan van Noord). **H.** Male inflorescences in anthesis of *P. canariensis* (credit: Jesús Cabrera). Scale bar: A–D=1 mm; E–H not to scale.</figcaption></figure>
### TERMINALIA
Body of penis elongate (W: L ratio: 0.27), about 1.5 × as long as apodemes; sides slightly convex in dorsal view, widening from base to apical 1/3, from there converging apicad, converging abruptly in apical 1/6, apex rounded; in lateral view curvature stronger in basal 2/5 of length, width narrowing regularly from near 1/2 of length, apex acute, straight (Fig. 25E).

## Sexual dimorphism

Females can be distinguished from males by the rostrum which is more shiny and distinctly narrowed near base. In lateral view, rostrum is distinctly longer (1.25 ×) than pronotum in females (1.1 × in males; Fig. 25C-D). The antennae are inserted near the apical 1/3 in females (1/4 in males).

## Life history

This species is the specific brood-site pollinator of *Phoenix canariensis* (Arecaceae), the Canary Island date palm. Adults visit male inflorescences at anthesis stage. Development of larvae and pupae takes place in the enclosed flowers. The development continues on the ground after flower abscission, the declining flower forming a cell protecting the pupa. Each flower hosts only one larva (Piry & Gompel 2002). In this pollination by deceit system, adults carrying pollen and searching for male inflorescences are inadvertently attracted by female inflorescences (producing similar olfactory clues) where they transfer pollen to stigma (Meekijjaroenroj & Anstett 2003). *Derelomus piriformis* was also recorded on male inflorescences of *Chamaerops humilis* in the South of France (Veyret 1940, identified as *D. subcostatus* in the latter paper), but there is no evidence that it can use this palm for larval development. Adult specimens may be found active on its hosts as long as male inflorescences in anthesis are available. This flowering phenology seems to vary across areas of introduction, from late summer to late autumn in the south of France (Piry & Gompel 2002), all year round in Israel (Friedman 2006). Adults are attracted by UV lights (Piry & Gompel 2002).

## Distribution

This species is native to the Canary Island where *Phoenix canariensis* originates. This palm is widely traded and planted for ornamental purposes, and *D. piriformis* has probably been transported simultaneously. It is now widely distributed in the Mediterranean region and Madeira Island (sorted by date of first record): Morocco, 1930 (Hoffmann, 1938); Madeira (Portugal) & France, 1936 (Hoffmann 1938; Veyret, 1940; PW collection); Italy, 1966 (CBGP coll.; Abbazzi & Osella 1992); Israel, 1976 (Friedman 2006); Greece, 1990 (Kakiopoulos *et al.* 2022); Corsica Island (France), 2015 (CBGP coll.). Further records: Canary Islands and Spanish mainland (Caladara 2013). Introduced to South America (Chile) with its host plant (Elgueta & Marvaldi 2006) and to North America (California).

## Remarks

In A. Hoffmann's collection housed at MNHN, three specimens corresponding in all aspects to the material listed in the original description of *D. piriformis* and bearing a red type label were located. As no specific specimen was designed in the type series, a specimen (Funchal, Madeira) was designated as the lectotype for this species [here designated] and was relabelled accordingly. The two other specimens from the series (Rabat, Morocco) were labelled as paralectotypes. Due to the uncommon sexual dimorphism in this species (males have the elytra more convex than females, a condition rare in weevils), the specimens considered by A. Hoffmann as males are in reality females, and conversely (Piry & Gompel 2002). The sex of specimens reported above corresponds to verified specimens, based on dissection when necessary. In the same collection, a female specimen labelled "*Derelomus kocheri*" by the hand of Hoffmann and bearing a red type label was located. For the reasons described above, this specimen is not a male as reported in the original description but a female one. This specimen is the holotype of *Derelomus kocheri* Hoffmann, 1957 and was labelled accordingly. Detailed examination of external and internal morphological features of this specimen and comparison with type and non-type
material of *D. piriformis* from various regions revealed no relevant differences. In addition, the type
locality of *D. kocheri* (Rabat, Morocco) is the same as the paralectotype of *D. piriformis*. As a result of
this, the species name *Derelomus kocheri* Hoffmann, 1957 is proposed as a junior synonym of *Derelomus
piriformis* (Hoffmann, 1938) [new synonymy].

Due to the early introduction of *D. piriformis* around the Mediterranean Sea and to the unclear taxonomic definition of *D. subcostatus* Boheman, 1844 (= *D. chamaeropis* (Fabricius, 1798), see Remarks section under that species), many specimens referred to as *D. subcostatus* in early collections and publications correspond to *D. piriformis* in reality. This is relevant to the publication of Veyret (1940).

*Derelomus peglerae* Haran sp. nov.

urn:lsid:zoobank.org:act:D4FD6AE5-7DD0-4898-AED8-8A9817BFEB0C

Fig. 26

Diagnosis

In the *D. ephippiger* group, this species can be distinguished by the combination of very long and almost straight rostrum, only slightly downcurved in lateral view (Fig. 26A–C) and an elytral pattern forming longitudinal dark strips near the middle of the length of interstriae 3–4. This species is unique in having a series of only 3 granules on the stridulatory plate (Fig. 26E). GenBank accession number for the corresponding DNA barcode: ON553433/PV698480–81.

Etymology

The species is dedicated to Alice Marguerite Pegler (1861–1929), pioneer field botanist and entomologist in the Eastern Cape Province of the Republic of South Africa. Through observations and specimens’ collection, she was an early contributor to the recognition of the importance of beetles, including weevils in the pollination of Cycads. She also collected the series used by Hesse to describe *D. postfasciatus*.

**Material examined**

Holotype

REPUBLIC OF SOUTH AFRICA – **Limpopo Province** • ♂; “South Africa, Tvl.; Hans Merensky Nat.; Res. 23.42S 30.44E [23°41′42.0″ S 30°40′05.2″ E]; 23-25.i.1987; R. Oberprieler” “collected on; leaves of; *Phoenix*; *reclinata*” “National coll.; of insects; Pretoria, S. Afr” “Holotype ♂; *Derelomus peglerae*; Haran, 2025”; SANC.

Paratypes

ANGOLA • 1 ♂, 1 ♀; no precise locality; 7 Jun. 1928; NHMUK.

ETHIOPIA • 1 ♀; “Abyssinie” [no precise locality]; 1882; A. Raffray; MNHN.

KENYA • 2 ♂♂; Nairobi, Scottlab [now the National Agricultural Research Laboratories, NAR]; 1°15′29.2″ N, 36°46′23.1″ E; 30 March 1947; R.H. Le Pelley coll.; on *Tecomia* Juss. sp.; NHMUK.

REPUBLIC OF SOUTH AFRICA – **Eastern Cape Province** • 1 ♂, 1 ♀; Mbotyi campsite; 31°27′54.0″ S, 29°43′48.0″ E; Dec. 2009; Ş. Procheş coll.; male and female inflorescences of *Phoenix* sp.; JHAR07433; CBGP. – **KwaZulu-Natal Province** • 4 ♂♂, 2 ♀♀, 2 specs (preserved in ethanol); St Lucia; 28°22′59.8″ S, 32°24′33.4″ E; 5 Jan. 2019; J. Haran coll.; male inflorescence of *Phoenix reclinata*; JHAR02043; CBGP • 1 ♂; iSimangaliso Wetland Park, Ozabeni, 3 km SSW of Sodwana Camp; 27°38′10.8″ S, 32°34′55.2″ E; 22 Oct. 2014; E Grobbelaar coll.; SANC-MCOL-0374/JHAR02335; SANC. – **Limpopo Province** • 1 ♀; Nwanedi Provincial Park; 22°37′34.8″ S, 30°24′21.6″ E; 8 Feb.
1994; R. Oberprieler coll.; collected from *Phoenix reclinata*; SANC • 1 ♂; Hans Merensky Nature Reserve; 23°41'42.0" S, 30°40'05.2" E; 23–25 Jan. 1987; R. Oberprieler coll.; collected on leaves of *Phoenix reclinata*; SANC • 5 ♂♂, 2 ♀♀; same collection data as for preceding; ANIC • 1 ♂, 2 ♀♀; Mmabolela Estate; 22°37'52" S, 28°16'34" E; 20–24 Nov. 1994; F. Genier coll.; savanna light trap; CMNC. – *Mpumalanga Province* • 1 ♂; Kruger Park, Satara Camp; 24°23'40.0" S, 31°46'41.6" E; 15–17 Dec. 1985; S. and J. Peck coll.; *Acacia* grasslands; CMNC.

UGANDA • 2 ♀♀; Kawanda; 0°25'12.0" N, 32°32'24.0" E; 15 Feb.–6 Mar. 1958; P. Whalley; mercury vapor light trap; NHMUK.

**Redescription (♂)**

MEASUREMENTS. Body length 2.8–3.6 mm.

COLOR. Body integument uniformly pale brown, club, medial line of pronotum dark brown, apex of rostrum, postocular strip also dark in some specimens, more rarely with a longitudinal dark strip near middle of length of interstriae 3–4; elytra and pronotum with minute recumbent whitish setae, glabrous in appearance.

HEAD. Rostrum distinctly longer than pronotum in lateral view (1.28×), slightly and regularly downcurved; in dorsal view about 5× as long as wide, integument densely punctate, forming 5 longitudinal carinae; antennae inserted near apical ¼ of length; head capsule densely punctate in dorsal view, forehead flat; eyes convex, exceeding lateral curve of head capsule in dorsal view; antennal funicle with first segment 2.5× as long as wide, slightly longer than segments 2–3 together, 3–4 isodiametric, 5–7 transverse.

PRONOTUM. Wider than long (W:L ratio: 1.35–1.41), widest near base, narrower there than elytra at humeral angles (0.75×), side slightly rounded, converging apicad, forming notch or blunt tooth near middle of length; apical constriction superficial, less deep than width of scape in middle of length; integument with punctures rounded, space between punctures rough, shiny, micropunctate, wider or narrower than diameter of punctures in middle, narrower laterally.

METATHORAX. Metanepisterna with recumbent white setae, non-contiguous.

ELYTRA. Longer than wide (W:L ratio: 0.73–0.77); sides slightly convex, widest near middle of length; humeri raised; apex rounded, notched or jointed at level of suture; striae with punctures about ⅓ to ¼ as wide as width of interstriae; interstriae flat or slightly convex, 5 raised into carina; scutellar shield rounded, concolorous, coated with small recumbent scales not concealing integument.

ABDOMEN. Underside uniformly pale brown, with minute recumbent whitish setae, not contiguous. Stridulatory plate with lines of 3 granules ⅔× as long as median line from base, space between granules 2–3× as wide as diameter of a granule. Central sclerotized area wide, wider than long, truncate at base (Fig. 26E).

LEGS. Profemora moderately thickened near middle of length; protibiae with external margin straight, internal bisinuate, setae on apical half of internal margin recumbent, at most as long as claws; claws simple.

TERMINALIA. Body of penis elongate (W:L ratio: 0.26), 2× as long as apodemes; sides concave in basal ⅔, convex in apical ⅓ in dorsal view, narrowing apicad in apical ¼, apex truncate; in lateral view curvature stronger in basal ⅓ of length, width narrowing regularly from apical ¼, apex curved upward (Fig. 26D).
<figure><img src="image_74.png" /><figcaption>**Fig. 26.** Habitus of *Derelomus peglerae* Haran sp. nov. **A–B, D–E.** Paratype, ♂ (Republic of South Africa) (JHAR02043-03, CBGP). **C.** Paratype, ♀ (same locality) (JHAR02043-01, CBGP). **A.** Habitus in dorsal view. **B–C.** Habitus in lateral view. **D.** Penis in dorsal (left) and lateral (right) views. **E.** Stridulatory plate. Scale bar: A–C = 1 mm; D–E not to scale.</figcaption></figure>
**Sexual dimorphism**

Female can be distinguished from males by the longer rostrum, 1.5 × as long as pronotum in lateral view (1.28 × in males). The antennae are inserted at the apical 2/5 of the rostrum (1/4 in males).

**Life history**

Adults are found on male inflorescences of *Phoenix reclinata* (Arecaceae), which probably constitutes the habitat for the larvae. This species is found on its host in sympatry with *D. costiger* Marshall, 1958 and *D. bivirgatus* Marshall, 1951. Adults were recorded in June, October and from December to March.

**Distribution**

*Derelomus peglerae* sp. nov. is known from Angola, Ethiopia, the Republic of South Africa and Uganda. In the Republic of South Africa, it was recorded from the coastal regions of the Eastern Cape and KwaZulu-Natal provinces, and inland in the Limpopo Province.

**_Derelomus baka_ Haran sp. nov.

urn:lsid:zoobank.org:act:288FC910-5687-42E8-A423-8C0643C74AC7

Fig. 27

**Diagnosis**

In the *D. ephippiger* group, this species can be distinguished by the combination of a rostrum distinctly longer than the pronotum in lateral view, the concolorous club and the elytra with a single dark spot near the middle of the suture. This species is close to *D. antonioui*, but in the latter species, the protibiae lack a comb of erect setae in the male and the body of the penis is more elongate.

**Etymology**

The species is dedicated to the Baka people, semi-nomadic hunter-gatherers of central Africa, including Cameroon. These people live in a remarkable and non-destructive equilibrium with forest ecosystems enabling the maintaining of the high biodiversity of these biomes.

**Material examined**

**Holotype**

CAMEROON • ♂; "Foumbot [5°30'21.6" N 10°38'02.4" E]; fév. [February] 67 [1967]" "Cameroun; B. de Miré [P. Bruno de Miré, french entomologist]" "Holotype ♂; *Derelomus baka*; Haran, 2025"; MNHN.

**Description (♂)**

**MEASUREMENTS.** Body length 3.0 mm.

**COLOR.** Body integument pale brown, apex of rostrum and reversed triangular spot extending on middle of length of interstriae 1–4 dark brown, head and pronotum darker than elytra; elytra and pronotum with minute recumbent whitish setae, glabrous in appearance.

**HEAD.** Rostrum distinctly longer than pronotum in lateral view (1.22 ×), slightly and regularly downcurved; in dorsal view about 4.5 × as long as wide, integument densely punctate, forming 5 longitudinal carinae; antennae inserted near apical 1/4 of length; head capsule densely punctate in dorsal view, forehead slightly concave; eyes convex, exceeding lateral curve of head capsule in dorsal view; antennal funicle with first segment 2 × as long as wide, slightly longer than segments 2–3 together, 3–7 transverse.
<figure><img src="image_75.png" /><figcaption>**Fig. 27.** Habitus of *Derelomus baka* Haran sp. nov., holotype, ♂ (Cameroon) (MNHN). **A.** Habitus in dorsal view. **B.** Habitus in lateral view. **C.** Penis in dorsal (left) and lateral (right) views. Scale bar: A–B=1 mm; C not to scale.</figcaption></figure>
**PRONOTUM.** Wider than long (W:L ratio: 1.32), widest near base, slightly narrower there than elytra at humeral angles (0.85×), side slightly rounded, converging apicad, forming a small notch near middle of length; apical constriction as deep as width of scape in middle of length; integument with punctures rounded, space between punctures rough, shiny, micropunctate, generally narrower than diameter of punctures in middle.

**METATHORAX.** Metanepisterna with recumbent white setae, non-contiguous.

**ELYTRA.** Longer than wide (W:L ratio: 0.76); sides convex, widest near middle of length; humeri raised; apex rounded, notched at level of suture; striae with punctures about 1/3 to 1/4 as wide as width of interstriae; interstriae flat or slightly convex, 5 raised into carina; scutellar shield rounded, concolorous, coated with small recumbent scales not concealing integument.

**ABDOMEN.** Underside uniformly pale brown, with minute recumbent whitish setae, not contiguous.

**LEGS.** Profemora moderately thickened near middle of length; protibiae with external margin straight, internal bisinuate, with comb of erect setae on apical 1/2 of length, setae longer than a claw; claws simple.

**TERMINALIA.** Body of penis elongate (W:L ratio: 0.27), about 1.5× as long as apodemes; sides subparallel in basal 2/3 of length, converging in apical 1/3 in dorsal view, forming short spatula in apical 1/7, apex rounded; in lateral view curvature stronger in basal 1/3 of length, width narrowing regularly from middle of length, apex bisinuate (Fig. 27C).

## Life history

Host plant unknown, the holotype was collected in February.

## Distribution

Cameroon.

## Remarks

A unique male specimen is known for this species. The stridulatory plate was damaged during the dissection and is therefore not displayed.

*Derelomus bivirgatus* Marshall, 1951

Fig. 28

*Derelomus bivirgatus* Marshall, 1951: 841.

## Diagnosis

In the *D. ephippiger* group, this species can be distinguished by the strong curvature of the rostrum in lateral view (Fig. 28B–C) and the sides of the elytra are very convex in the male. Such a strong curvature of the rostrum is only met in *D. brevis*, but in the latter species, the rostrum and the body are shorter and the sides of the elytra are moderately convex in the males (Fig. 21A–C). GenBank accession numbers for the corresponding DNA barcodes: PV698441–42–43.
**Material examined**

**Lectotype** (here designated)

KENYA • ♂; "Type [red label]" "Mac Arthur; Makindu [Kenya, 2°16′37.2″ S 37°49′08.4″ E] 5 37 [May 1937]" "Press by; Com Inst Ent; B M 1951-179" *Derelomus; bivirgatus* Mshl.; type ♂" "LECTOTYPE ♂; *Derelomus; bivirgatus*; Marshall, 1951; Haran des. 2025"; NHMUK.

**Paralectotypes**

KENYA • 6 ♂♂, 7 ♀♀; Makindu; 2°16′37.2″ S, 37°49′08.4″ E; May 1937; Mac Arthur coll.; NHMUK.

**Other material**

REPUBLIC OF SOUTH AFRICA – KwaZulu-Natal Province • 5 ♂♂, 4 ♀♀, 2 specs (preserved in ethanol); St Lucia; 28°22′59.8″ S, 32°24′33.4″ E; 5 Jan. 2019; J. Haran coll.; male inflorescence of *Phoenix reclinata*; JHAR02044; CBGP • 2 ♀♀; Kloof; 29°47′06.0″ S, 30°50′10.8″ E; Aug. 1926; R.E. Turner coll.; NHMUK.

UGANDA • 2 ♂♂, 1 ♀; Kawanda; 0°25′12.0″ N, 32°32′24.0″ E; 15 Feb.–6 Mar. 1958; P. Whalley; mercury vapor light trap; NHMUK.

**Redescription** (♂)

MEASUREMENTS. Body length 2.5–3.2 mm.

COLOR. Body integument pale brown, club, apex of rostrum, post-ocular area, scutellar shield and median line of pronotum usually darker or dark brown, basal 3/5 of 4th interstriae and 1st interstriae at 3/5 of length generally dark brown, forming empty rectangle on elytra; dorsum with minute recumbent whitish setae, glabrous in appearance.

HEAD. Rostrum slightly longer than pronotum in lateral view (1.07×), strongly and regularly downcurved, almost forming quarter circle; in dorsal view about 3.5–3.7× as long as wide, integument densely punctate, forming 5 longitudinal carinae; antennae inserted at apical 1/4 of length in lateral view; head capsule densely punctate in dorsal view, forehead flat; eyes convex, exceeding lateral curve of head capsule in dorsal view; antennal funicle with first segment 2× as long as wide, as long as segments 2–3 together, 3–7 transverse.

PRONOTUM. Wider than long (W : L ratio: 1.54), widest near base or at basal 1/3 of length, a little narrower there than elytra at humeral angles (0.83×), side rounded, converging apicad, forming notch and blunt tooth near middle of length; apical constriction distinct, about as deep as width of scape in middle of length; integument with punctures rounded, space between punctures rough, shiny, micropunctate, wider or narrower than diameter of punctures in middle, narrower laterally.

METATHORAX. Metanepisterna with recumbent white setae, non-contiguous.

ELYTRA. A little longer than wide (W : L ratio: 0.90); sides strongly convex, widest near basal 2/3 of length; humeri not distinct from the lateral curvature of elytra; apex rounded, notched or jointed at level of suture; striae with punctures about 1/3 to 1/4 as wide as width of interstriae; interstriae flat, 5 slightly raised on basal 3/5 of length; scutellar shield rounded, generally darker than elytra, coated with small recumbent scales not concealing integument.

ABDOMEN. Underside uniformly pale brown, with minute recumbent whitish setae, not contiguous. Stridulatory plate with lines of 9 granules more than 3/4 as long as median line from base, space between granules 2–3 × as wide as diameter of a granule. Central sclerotized area isodiametric, truncate at base (Fig. 28E).
<figure><img src="image_76.png" /><figcaption>**Fig. 28.** Habitus of *Derelomus bivirgatus* Marshall, 1951. **A–B, D–E.** ♂ (Republic of South Africa) (JHAR02044-03, CBGP). **C.** ♀ (same locality) (JHAR02044-04, CBGP). **A.** Habitus in dorsal view. **B–C.** Habitus in lateral view. **D.** Penis in dorsal (left) and lateral (right) views. **E.** Stridulatory plate. Scale bar: A–C=1 mm; D–E not to scale.</figcaption></figure>
LEGS. Profemora moderately thickened near middle of length; protibiae with external margin straight, internal bisinuate, setae on apical half of internal margin recumbent, shorter than length of claws; claws simple.

TERMINALIA. Body of penis elongate (W : L ratio: 0.24), about 2 × as long as apodemes; sides subparallel, in dorsal view, narrowing apicad in apical ¼ of length, apex rounded; in lateral view curvature moderate but stronger in basal ½ of length, width narrowing regularly apicad from ½ of length, apex acute, straight (Fig. 28D).

**Sexual dimorphism**

Female can be distinguished from males by the rostrum slightly more downcurved in lateral view and with antennae inserted at the apical ⅓ of the length (¼ in males). The elytra of females are also longer (W : L ratio: 0.78–0.83) than those of males (W : L ratio: 0.90).

**Life history**

This species was collected on male inflorescences of *Phoenix reclinata*, which probably constitutes the substrate for the larvae. Adults were collected in January, February, March, May and August.

**Distribution**

In our current knowledge only known from Kenya, Uganda and in the coastal region of the KwaZulu-
Natal Province of the Republic of South Africa.

**Remarks**

In the collection housed at NHMUK, a male specimen from Makindu labelled "*Derelomus bivirgatus*
Mshl., Type ♂" and bearing a red type label was located. As the description of this species does not
refer to a specific type specimen, this male is designated as the lectotype for *Derelomus bivirgatus*
Marshall, 1951 [here designated] and was labelled accordingly. The rest of the specimens were labelled
as paralectotypes. Specimens from Kenya and the Republic of South Africa show slight morphological
differences, in the latter the rostrum is more downcurved basally in both sexes. These differences are
related to regional variation, as observed in other species distributed in Eastern and Southern Africa
(*D. fasciatus*, *D. pallidus*, etc).

*Derelomus crypticus* Haran sp. nov.

urn:lsid:zoobank.org:act:D4075549-2910-459F-877E-6C443AD6F105

Fig. 29

**Diagnosis**

In the *D. ephippiger* group, this species can be distinguished by the comparatively strong curvature of the rostrum in lateral view (Fig. 29B–C) and the sides of the elytra moderately convex in males in dorsal view. This species is very close to and sympatric with *D. bivirgatus* but the latter is larger and has a longer rostrum (Fig. 28A–C). The new species is genetically well distinguished from *D. bivirgatus* (Fig. 34). GenBank accession number for the corresponding DNA barcode: ON553424.

**Etymology**

The specific epithet refers to the cryptic nature of this rare species. Specimens were first identified as aberrant forms of *D. bivirgatus* and the use of molecular data was critical for its recognition as a distinct species.
**Material examined**

**Holotype**

KENYA • ♂; "Mac Arthur; Makindu [Kenya, 2°16'37.2" S 37°49'08.4" E] 5 37 [May 1937]" "Press by; Com Inst Ent; B.M. 1981-315" "HOLOTYPE ♂; *Derelomus; crypticus*; Haran 2025"; NHMUK.

**Paratype**

REPUBLIC OF SOUTH AFRICA – **Western Cape Province** • 1 ♀; Pringle Bay; 34°21'00.0" S, 18°49'48.0" E; 28 Nov. 2019; J. Haran coll.; on male inflorescence of *Phoenix reclinata*; JHAR03078-03; CBGP

**Description (♂)**

**MEASUREMENTS.** Body length 2.0 mm.

**COLOR.** Body integument pale brown, head, club, the median line of pronotum slightly darker; pattern on elytra forming elongated dark spot beyond middle of length of interstriae 4, slightly extending on interstriae 3 and 5; dorsum with minute recumbent whitish setae, glabrous in appearance.

**HEAD.** Rostrum as long as pronotum in lateral view, strongly and regularly downcurved, almost forming a quarter circle; in dorsal view about 3.5 × as long as wide, integument densely punctate, forming 5 longitudinal carinae; antennae inserted at apical ¼ of length in lateral view; head capsule densely punctate in dorsal view, forehead flat; eyes convex, exceeding lateral curve of head capsule in dorsal view; antennal funicle with first segment 1.5 × as long as wide, slightly longer than segments 2–3 together, 3–7 transverse.

**PRONOTUM.** Wider than long (W : L ratio: 1.43), widest in basal ⅔ of length, slightly narrower there than elytra at humeral angles (0.95 ×), side subparallel in basal ⅔, converging apicad in apical ¼, forming blunt tooth beyond middle of length; apical constriction distinct, shallow, less deep than width of scape in middle of length; integument with punctures rounded, space between punctures rough, shiny, micropunctate, generally wider or narrower than diameter of punctures in middle, narrower laterally.

**METATHORAX.** Metanepisterna with recumbent white setae, non-contiguous.

**ELYTRA.** Longer than wide (W : L ratio: 0.76); sides a little convex, widest near middle of length; humeri distinct; apex rounded and jointed at level of suture; striae with punctures about ½ to ⅓ as wide as width of interstriae; interstriae slightly convex; scutellar shield rounded, a little darker than elytra, coated with small recumbent scales not concealing integument.

**ABDOMEN.** Underside uniformly pale brown, with minute recumbent whitish setae, not contiguous. Stridulatory plate with lines of 6–7 granules about ¾ as long as median line from base, space between granules 2–3 × as wide as diameter of a granule. Central sclerotized area isodiametric, rounded at base.

**LEGS.** Profemora moderately thickened near middle of length; protibiae with external margin straight, internal bisinuate, setae on apical half of internal margin recumbent, shorter than length of claws; claws simple.

**Sexual dimorphism**

The female can be distinguished from males by the rostrum which slightly more downcurved basally in lateral view and by the antennae inserted at the apical ⅓ of the length (¼ in the male).
<figure><img src="image_77.png" /><figcaption>**Fig. 29.** Habitus of *Derelomus crypticus* Haran sp. nov. **A–B.** Holotype, ♂ (Kenya) (NHMUK). **C–D.** Paratype, ♀ (Republic of South Africa) (JHAR03078-03, CBGP). **A, C.** Habitus in dorsal view. **B, D.** Habitus in lateral view. Scale bar: A, C = 1 mm; B–D not to scale.</figcaption></figure>
**Life history**

This species was collected on male inflorescences of *Phoenix reclinata*, which probably constitutes the
substrate for the larvae. Adults were collected in May and November.

**Distribution**

In our current knowledge only known from Kenya, and the Western Cape Province of the Republic of South Africa. The population from Western Cape Province is likely an introduction since no palm is indigenous to this area.

**Remarks**

This species is known from very few specimens, it is seemingly rare compared to the other species
found on male inflorescences of *Phoenix reclinata*. The elytral pattern and measurements are probably
more variable than those provided in the description. The male from Kenya and the female from the
Republic of South Africa are quite divergent regarding the shape of their rostrum, beyond what is
expected for sexual dimorphism. Populations from these two areas are probably morphologically slightly
divergent, similarly to what is observed in other species showing such distribution patterns. The penis
and stridulatory plate of the only male available was unfortunately damaged during dissection and could
not be used for the description.

*Derelomus costiger* Marshall, 1958
Fig. 30

*Derelomus costiger* Marshall, 1958: 741.

*Psilocaulus elatus* Richard, 1958: 50. **Syn. nov.**

*Derelomus costiger* – Haran *et al*. 2022a (phylogenetic relationships, life history); 2023a (life history).

**Diagnosis**

In the *D. ephippiger* group, this species can be distinguished by the combination of rostrum comparatively moderately long and downcurved (Fig. 30C-D), only 3.5 × as long as wide in dorsal view in males (Fig. 30A) and by the males lacking patterns of elytra and internal comb of elongate setae on protibiae. *Derelomus costiger* is similar to *D. discus*, but in the latter species males have a longer rostrum (4.75 × as long as wide) and bears combs of setae on protibiae (Fig. 31D). It is also close to *D. ephippiger*, but the latter species exhibit a distinct dark pattern at the base of elytra (Fig. 32A). The body of penis and the stridulatory plate are diagnostic for these species. GenBank accession numbers for the corresponding DNA barcodes: ON553421-22-23/PV698475-76.

**Material examined**

Lectotype of *Derelomus costiger* Marshall, 1958 (here designated)

REPUBLIC OF SOUTH AFRICA • ♂; “Holotype [red label]” “South Africa; Stellenbosch [Western Cape Province, 33°55'55.2" S 18°52'30.0" E]; 9.4.1957; J. G. Theron” “*Derelomus; costiger*. Mshl; Type ♂” “LECTOTYPE ♂; *Derelomus; costiger*; Marshall, 1958; Haran des. 2025”; NHMUK.

Holotype of *Psilocaulus elatus* Richard, 1958

COMORO ISLANDS • ♂; “Moheli; Fomboni[12°17'06.5" S 43°44'38.9" E]; XI.55-A.R” “Institut; Scientifique: Madagascar” “TYPE[red label]” “*Psilocaulus; elatus* n; type; Det. R. Richard” “*Derelomus; costiger* Mshl; Haran det. 2024”; MNHN.
**Paralectotypes**

REPUBLIC OF SOUTH AFRICA – *Western Cape Province* • 1 ♂; Stellenbosch; 33°56′20.4″ S, 18°52′08.4″ E; 9 Apr. 1957; J. G. Theron coll.; CIE. n° 15595; NHMUK • 3 ♂♂; same data as for preceding; SAMC.

**Other material**

ANGOLA • 1 ♂; no precise locality; 7 Jun. 1928; NHMUK • 1 ♂, 1 ♀; Huila district; 14°47′53.2″ S, 14°54′10.8″ E; Mus. Murray; NHMUK.

IVORY COAST • 1 ♂; Toumodi; 6°33′21.6″ N, 5°01′42.0″ W; 5 Oct 1963; R. Vuattoux coll.; flower of *Phoenix* sp.; MNHN.

KENYA • 2 ♂♂, 2 ♀♀; Makindu; 2°16′37.2″ S, 37°49′08.4″ E; May 1937; Mac Arthur coll.; NHMUK.

MOZAMBIQUE • 1 ♀; Maputo Special Reserve, west gate; 26°30′14.4″ S, 32°43′05.0″ E; 21–30 Nov. 2016; Aristophanous M, Cristovae J Laszio G and Miles W coll.; light trap; NHMUK.

REPUBLIC OF SOUTH AFRICA – *Western Cape Province* • 1 ♂, 1 ♀, 8 specs (preserved in ethanol); Stellenbosch, J. Marais Nature reserve; 33°55′55.2″ S, 18°52′30.0″ E; 18 Jul. 2018; J. Haran coll.; on flowers of *Buddleja auriculata* (Buddlejaceae); JHAR01181; CBGP • 1 ♀, 11 specs (preserved in ethanol); same data as for preceding; JHAR01182; CBGP • 1 ♂, 1 ♀; Pringle Bay; 34°21′00.0″ S, 18°49′48.0″ E; 28 Nov. 2019; J. Haran coll.; on male inflorescence of *Phoenix reclinata*; JHAR03078-01/04; CBGP. – *Eastern Cape Province* • 1 ♂, 2 ♀♀; Kentani; 32°30′29.8″ S, 28°19′37.2″ E; H.P. Abernethy coll.; SAM-COL-A050313; SAMC • 1 ♂, 3 ♀♀; Mbotyi, 31°27′10.8″ S, 29°44′49.2″ E; Dec. 2009; Ş. Procheş coll.; on male and female inflorescences of *Phoenix* sp.; JHAR07434; CBGP. – *Gauteng Province* • 1 ♂, 1 ♀; Pretoria; 25°46′23.2″ S, 28°12′32.4″ E; 22 Jun. 1990; S. Neser coll.; in flowers of *Phoenix reclinata*, SANC. – *KwaZulu-Natal Province* • 4 ♂♂, 1 ♀, 30 specs (preserved in ethanol); St Lucia; 28°22′59.8″ S, 32°24′33.4″ E; 5 Jan. 2019; J. Haran coll.; male inflorescence of *Phoenix reclinata*; JHAR02041-0½042/2044; CBGP • 1 ♂, 1 ♀, 7 specs (preserved in ethanol); iSimangaliso Wetland Park, Ozabeni, 3km SSW Sodwana Camp; 27°38′10.8″ S, 32°34′55.2″ E; 22 Oct. 2014; E Grobbelaar coll.; SANC-MCOL-0374/JHAR02335; SANC. – *Limpopo Province* • 1 ♂; Mmabolela Estate; 22°37′52″ S, 28°16′34″ E; 20–24 Nov. 1994; F. Genier coll.; savana light trap; CMNC.

REUNION ISLAND (FRANCE) • 1 ♂, 3 ♀♀; Sainte Marie; 20°53′53.2″ S, 55°32′56.4″ E; Feb. 2022; C. Lemagnen coll.; UV light trap in a garden; JHAR07813; EL.

SENEGAL • 6 ♂♂, 30 ♀♀; Kayar; 14°54′57.6″ N, 17°07′08.4″ W; 8 Apr. 1951; Villiers coll.; in flowers of *Phoenix dactylifera*; MNHN • 3 ♂♂; same collection data as for preceding; NHMUK.

UNITED STATES OF AMERICA • 1 ♂; Florida, Dade County Kendall, Old Cutler Road; 25°37′55.0″ N, 80°18′39.9″ W; 25 Jul.–1 Aug. 2016; S. Haltbert coll.; air suction trap; CMNC • 1 ♀; Florida, Palm Beach County, Delary Beach; 26°25′56.8″ N, 80°05′52.2″ W; 4 Jul. 2020; V. Golia coll.; porch light; CMNC • 1 ♂; Florida, Palm Beach County, Delary Beach; 26°25′56.8″ N, 80°05′52.2″ W; 9 Nov. 2020; V. Golia coll.; porch light; CMNC.

ZIMBABWE • 1 ♂; Victoria Falls; 17°56′42.0″ S, 25°50′21.6″ E; Aug. 1929; G.A.K. Marshall coll.; NHMUK.

**Redescription (♂)**

MEASUREMENTS. Body length 2.5–2.8 mm.
**COLOR.** Body integument uniformly pale brown or with apical 2/3 of rostrum, club and post ocular area dark brown, some specimens with median line of pronotum darker than rest of integument; dorsum with minute recumbent whitish setae, glabrous in appearance.

**HEAD.** Rostrum as long as pronotum in lateral view, slightly and regularly downcurved; in dorsal view about 3.5 × as long as wide, integument densely punctate, forming 5 longitudinal carinae; antennae inserted at apical 1/5 of length in lateral view; head capsule densely punctate in dorsal view, forehead flat; eyes convex, exceeding lateral curve of head capsule in dorsal view; antennal funicle with first segment 2 × as long as wide, as long as segments 2–3 together, 3–7 transverse.

**PRONOTUM.** Wider than long (W:L ratio: 1.50–1.66), widest near base, a little narrower there than elytra at humeral angles (0.82×), side rounded, converging apicad, forming notch and blunt tooth near middle of length; apical constriction distinct, about as deep as width of scape in middle of length; integument with punctures rounded, space between punctures rough, shiny, micropunctate, wider or narrower than diameter of punctures in middle, narrower laterally.

**METATHORAX.** Metanepisterna with recumbent white setae, non-contiguous.

**ELYTRA.** A little longer than wide (W:L ratio: 0.84–0.92); sides strongly convex, widest near basal 2/5 of length; humeri not distinct from the lateral curvature of elytra; apex rounded, notched or jointed at level of suture; striae with punctures about 1/3 to 1/4 as wide as width of interstriae; interstriae flat, 5 slightly raised on basal 3/5 of length; scutellar shield rounded, generally slightly darker than elytra, coated with small recumbent scales not concealing integument.

**ABDOMEN.** Underside uniformly pale brown, with minute recumbent whitish setae, not contiguous. Stridulatory plate with lines of 7–8 granules about 2/3 × as long as median line from base, space between granules 1–2 × as wide as diameter of a granule. Central sclerotized area slightly longer than wide, truncate at base (Fig. 30F).

**LEGS.** Profemora moderately thickened near middle of length; protibiae with external margin straight, internal bisinuate, setae on apical half of internal margin recumbent, short, not contrasting with the setae of the rest of tibiae; claws simple.

**TERMINALIA.** Body of penis elongate (W:L ratio: 0.23), about 1.5 × as long as apodemes; sides slightly convex in dorsal view, widening regularly from base to apical 1/3, from there narrowing apicad, apex rounded; in lateral view curvature moderate but stronger in basal 1/2 of length, width narrowing regularly from basal 1/2 of length to apex, apex bisinuate (Fig. 30E).

## Sexual dimorphism

Females can be distinguished from males by the rostrum which is slightly longer (1.15 × as long as pronotum in lateral view) and narrower (Fig. 30D). The antennae are inserted at apical 1/3 of rostrum in lateral view (1/4 in males). Elytra of females are also more elongate (W:L ratio: 0.70) than in males (Fig. 30B).

## Variation

This species varies in color geographically. Populations from the Republic of South Africa have the rostrum, the post-ocular area and the club dark brown while those from west Africa (Ivory Coast and Senegal) are uniformly pale brown. These variations are not associated with divergent external and internal morphology in specimens examined.
<figure><img src="image_78.png" /><figcaption>**Fig. 30.** Habitus and life history of *Derelomus costiger* Marshall, 1958. **A, C, E–F.** Paralectotype, ♂ (Republic of South Africa) (SAMC). **B, D.** ♀ (Republic of South Africa) (JHAR01182-02, CBGP). **A–B.** Habitus in dorsal view. **C–D.** Habitus in lateral view. **E.** Penis in dorsal (left) and lateral (right) views. **F.** Stridulatory plate. **G–H.** Adults in natura on inflorescence of *Phoenix reclinata* Jacq. in the Republic of South Africa. Scale bar: A–D=1 mm; E–H not to scale.</figcaption></figure>
**Life history**

Adults of this species were collected in various sites on male inflorescences of *Phoenix reclinata* (Arecaceae), which probably constitutes the substrate for the larvae (Fig. 30H). It has been sporadically collected on flowers of *Buddleja auriculata* Benth. (Scrophulariaceae). Adults are attracted by UV lights. This species is the most abundant among those found on male flowers of *P. reclinata*, at least in the Republic of South Africa. Adults were collected all year round.

**Distribution**

In our current knowledge, this species is known from Angola, Comoros and La Reunion Islands, Ivory Coast, Kenya, Mozambique, the Republic of South Africa, Senegal, USA (Florida) and Zimbabwe, but its host is widely distributed in eastern and western Africa, and it is likely that this species is more widespread. In La Reunion Island, this species has only been collected since 2010, suggesting a rather recent introduction on this island (E. Lemagnen pers. com.). This species has also been introduced to North America in recent years.

**Remarks**

*Derelomus costiger* was described based on a series of 6 male and 3 female specimens without designation of a holotype (Marshall 1958). In the collection housed at NHMUK, the type series from Stellenbosch was located. A male of this series bearing a red type label is designated as the lectotype for *Derelomus costiger* Marshall, 1958 [here designated] and was relabelled accordingly. The rest of the series was labelled as paralectotypes. *Psilocaulus elatus* Richard, 1958 is a derelomine weevil described from Comoros Islands a few months after Marshall's description of *D. costiger*. The examination of the holotype of this species revealed no difference with the lectotype and paralectotypes of *D. costiger*. As a result, the species name *Psilocaulus elatus* Richard, 1958 is proposed as a junior synonym of *Derelomus costiger* Marshall, 1958 [new synonymy].

*Derelomus discus* Haran sp. nov.

urn:lsid:zoobank.org:act:709DB3EC-E923-453D-BD88-C1C31004CF04

Fig. 31

**Diagnosis**

In the *D. ephippiger* group, this species can be distinguished by the uniformly pale brown pronotum and elytra in the males, their shape almost forming a disc, and the comb of erect setae on the protibiae (Fig. 31D). It is closest to *D. ephippiger* and *D. costiger*, but the rostrum is longer than in those two species which also lack a comb on the protibiae in the males. The body of the penis and the stridulatory plate are diagnostic in all these species. Females are unknown.

**Etymology**

This species is named in reference to the very wide elytra in male, almost forming a disc with the pronotum.

**Material examined**

**Holotype**

REPUBLIC OF SOUTH AFRICA – **Limpopo Province** • ♂; “SOUTH AFRICA, Venda; Nwanedi Waterfall; 22.38S 30.24E [22°37′34.8″ S 30°24′21.6″ E]; 8.ii.1994 R. Oberprieler” “collected from; *Phoenix reclinata*; (ARECACEAE)” “National coll.; of insects; Pretoria, S. Afr” “Holotype ♂; *Derelomus discus*; Haran, 2025”; SANC.
**Paratype**

REPUBLIC OF SOUTH AFRICA – **Limpopo Province** • 1 ♂; Hans Merensky Nature Reserve; 23°41′42.0″ S, 30°40′04.8″ E; 23–25 Jan. 1987; R. Oberprieler coll.; collected on leaves of *Phoenix reclinata*; ANIC.

**Description (♂)**

**MEASUREMENTS.** Body length 3.2 – 3.3 mm.

**COLOR.** Body integument uniformly pale brown, apical ¾ of rostrum and club dark brown, post ocular area slightly darker than rest of head capsule; dorsum with minute recumbent whitish setae, glabrous in appearance.

**HEAD.** Rostrum a little longer than pronotum in lateral view (1.1 ×), regularly downcurved; in dorsal view about 4.75 × as long as wide, integument densely punctate, forming 5 longitudinal carinae; antennae inserted at apical ½ of length in lateral view; head capsule densely punctate in dorsal view, forehead flat; eyes convex, exceeding lateral curve of head capsule in dorsal view; antennal funicle with first segment 2.5 × as long as wide, as long as segments 2–3 together, 3–7 transverse.

**PRONOTUM.** Wider than long (W : L ratio: 1.85), widest near base, a little narrower there than elytra at humeral angles (0.85 ×), side rounded, converging apicad, forming deep notch near middle of length; apical constriction distinct, about as deep as width of scape in middle of length; integument with punctures rounded, space between punctures rough, shiny, micropunctate, wider or narrower than diameter of punctures in middle, narrower laterally.

**METATHORAX.** Metanepisterna with recumbent white setae, non-contiguous.

**ELYTRA.** A little longer than wide (W : L ratio: 0.92); sides strongly convex, widest near basal ¼ of length; humeri not distinct from lateral curvature of elytra; apex rounded, jointed at level of suture; striae with punctures about ½ to ⅓ as wide as width of interstriae; interstriae flat, 5 slightly raised on basal ⅔ of length; scutellar shield rounded, coated with small recumbent scales not concealing integument.

**ABDOMEN.** Underside uniformly pale brown, with minute recumbent whitish setae, not contiguous. Stridulatory plate with lines of 11–12 granules about as long as median line from base, space between granules 1–2 × as wide as diameter of a granule. Central sclerotized area slightly longer than wide, rounded at base (Fig. 31E).

**LEGS.** Profemora moderately thickened near middle of length; protibiae with external margin straight, internal bisinuate, with a comb of erect setae on apical ½ of length, setae longer than a claw (Fig. 31D); claws simple.

**TERMINALIA.** Body of penis elongate (W : L ratio: 0.33), about as long as apodemes; sides subparallel in basal ⅔, converging apicad in apical ⅓ in dorsal view, apex rounded; in lateral view curvature moderate, stronger in basal ⅓ of length, width narrowing regularly from apical ¼ of length to apex, apex downcurved (Fig. 31C).

**Life history**

Adults were collected on *Phoenix reclinata* (Arecaceae) in January and February.

**Distribution**

This species is known from two male specimens from the Limpopo Province in the Republic of South Africa. Its putative host, *P. reclinata* is widely distributed in Africa.
<figure><img src="image_79.png" /><figcaption>**Fig. 31.** Habitus of *Derelomus discus* Haran sp. nov., holotype, ♂ (Republic of South Africa) (SANC). **A.** Habitus in dorsal view. **B.** Habitus in lateral view. **C.** Penis in dorsal (left) and lateral (right) views. **D.** Protibia with internal comb of erect setae. **E.** Stridulatory plate. Scale bar: A–B=1 mm; C–E not to scale.</figcaption></figure>
*Derelomus ephippiger* Gyllenhal, 1836
Fig. 32

*Derelomus ephippiger* Gyllenhal, 1836: 629.

*Derelomus auberti* Hustache, 1932: 68. **Syn. nov.**

**Diagnosis**

In the *D. ephippiger* group, this species can be distinguished by the combination of a square dark spot at the base of the elytra between interstriae 1–4, the club similar in color to the funicle and the rostrum at most as long as the pronotum in lateral view (Fig. 32B–C). It is closest to *D. discus* sp. nov. and *D. costiger*, but in the latter species, the integument of the elytra is uniformly pale brown in males and the rostrum has a distinct size and curvature (Figs 30C, 31B). The body of the penis and the stridulatory plate are diagnostic in all these species.

**Material examined**

**Lectotypes**

COUNTRY(?) • ♂; "Typus [red label]" "♂" "Barbaria ? [Northern coast of Africa, from Morocco to Lybia, see discussion in the Remarks section]; Quensel [Conrad Quensel, Swedish entomologist]" "NHRS-JLKB000073801" "LECTOTYPE♂; *Derelomus*; *ephippiger*; Gyllenhal, 1836; Des. Haran, 2025"; NHRS.

GABON • ♀; "Museum Paris; Gabon; Collection Léon Fairmaire; 1906" "LECTOTYPE; *Derelomus*; *auberti*; Hust. 1932; Haran des. 2025" "*Derelomus*; *ephippiger*; Gyllenhal, 1836; Haran det. 2025"; MNHN.

**Paralectotypes**

COUNTRY (?) • 1 ♂, 1 ♀; data as for lectotype; NHRS.

GABON • 1 ♀; same collection data as for lectotype of *D. auberti*; MNHN.

**Other material**

NIGERIA • 1 ♂; Jos; 9°54′00.0″ N, 8°51′10.8″ E; 13 Oct. 1955; J. Bechyně coll.; expedition Museum G. Frey Nigeria-Cameroon; NHRS.

SENEGAL • 1 ♀; Kayar; 14°54′57.6″ N, 17°07′08.4″ W; 8 Apr. 1951; Villiers coll.; MNHN • 1 ♂, 1 ♀; same collection data as for preceding; NHMUK.

**Redescription (♂)**

MEASUREMENTS. Body length 2.8–3.0 mm.

COLOR. Body integument pale brown, pronotum and head usually slightly darker than the rest of body, basal ⅜ of interstriae 1–4 dark brown, forming dark rectangle contrasting with rest of elytra; dorsum with minute recumbent whitish setae, glabrous in appearance.

HEAD. Rostrum as long as pronotum in lateral view, slightly and regularly downcurved; in dorsal view about 3 × as long as wide, integument densely punctate, forming 5 longitudinal carinae; antennae inserted near apical ¼ of length in lateral view; head capsule densely punctate in dorsal view, forehead flat; eyes convex, exceeding lateral curve of head capsule in dorsal view; antennal funicle with first segment 2 × as long as wide, as long as segments 2–3 together, 3–7 transverse.
<figure><img src="image_80.png" /><figcaption>**Fig. 32.** Habitus of *Derelomus ephippiger* Gyllenhal, 1836. **A–B, D–E.** Holotype, ♂ (NHRS). **C.** Paralectotype, ♀ (Gabon) (MNHN). **A.** Habitus in dorsal view. **B–C.** Habitus in lateral view. **D.** Penis in dorsal (left) and lateral (right) views. **E.** Stridulatory plate. Scale bar: A–C=1 mm; D–E not to scale.</figcaption></figure>
29. Body wider (Fig. 28A); elytra in males very convex laterally, distinctly wider than width of prothorax. Prothorax wider (W : L ratio: 1.54). Rostrum bicoloured, showing black areas laterally and apically (Fig. 28B–C). Club black. On *Phoenix reclinata* .................................... *D. bivirgatus* Marshall, 1951

– Body narrower (Fig. 29A–B); elytra in males moderately convex laterally, not much wider than width of prothorax. Prothorax narrower (W : L ratio: 1.43–1.50). Rostrum uniformly brown, (Fig. 29B–D), lacking contrasting black areas. Club pale brown, concolorous with funicles segments in females (Fig. 29C). Brown in males (Fig. 29A). On *Phoenix reclinata* .................... *D. crypticus* Haran sp. nov.

30. Elytra in male uniformly pale brown (Fig. 30A). Club dark brown. Stridulatory plate with series of 7 or more granules (Figs 30F, 31E) .................................................................................................................. 31

– Elytra with a large brown scutellar spot, located between 4th interstriae and extending beyond middle of length in both sexes (Fig. 32A). Club and funicle concolorous, pale brown. Stridulatory plate with series of 5–6 granules (Fig. 32E) .................................................................... *D. ephippiger* Gyllenhal, 1836

31. Rostrum in male as long as pronotum in lateral view, slightly and regularly downcurved (30C); in dorsal view about 3.5 × as long as wide. Internal margin of protibiae in male with short and recumbent setae, not contrasting with the rest of the coating. Body of penis bisinuate at apex in lateral view (Fig. 30E). Stridulatory plate with series of 7–8 granules (Fig. 30F). On *Phoenix reclinata* .... .................................................................................................................... *D. costiger* Marshall, 1958

– Rostrum in male longer than pronotum in lateral view, comparatively more downcurved (Fig. 31B); in dorsal view about 4.75 × as long as wide. Internal margin of protibiae in male with a comb of long erect setae, longer than length of claw (Fig. 31D). Body of penis downcurved at apex in lateral view (Fig. 31C). Stridulatory plate with series of 11–12 granules (Fig. 31E). On *Phoenix reclinata* .................................................................................................... *D. discus* Haran sp. nov.

## Molecular analyses

DNA-grade specimens were obtained for 19 of the 32 known species of Afrotropical and Mediterranean *Derelomus*. For COI data alone, the lowest interspecific distance was observed between *D. caldarai* and *D. schoedli* (7.6%), while the highest intraspecific distance was observed in *D. chamaeropis* between the Moroccan and Spanish populations (3.4%). The species concepts recovered by morphology were consistent with the best-fit ML tree inferred in IQ-TREE from the analysis of the concatenated dataset, where all species represented by several individuals were found to be monophyletic (Fig. 33). The genus *Derelomus* was recovered monophyletic, with moderate support (SH-aLRT of 83.4% and uBV of 64%). Within *Derelomus*, interspecific relationships were not highly supported (only four nodes out of 18 have both SH-aLRT ≥80% and uBV values ≥95%); however, nodes leading to specimens belonging to multiple specimens of the same species were generally highly supported (nine nodes out of 12 have both SH-aLRT > 80% and uBV values > 95%).

Molecular species delimitation analyses based on the COI dataset were highly consistent with the morphological concepts of species, as the results of most SD analyses support extant species boundaries (Fig. 34). A potential lumping of species was inferred only once with the ASAP analysis, between *D. pallidus* and *D. prochesi*. Potential intraspecific splits were found twice in the bPTP analysis (for *D. schoedli* and *D. trinotatus*), once in the PTP(ML) analysis (for *D. schoedli*) and three times in the GMYCm analysis (for *D. fasciatus*, *D. peglerae* and *D. prochesi*).

## Discussion

A total of 32 valid *Derelomus* species are recognised in the Afrotropical and Mediterranean regions, 14 of which are described here as new. Five species names were also set in synonymy after comparison with the type material. Based on morphological features and partly on the results of molecular
phylogenetic analyses, *Derelomus* species from the study area can be grouped into four main species groups (*D. signatus*, *D. pallidus*, *D. nigrovariegatus* and *D. ephippiger* groups) which also show a general consistency regarding their host repertoire and distribution range (Table 1). The affinities of three species (*D. trinotatus*, *D. fasciatus* and *D. languidus*) are unclear yet and they are therefore kept as separate species groups. The molecular phylogeny based on nuclear and mitochondrial gene fragments recovered these groups with high support for the *D. signatus* group (three spp. included; SH-aLRT of 98.7% and uBV of 100%) and the *D. pallidus* group (two spp. included; SH-aLRT of 96.4% and uBV of 100%). The *D. nigrovariegatus* and *D. ephippiger* groups were either too sparsely sampled or recovered polyphyletically (though with low support), respectively. The low support at the crown of the phylogenetic tree of *Derelomus* is consistent with a previous inference based on similar dataset but with a larger sampling of genera in Derelomini (Haran *et al.* 2022a). This pattern was not found in related genera analysed with similar datasets (*Elaeidobius* and *Ebenacobius*), which suggests that *Derelomus* may have undergone rapid radiation during its early stages of diversification. The lack of a strong support overall for the inferred tree precludes further interpretation, so future studies using larger molecular datasets and denser species sampling are needed to elucidate the evolution of this genus. Overall, the results of the molecular SD analyses strongly support the species status that are based on morphological concepts. Out of the seven SD methods implemented, lumping was only suggested by the ASAP approach while splitting was only inferred six times with tree-based approaches (bPTP, PTP(ML) and GMYCm). This pattern is expected as distance-based SD methods tend to over-lump while tree-based SD methods are generally known to over-split (Pentinsaari *et al.* 2017; Renner *et al.* 2017; Dellicour & Flot 2018; Le Ru *et al.* 2022; Hévin *et al.* 2024).

The taxonomy of *Derelomus* remains challenging and several species concepts proposed here would benefit from more investigations. This applies to certain taxa in the *D. ephippiger*, *D. fasciatus* and *D. signatus* species groups for which the species status of morphologically slightly divergent populations from eastern and southern Africa was often difficult to establish. The most parsimonious approach was used here (i.e., avoiding species description when based solely on slight morphological divergences), but it is likely that molecular analyses used on a denser sampling of populations will uncover cryptic diversity in these lineages. In the *D. ephippiger* and *D. signatus* groups in particular, many species are known from very few specimens or localities and it is expected that there are more species in these groups than are reported in this work. When studying *Derelomus*, it should also be noted that many species are human-transported with their hosts outside their native range (e.g., *D. piriformis* with *Phoenix canariensis* in the Mediterranean region; *D. costiger*, *D. postfasciatus* and *D. strangulatus* with *P. reclinata* in the Western Cape Province of the Republic of South Africa, etc.), which may result in reconnection of allopatric neighbouring species and the formation of hybrids. A few poorly sclerotized specimens from Kenya with hybrid phenotypes between species of the *D. ephippiger* group suggest that this process may have arisen at least sporadically.

The data assembled in this study revealed some trends in the ecology and evolution of the genus *Derelomus*. A first notable point is the ratio of species with non-palm hosts. Though typically seen as a palm-associated genus (Lepesme 1947; Franz & Valente 2005), we show here that about 40% of *Derelomus* species with known host are in fact associated with other monocot (Strelitziaceae) or dicot families (Achariaceae, Anacardiaceae, Ebenaceae and Fabaceae). This observation confirms the findings of Haran *et al.* (2022a), who revealed that *Derelomus* species frequently shift from palms (Arecaceae) to other plant lineages. In the case of the *D. signatus* group, the shift to *Vachellia* (Fabaceae) has seemingly led to an increase of diversification, a process probably still ongoing owing to the close relatedness of species in this group. Yet, the role of *Derelomus* species in the pollination of their host has only been established for *D. chamaeropis* and *Chamaerops humilis* and for *D. piriformis* and *Phoenix canariensis* (Anstett 1999; Meekijjaroenroj & Anstett 2003). Although weevils generally exhibit niche conservatism at the genus level for such interactions (Haran *et al.* 2023a), detailed ecological studies are needed to
<figure><img src="image_81.png" /><figcaption>**Fig. 33.** Best-fit ML tree of Afrotropical and Mediterranean *Derelomus* Schoenherr, 1825, inferred from nuclear and mitochondrial genes. Node support refers to SH-like approximate likelihood ratio tests (SH-aLRT) and ultrafast bootstrap values (uBV), in that order.</figcaption></figure>
<figure><img src="image_82.png" /><figcaption>**Fig. 34.** Results from molecular species delimitation analyses for the 19 species of *Derelomus* Schoenherr, 1825 with COI sequence available. Grey vertical lines on the right refer to morphological concepts of species and colored lines refer to the results of molecular species delimitation analyses. Blue and red boxes highlight species lumped (L) or splitted (S) respectively for the corresponding analyses.</figcaption></figure>
<table><caption>**Table 1** (continued on next page). Distribution range and host plant associations of the *Derelomus* Shoenherr, 1825 from the Afrotropical and Mediterranean regions. * = larvae bred from tissues of inflorescences.</caption><thead><tr><th>Group</th><th>Species</th><th>Distribution</th><th>Host</th><th>Host fam.</th></tr></thead><tbody><tr><td rowspan="6"><i>D. signatus</i></td><td><i>D. signatus</i> Gyllenhal, 1836</td><td>Ethiopia, Kenya, S Africa, Zambia</td><td><i>Vachellia seyal</i>, <i>V. hockii</i>, <i>V. sp.</i></td><td>Fabaceae</td></tr><tr><td><i>D. karoensis</i> sp. nov.</td><td>S Africa</td><td><i>Vachellia karroo</i></td><td>Fabaceae</td></tr><tr><td><i>D. acuminatus</i> sp. nov.</td><td>Ethiopia, Kenya, S Africa (?)</td><td><i>Vachellia xanthophloea</i></td><td>Fabaceae</td></tr><tr><td><i>D. abyssinicus</i> Hustache, 1936</td><td>Ethiopia, Kenya, Rwanda, Tanzania, Uganda</td><td><i>Vachellia seyal</i>, <i>V. hockii</i>, <i>V. sp.</i></td><td>Fabaceae</td></tr><tr><td><i>D. caldarai</i> sp. nov.</td><td>S Africa, Tanzania</td><td><i>Mimosa sp.</i></td><td>Fabaceae</td></tr><tr><td><i>D. lindbergi</i> Roudier, 1957</td><td>Burkina F., Cameroun, Cape Verde Archipelago, Senegal</td><td><i>Vachellia sp.</i></td><td>Fabaceae</td></tr><tr><td><i>D. pericarti</i> Hoffmann, 1968</td><td>Cameroun, Chad</td><td><i>Vachellia nilotica</i></td><td>Fabaceae</td></tr><tr><td><i>D. schoedli</i> Rheinheimer, 2005</td><td>Ethiopia, Saudi Arabia</td><td><i>Vachellia sp.</i></td><td>Fabaceae</td></tr><tr><td rowspan="5"><i>D. pallidus</i></td><td><i>D. pallidus</i> Fähraeus, 1844</td><td>Angola, Kenya, S Africa, Tanzania, Uganda</td><td><i>Euclea racemosa</i></td><td>Ebenaceae</td></tr><tr><td>= <i>D. pallidus</i> Hartmann, 1904</td><td></td><td></td><td></td></tr><tr><td>= <i>D. hartmanni</i> Klima, 1934</td><td></td><td></td><td></td></tr><tr><td><i>D. setifer</i> sp. nov.</td><td>S Africa</td><td></td><td>–</td></tr><tr><td><i>D. prochesi</i> sp. nov.</td><td>S Africa</td><td><i>Kiggelaria africana</i></td><td>Achariaceae</td></tr><tr><td rowspan="5"><i>D. nigrovariegatus</i></td><td><i>D. strelitziae</i> sp. nov.</td><td>S Africa</td><td><i>Strelitzia nicolai*</i></td><td>Strelitziacae</td></tr><tr><td><i>D. strangulatus</i> sp. nov.</td><td>S Africa</td><td><i>Phoenix reclinata</i></td><td>Arecaceae</td></tr><tr><td><i>D. zonatus</i> Marshall, 1950</td><td>Democratic Republic of Congo</td><td><i>Phoenix reclinata</i></td><td>Arecaceae</td></tr><tr><td><i>D. brunneus</i> sp. nov.</td><td>Democratic Republic of Congo</td><td><i>Phoenix reclinata</i></td><td>Arecaceae</td></tr><tr><td><i>D. ater</i> Marshall, 1950</td><td>Democratic Republic of Congo</td><td><i>Phoenix reclinata</i></td><td>Arecaceae</td></tr><tr><td><i>D. nigrovariegatus</i> Hustache, 1936</td><td>Ethiopia</td><td></td><td>–</td></tr><tr><td><i>D. trinotatus</i></td><td><i>D. trinotatus</i> sp. nov.</td><td>Cameroun, Gabon, Liberia, Nigeria</td><td></td><td>–</td></tr><tr><td><i>D. fasciatus</i></td><td><i>D. fasciatus</i> Hartmann, 1904</td><td>S Africa, Tanzania</td><td><i>Searsia glauca</i></td><td>Anacardiaceae</td></tr><tr><td><i>D. languidus</i></td><td><i>D. languidus</i> Fähraeus, 1844</td><td>Democratic Republic of Congo, S Africa</td><td><i>Phoenix reclinata</i></td><td>Arecaceae</td></tr><tr><td></td><td>= <i>D. rectirostris</i> Hustache, 1937</td><td></td><td></td><td></td></tr></tbody></table>
<table><caption>**Table 1 (continued).** Distribution range and host plant associations of the *Derelomus* Shoenherr, 1825 from the Afrotropical and Mediterranean regions.
* = larvae bred from tissues of inflorescences.</caption><thead><tr><th>Group</th><th>Species</th><th>Distribution</th><th>Host</th><th>Host fam.</th></tr></thead><tbody><tr><td rowspan="10">D. ephippiger</td><td>D. brevis sp. nov.</td><td rowspan="4">Kenya, Tanzania, La Réunion, Mayotte islands</td><td>Cocos nucifera</td><td>Arecaceae</td></tr><tr><td>D. antonioui Alizar, 2007</td><td>Cyprus, Saudi Arabia</td><td>Phoenix canariensis, P. dactylifera</td><td>Arecaceae</td></tr><tr><td>D. postfasciatus Hesse, 1929</td><td>S Africa</td><td>Phoenix reclinata</td><td>Arecaceae</td></tr><tr><td>D. chamaeropis (Fabricius, 1789)<br>= D. subcostatus Boheman, 1844<br>= Ochrinulus antigae Reitter, 1887</td><td>Western Mediterranean region</td><td>Chamaerops humilis*</td><td>Arecaceae</td></tr><tr><td>D. piriformis (Hoffmann, 1938)<br>= D. kocheri Hoffmann, 1957</td><td>Canary Island, Mediterranean region (intr.),<br>USA (intr.)</td><td>Phoenix canariensis*</td><td>Arecaceae</td></tr><tr><td>D. peglerae sp. nov.</td><td rowspan="2">Angola, Ethiopia, Kenya, S Africa, Uganda</td><td rowspan="2">Phoenix reclinata</td><td rowspan="2">Arecaceae</td></tr><tr><td>D. baka sp. nov.</td></tr><tr><td>D. bivirgatus Marshall, 1951</td><td>Cameroon</td><td>Phoenix reclinata</td><td>Arecaceae</td></tr><tr><td>D. crypticus sp. nov.</td><td>Kenya, S Africa</td><td>Phoenix reclinata</td><td>Arecaceae</td></tr><tr><td>D. costiger Marshall, 1958<br>= Psilocaulus elatus Richard, 1959<br>= D. discus sp. nov.</td><td>Kenya, S Africa</td><td>Phoenix reclinata</td><td>Arecaceae</td></tr><tr><td>D. ephippiger Gyllenhal, 1836<br>= D. auberti Hustache, 1932</td><td>Mozambique, Senegal, S Africa, USA (intr.),<br>Zimbabwe<br>S Africa</td><td>Phoenix reclinata</td><td>Arecaceae</td></tr><tr><td>incertae sedis</td><td>D. thalioculus Poinar &amp; Legalov, 2015</td><td>Gabon, Nigeria, Senegal</td><td>–</td><td>–</td></tr><tr><td></td><td></td><td></td><td>Dominican amber</td><td></td></tr></tbody></table>
clarify whether such mutualism is at work with plant lineages outside palms. This study also revealed the prominent role of one palm species, *Phoenix reclinata*, in hosting diverse species of Afrotropical *Derelomus*. At least a third of the species treated here (11 in total) were found associated with this host, on male inflorescences or fruits, either in sympatry or in apparent allopatry. *Phoenix reclinata* is widely distributed in Africa and can be found in a range of ecosystems from coastal habitat up to 2000 m high in the Cameroon volcanic line (T. Couvreur pers. com.). This case merits further investigation into the level of dependence on *Derelomus* species for the pollination of this palm as it would provide a relevant model for studying whether this host uses different locally-adapted pollinators to ensure reproduction across contrasting regions. More generally, only a few palms in the comparatively poor Afrotropical region (Couvreur et al. 2011) appear to be associated with *Derelomus*, a pattern likely reflecting the reality because these plants are well sampled by entomologists due to their economic and ornamental value and their conspicuous inflorescences (Lepesme 1947; Franz & Valente 2005). In all, only five species (*Chamaerops humilis*, *Cocos nucifera*, *Phoenix canariensis*, *P. dactylifera* and *P. reclinata*) are associated at least with one specialised species of *Derelomus* developing in its male inflorescences and thus pollinating or possibly pollinating it. As such, ciophily in palms clearly departs from the one-to-one pollination mutualism usually expected in this kind of system (Herre et al. 2008, but see Cruaud et al. 2012), and the majority of species probably use other systems to ensure their pollination.

A second remarkable point is the distribution of the genus in the study area. *Derelomus* species are mostly recorded in Mediterranean, sub-arid or subtropical regions and are seemingly almost absent from tropical rainforests of Central and West Africa, even though these ecosystems host most of the diversity of African palms (Dransfield *et al.* 2008), the primary hosts for *Derelomus*. This genus is possibly poorly adapted to the conditions in these forests, as suggested by the fact that palms typical of these ecosystems such as the African oil palm (*Elaeis guineensis* Jacq.) are associated with other genera in Derelomini (*Elaeidobius* and *Prosoestus*; Lepesme 1947). Either *Derelomus* is not coping well with tropical wet conditions, or – in case of ciophily – plants in arid or semi-arid biomes are more prone to engage in interactions with this genus.

*Derelomus* species exhibit several morphological features related to their peculiar lifestyle. The males of many species bear a distinct comb of erect setae on the protibiae. Such structures are generally interpreted as pollen-carrying systems to improve pollination in ciophily (de Medeiros & Farrell 2020; Hsiao & Oberprieler 2022). Dimorphism in males also manifests by wide and costate elytra, resulting in a wider and more convex body shape compared to females, which contrasts with what is seen in weevils in general. This apparent reversed dimorphism has led to a misinterpretation of species' concepts in some cases (see the Remarks section under *D. piriformis*). The shape of these enlarged elytra mimics remarkably the structure of the petals of male flowers of the palm hosts of the species exhibiting these features, which probably serve as camouflage to avoid predation. Enlarged elytra can also be interpreted as an adaptation to fighting between males, since they have been observed trying to dislodge and replace other males in copula by lifting them laterally in other Derelomini (*Elaeidobius*; JH obs.). The presence of major and minor males for this feature in several species (*D. antonioui*, *D. chamaeropis*, *D. costiger etc*) tends to confirm this hypothesis. Finally, the stridulatory plates in males showed some degree of variation between closely related species (number and distance between granules) found in sympatry on the same host. These divergences suggest that acoustic communication via stridulation is used by species for intra- and/or inter-specific recognition.

The fossil *Derelomus thalioculus* Poinar & Legalov, from the Dominican amber is distinct from neotropical lineages of “Derelomini” sensu Franz 2006) and agrees in several respects with features of *Derelomus* as currently understood: pronotum with distinct lateral carina (feature absent from neotropical Derelomini), pygidium concealed by elytra, scrobes directed toward eyes and eyes convex (Poinar & Legalov 2015). However, none of these features are specific to *Derelomus* and they are observed at least in two other genera in Derelomini sensu stricto (i.e., *Elaeidobius* and *Ebenacobius*). In addition, the
distinct fovea on forehead and the campanuliform shape of pronotum (Fig. 2; Poinar & Legalov 2015) are features absent in the genus *Derelomus* in its current concept. The fovea is found in *Ebenacobius* and some *Elaeidobius*, and the campanuliform pronotum is typical for *Elaeidobius*. The only distinctive feature enabling to ascertain the placement of this species in *Derelomus* is the narrowing of the base of the penis, but this character is not accessible for amber-preserved specimens, at least with traditional observation methods. Based on this, it is reasonable to set *Derelomus thalioculus* Poinar & Legalov in “Derelomini incertae sedis”, as it might represent an intermediate lineage set between *Elaeidobius* and *Derelomus*.

Two hundred years after the description of the genus by Schoenherr, this work presents an updated taxonomic framework for *Derelomus* in Africa and the Mediterranean region, incorporating new data on host associations. This body of information lays the foundation for the clarification of species complexes, the evolution of the group and the exploration of the pollination ecology of the associated plants.

## Acknowledgments

The following collection managers are acknowledged for providing access to the specimens housed in their institution: Bob Anderson (CMNC), Max Barclay (NHMUK), Johannes Bergsten (NHRS), Lourdes Chamorro (USNM), Hermes Escalona (ANIC), Christoph Germann (NMB), Stéphane Hanot (RMCA), Michael Kuhlmann (ZMK), Rich Leschen (NZAC), Antoine Mantilleri and Hélène Perrin (MNHN), Ruth Müller (TMSA), Mandy Schröter (MTD), Werner Strümpher (TMSA) and Simon van Noort (SAMC). We also thank Roman Borovec, Patrick Weill, Dominique Menet and Emmanuel Lemagnen for providing specimens from their private collections. We acknowledge Rémi Allio (INRAe, France) for support in canopy access and sampling in Gabon. The following services and reserve managers are acknowledged for access and collecting permits: the Western Cape Nature Conservation Board (Permit No. CN44-30-4229; Republic of South Africa), the Cape Research Centre, South African National Parks (Permit No. CRC/2019-2020/012–2012/V1), Ezemvelo KZN Wildlife permits office (OP1382-2019); Paula Strauss and Michael Lutzeyer (Grootbos Private Nature Reserve); Pr. Alfred Ngomanda, CENAREST Gabon (research permit n°AA25/23). This study was supported by recurring funding from Cirad (JMH) and INRAe (GJK).

## References

Alonso-Zarazaga M.A. & Lyal C.H.C. 1999. *A World Catalogue of Families and Genera of Curculionoidea (Insecta: Coleoptera) (Excepting Scolytidae and Platypodidae)*. Entomopraxis, Barcelona.

Alziar G. 2007. Description de *Derelomus antonioui* n. sp. de Chypre et nouvelles données sur la faune de l'île (Coleoptera: Curculionoidea). Contribution à la connaissance de l'histoire naturelle de l'île de Chypre. Coleoptera: Curculionoidea V. *Biocosme Mésogéen* 24 (3): 107-120.

Anstett M-C. 1999. An experimental study of the interaction between the dwarf palm (*Chamaerops humilis*) and its floral visitor *Derelomus chamaeropsis* throughout the life cycle of the weevil. *Acta Oecologica* 20 (5): 551-558. https://doi.org/10.1016/S1146-609X(00)86622-9

Bondar G.G. 1941. Notas Entomológicas da Baía. VII. *Revista de Entomologia* 12: 268-303.

Boheman C.H. 1844. [New taxa]. In: Schoenherr C.J. (ed.) *Genera et species curculionidum, cum synonymia hujus familiae species novae aut hactenus minus cognitae, descriptionibus a Dom. L. Gyllenhal, C.H. Boheman, O. J. Fåhraeus, et entomologiis aliis illustratae*. *Tomus octavus*. – *Pars prima*. Supplementum continens. Roret, Paris [Parisiis] / Fleischer, Leipzig [Lipsiae].

Caldara R. 2013. Curculionidae: Curculioninae. In: Löbl I. & Smetana A. (eds) *Catalogue of Palaearctic Coleoptera, Volume 8*: 51–56, 117–172. Brill, Leiden.
Caldara R., Franz N.M. & Oberprieler R.G. 2014. Curculioninae Latreille, 1802. *In*: Leschen R.A.B. & Beutel R.G. (eds) *Handbook of Zoology. Coleoptera, Beetles - Morphology and Systematics*: 589-628. De Gruyter, Berlin.

Colonnelli E. 2014. Apionidae, Nanophyidae, Brachyceridae and Curculionidae except Scolytinae (Coleoptera) from Socotra Island. *Acta Entomologica Musei Nationalis Pragae* 54: 295-422.
https://doi.org/10.5281/zenodo.5313125

Couvreur T.L.P., Forest F. & Baker W.J. 2011. Origin and global diversification patterns of tropical rain forests: inferences from a complete genus-level phylogeny of palms. *BMC Biology* 9: e44.
https://doi.org/10.1186/1741-7007-9-44

Cruaud A., Cook J., Da-Rong Y., Genson G., Jabbour-Zahab R., Kjellberg F., Pereira R.A.S., Rønsted N., Santos-Mattos O., Savolainen V., Ubaidillah R., van Noort S., Yan-Qiong P. & Rasplus J-Y. 2012. Fig-fig wasp mutualism: the fall of the strict cospeciation paradigm? *In*: Patiny S. (ed.) *Evolution of Plant-Pollinator Interactions*: 68-102. Cambridge University Press, Cambridge.
https://doi.org/10.1017/CBO9781139014113.005

De Cauwer I., Delle-Vedove R., Buatois B., Godé C. & Dufaÿ M. 2025. Striking variation of pollinator attracting scent within a highly specialized pollination system. *Journal of Ecology* 113 (4): 856-868.
https://doi.org/10.1111/1365-2745.14493

Dellicour S. & Flot J.-F. 2018. The hitchhiker's guide to single-locus species delimitation. *Molecular Ecology Resources* 18: 1234-1246. https://doi.org/10.1111/1755-0998.12908

de Medeiros B.A.S. & Farrell B.D. 2020. Evaluating insect-host interactions as a driver of species divergence in palm flower weevils. *Communications Biology* 3: e749.
https://doi.org/10.1038/s42003-020-01482-3

Dransfield J., Uhl N.W., Asmussen C.B., Baker J.W., Harley M.M. & Lewis C.E. 2008. *Genera Palmarum: The Evolution and Classification of Palms*. KewPublishing, Kew.

Dufaÿ M. & Anstett M.C. 2004. Cheating is not always punished: killer female plants and pollination by deceit in the dwarf palm *Chamaerops humilis*. *Journal of Evolutionary Biology* 17 (4): 862-868.
https://doi.org/10.1111/j.1420-9101.2004.00714.x

Dufaÿ M., Hossaert-McKey M. & Anstett M.C. 2002. When leaves act like flowers: how dwarf palms attract their pollinators. *Ecology Letters* 6 (1): 28-34. https://doi.org/10.1046/j.1461-0248.2003.00382.x

Elgueta M. & Marvaldi A.E. 2006. Lista sistemática de las especies de Curculionoidea (Insecta: Coleoptera) presentes en Chile, y su sinonimia. *Boletín Museo Nacional de Historia Natural* 55: 113-153. https://doi.org/10.54830/bmnhn.v55.2006.274

Fabricius J.C. 1798. *Supplementum Entomologiae Systematicae*. Proft et Storck, Hafniae [Copenhagen]. Available from https://www.biodiversitylibrary.org/item/132638#page/5/mode/lup [accessed 31 Jul. 2025].

Fåhraeus O.I. 1844. [New taxa] *In*: Schoenherr C.J. (ed.) *Genera et species curculionidum, cum synonymia hujus familiae species novae aut hactenus minus cognitae, descriptionibus a Dom. L. Gyllenhal, C.H. Boheman, O. J. Fåhraeus, et entomologiis aliis illustratae. Tomus octavus. - Pars prima*. Supplementum continens. Roret, Paris [Parisiis] / Fleischer, Leipzig [Lipsiae]. Available from https://www.biodiversitylibrary.org/item/24772#page/7/mode/lup [accessed 31 Jul. 2025].

Franz N.M. 2001. Description and phylogeny of *Staminodeus*, a new genus of Derelomini (Coleoptera: Curculionidae) associated with Cyclanthaceae. *The Coleopterists Bulletin* 55 (4): 411-432.
https://doi.org/10.1649/0010-065X(2001)055[0411:DAPOSA]2.0.CO;2
Franz N.M. 2003. Systematics of *Cyclanthura* gen. n., a new genus of Derelomini (Coleoptera: Curculionidae). *Insect Systematics and Evolution* 34 (2): 153–198.
https://doi.org/10.1163/187631203788964818

Franz N.M. 2006. Towards a phylogenetic system of derelomine flower weevils (Coleoptera: Curculionidae). *Systematic Entomology* 31 (2): 220–287.
https://doi.org/10.1111/j.1365-3113.2005.00308.x

Franz N.M. 2007a. Reproductive trade-offs in a specialized plant-pollinator system involving *Asplundia uncinata* Harling (Cyclanthaceae) and a derelomine flower weevil (Coleoptera: Curculionidae). *Plant Systematics and Evolution* 269: 183–201. https://doi.org/10.1007/s00606-007-0595-1

Franz N.M. 2007b. Pollination of *Anthurium* by derelomine flower weevils (Coleoptera: Curculionidae). *Revista de Biologia Tropical* 55 (1): 269–271. https://doi.org/10.15517/rbt.v55i1.6079

Franz N.M & O’Brien C.W. 2001a. *Ganglionus*, a new genus of Derelomini (Coleoptera: Curculionidae) associated with *Carludovica* (Cyclanthaceae). *Annals of the Entomological Society of America* 74 (6): 835–850. https://doi.org/10.1603/0013-8746(2001)094[0835:GANGOD]2.0.CO;2

Franz N.M. & O’Brien C.W. 2001b Revision and phylogeny of *Perelleschus* (Coleoptera: Curculionidae), with notes on its association with *Carludovica* (Cyclanthaceae). *Transactions of the American Entomological Society* 127 (2): 255–287.

Franz N.M. & Valente R.M. 2005. Evolutionary trends in derelomine flower weevils (Coleoptera: Curculionidae): from associations to homology. *Invertebrate Systematics* 19 (6): 499–530.
https://doi.org/10.1071/IS05026

Friedman A.L.L. 2006. *Derelomus piriformis* Hoffmann (Curculionoidea: Curculionidae: Curculioninae: Derelomini), a new invasive species in Israel. *Phytoparasitica* 34: 357–359.
https://doi.org/10.1007/BF02981022

Frost S.K. & Frost P.G.H. 1981. Sunbird pollination of *Strelitzia nicolai*. *Oecologia* 49 (3): 379–384.
https://doi.org/10.1007/BF00347603

García Y. Castellanos M.C. & Pausas J.G. 2018. Differential pollinator response underlies plant reproductive resilience after fires. *Annals of Botany* 122 (6): 961–971.
https://doi.org/10.1093/aob/mcy122

GBIF Secretariat: GBIF Backbone Taxonomy. https://doi.org/10.15468/39omei
Available from https://www gbif org/species/5284517 [accessed 15 Nov. 2024].

Guindon S., Dufayard J.F., Lefort V., Anisimova M., Hordijk W. & Gascuel O. 2010. New algorithms and methods to estimate maximum-likelihood phylogenies: assessing the performance of PhyML 3.0. *Systematic Biology* 59 (3): 307–321. https://doi.org/10.1093/sysbio/Syq010

Gyllenhal L. 1836. [New taxa] In: Schoenherr C.J. (ed.) *Genera et species curculionidum, cum synonymia hujus familiae species novae aut hactenus minus cognitae*, descriptionibus a Dom. L. Gyllenhal, C.H. Boheman, et entomologiis aliis illustratae. *Tomus quartus – Pars Secunda*: 601–1121. Roret, Paris [Parisiis]. Available from https://www.biodiversitylibrary.org/item/24769#page/615/mode/1up [accessed 31 Jul. 2025].

Haran J., Beaudoin-Ollivier L., Benoit L. & Kuschel G. 2020. Revision of the palm-pollinating weevil genus *Elaeidobius* Kuschel, 1952 (Curculionidae, Curculioninae, Derelomini) with descriptions of two new species. *European Journal of Taxonomy* 684: 1–32. https://doi.org/10.5852/ejt.2020.684

Haran J., Beaudoin-Ollivier L., Benoit L. & Kergoat G.J. 2021. The origin of an extreme case of sister-species sympatry in a palm-pollinator mutualistic system. *Journal of Biogeography* 48 (12): 3158–3169.
https://doi.org/10.1111/jbi.14273
Haran J., Procheş Ş., Benoit L. & Kergoat G.J. 2022a. From monocots to dicots: host shifts in Afrotropical derelomine weevils shed light on the evolution of non-obligatory brood pollination mutualism. *Biological Journal of the Linnean Society* 137 (1): 15–29. https://doi.org/10.1093/biolinnean/blac069

Haran J., Benoit L., Procheş Ş. & Kergoat G.J. 2022b. *Ebenacobius* Haran, a new southern African genus of flower weevils (Coleoptera: Curculioninae: Derelomini) associated with dicotyledonous plants. *European Journal of Taxonomy* 818: 1–54. https://doi.org/10.5852/ejt.2022.818.1771

Haran J., Kergoat G.J. & de Medeiros B.A.S. 2023a. Most diverse, most neglected: weevils (Coleoptera: Curculionoidea) are ubiquitous specialized brood-site pollinators of tropical flora. *Peer Community Journal* 3: e279. https://doi.org/10.24072/pcjournal.279

Haran J., Li X., Allio R., Shin S., Benoit L., Oberprieler R.G., Farrell B.D., Brown S.D.J., Leschen R.A.B., Kergoat G.J. & McKenna D.D. 2023b. Phylogenomics illuminates the phylogeny of flower weevils (Curculioninae) and reveals ten independent origins of brood-site pollination mutualism in true weevils. *Proceedings of the Royal Society B: Biological Sciences* 290: e20230889. https://doi.org/10.1098/rspb.2023.0889

Hartmann F. 1904. Neue Rüsselkäfer aus Ostafrika. *Deutsche Entomologische Zeitschrift* 1904 (2): 369–419. https://doi.org/10.5281/zenodo.16017081

Hebert P.D.N., Cywinska A., Ball S.L. & deWaard J.R. 2003. Biological identifications through DNA barcodes. *Proceedings of the Royal Society B: Biological Sciences* 270 (1512): 313–321. https://doi.org/10.1098/rspb.2002.2218

Herre A.E., Jandér K.C. & Machado C.A. 2008. Evolutionary ecology of figs and their associates: recent progress and outstanding puzzles. *Annual Review of Ecology, Evolution and Systematics* 39: 439–458. https://doi.org/10.1146/annurev.ecolsys.37.091305.110232

Hesse J.C. 1929. Some new species of Curculionidae from South Africa and Southwest Africa. *Annals of the South African Museum* 25: 475–536.

Hévin N.-M.C., Kergoat G.J., Zilli A., Capdevielle-Dulac C., Musyoka B.K., Sezonlin M., Conlong D., Van Den Berg J., Ndemah R., Le Gall P., Cugala D., Nyamukondiwa C., Pallangyo B., Njaku M., Goftishu M., Assefa Y., Mubenga Kandonda O., Bani G., Molo R., Chipabika G., Ong'amo G., Clamens A.-L., Barbut J., Le Ru B. 2024. Revisiting the taxonomy and molecular systematics of *Sesamia* stemborers (Lepidoptera: Noctuidae: Apameini: Sesamiina): updated classification and comparative evaluation of species delimitation methods. *Arthropod Systematics & Phylogeny* 82: 447–501. https://doi.org/10.3897/asp.82.e113140

Hewitt G. The genetic legacy of the Quaternary ice ages. *Nature* 405: 907–913. https://doi.org/10.1038/35016000

Hoffmann A. 1938a. Description d'un genre nouveau et de nouvelles espèces de Curculionidae de la région Paléarctique. *Bulletin de la Société entomologique de France* 43 (3-4): 46-52. https://doi.org/10.3406/bsef.1938.15241

Hoffmann A. 1938b. Observations diverses. *Bulletin de la Société entomologique de France* 43 (9-10): 107.

Hoffmann A. 1957. Descriptions de Curculionides nouveaux (Col.). *Bulletin de la Société entomologique de France* 62: 77–85. https://doi.org/10.3406/bsef.1957.20337

Hoffmann A. 1958. Faune de France. 62. *Coléoptères Curculionides, vol. 3*. Lechevalier, Paris, 1958.

Hoffmann A. 1968. Coléoptères Curculionides récoltés au Tchad et au Cameroun par mm. J. Gruvel et J. Péricart en 1962-1963. *Bulletin de l'Institut français de l'Afrique noire, série A* 30 (1): 151-187.
Hsiao Y. & Oberprieler R.G. 2022. Taxonomic revision of the genus *Miltotranes* Zimmerman, 1994 (Coleoptera: Curculionidae: Molytinae), the *Bowenia*-pollinating cycad weevils in Australia, with description of a new species and implications for the systematics of *Bowenia*. *Insects* 13 (5): e456. https://doi.org/10.3390/insects13050456

Hustache A. 1932. Curculionides nouveaux de l'Afrique Équatoriale (IIe partie). *Sborník entomologického oddelení Národního muzea v Praze* 10: 28–109.

Hustache A. 1933. Synopsis des curculionides de Madagascar, Premier Supplément. *Mémoires de l'Académie malgache* 15: 5–123.

Hustache A. 1936. Entomological expedition to Abyssinia 1926–27. Coleoptera: Curculionidae. *Annals and Magazine of Natural History* 10 (18): 483–505. https://doi.org/10.1080/00222933608655218

Hustache A. 1937. Curculionides du Congo (3e Série). *Revue de Zoologie et de Botanique africaines* 30: 181–260.

Hustache A. 1939. Deuxième supplément au Synopsis des Curculionides Malgaches. *Bulletin de l'Académie malgache* 21: 1–51.

International Commission on Zoological Nomenclature [ICZN]. 1999. International Code of Zoological Nomenclature. 4th edition. The International Trust for Zoological Nomenclature, London. https://doi.org/10.5962/bhl.title.50608

Jácome-Flores M.E., Delibes M., Wiegand T. & Fedriani J.M. 2018. Spatio-temporal arrangement of *Chamaerops humilis* inflorescences and occupancy patterns by its nursery pollinator, *Derelomus chamaeropsis*. *Annals of Botany* 121: 471–482. https://doi.org/10.1093/aob/mcx177

Kakiopoulos G., Zafeiriou S. & Demetriou J. 2022. First record of *Derelomus piriformis* (Coleoptera: Curculionidae), an alien species associated with palm trees, in Greece. *Natura Croatica* 31 (1):127–131.

Kalyaanamoorthy S., Minh B.Q., Wong T.K.F., von Haeseler A. & Jermiin L.S. 2017. ModelFinder: fast model selection for accurate phylogenetic estimates. *Nature Methods* 14: 587–589. https://doi.org/10.1038/nmeth.4285

Katoh K. & Standley D.M. 2013. MAFFT multiple sequence alignment software version 7: improvements in performance and usability. *Molecular Biology and Evolution* 30: 772–780. https://doi.org/10.1093/molbev/mst010

Klima A. 1934. *Coleopterorum Catalogus*. Vol. 28, Pars 140, Curculionidae: Erirhininae: 1–167. Junk, 's-Gravenhage.

Kojima H. & Morimoto K. 2005. Weevils of the tribe Acalyptini (Coleoptera: Curculionidae: Curculioninae): redefinition and a taxonomic treatment of the Japanese, Korean and Taiwanese species. *ESAKIA* 45: 69–115. https://doi.org/10.5109/2704

Kumar S., Stecher G. & Tamura K. 2016. MEGA7: Molecular Evolutionary Genetics Analysis version 7.0 for bigger datasets. *Molecular Biology and Evolution* 33 (7): 1870–1874. https://doi.org/10.1093/molbev/msw054

Lepesme P. 1947. *Les insectes des palmiers*. Paul Lechevalier, Paris.

Le Ru B., Hévin N., Capdevielle-Dulac C., Musyoka B.K., Sezonlin M., Conlong D., Van Den Berg J., Ndemah R., Le Gall P., Cugala D., Nyamukondiwa C., Pallangyo B., Njaku M., Goftishu M., Assefa Y., Bani G., Molo R., Chipapika G., Ong'amo G., Clamens A.-L., Barbut J., Kergoat G.J. 2022. Phylogenetics, integrative taxonomy and systematics of the *Sesamia cretica* Lederer, 1857 species group (Lepidoptera: Noctuidae: Apameini: Sesamiina), with the description of 23 new species from the Afrotropical region. *Annales de la Société entomologique de France (N.S.)* 58 (5): 387–454. https://doi.org/10.1080/00379271.2022.2113341
Lyal C.H.C. 2020. *Glossary of Weevil Characters*. International Weevil Community Website. Available from [http://weevil.info/glossary-weevil-characters](http://weevil.info/glossary-weevil-characters) [accessed 15 Oct. 2024].

Lyal C.H.C. & King T. 1996. Elytro-tergal stridulation in weevils (Insecta: Coleoptera: Curculionoidea). *Journal of Natural History* 30: 703–773. https://doi.org/10.1080/00222939600770391

Maddison W.P. & Maddison D.R. 2023. Mesquite: a modular system for evolutionary analysis. Version 3.81. Available from [http://www.mesquiteproject.org](http://www.mesquiteproject.org) [accessed 31 Jul. 2024].

Marshall G.A.K. 1928. LXVIII.—New Oriental Curculionidae. *Annals and Magazine of Natural History* 2 (12): 537–557. https://doi.org/10.1080/00222932808672918

Marshall G.A.K. 1935. Four new Derelominae (Col. Curc.). *Stylops* 4 (6): 137–140. https://doi.org/10.1111/j.1365-3113.1935.tb00577.x

Marshall G.A.K. 1950. New Curculionidae (Col.) from the Belgian Congo. *Revue de Zoologie et de Botanique africaine* 43: 260–266.

Marshall G.A.K. 1951. New Curculionidae from Tropical Africa. *Annals and Magazine of Natural History* 12 (4): 833–848. https://doi.org/10.1080/00222935108654215

Marshall G.A.K. 1958. New South African Curculionidae (Col.). *Annals and Magazine of Natural History* 10 (118): 737–751. https://doi.org/10.1080/00222935708656026

Meekijjaroenroj A. & Anstett M.C. 2003. A weevil pollinating the Canary Islands date palm: between parasitism and mutualism. *Naturwissenschaften* 90 (10): 452–455. https://doi.org/10.1007/s00114-003-0454-z

Minh B.Q., Nguyen M.A.T. & von Haeseler A. 2013. Ultrafast approximation for phylogenetic bootstrap. *Molecular Biology and Evolution* 30 (5): 1188–1195. https://doi.org/10.1093/molbev/mst024

Minh B.Q., Schmidt H.A., Chernomor O., Schrempf D., Woodhams M.D., von Haeseler A. & Lanfear R. 2020. IQ-TREE 2: new models and efficient methods for phylogenetic inference in the genomic era. *Molecular Biology and Evolution* 37: 1530–1534. https://doi.org/10.1093/molbev/msaa015

Monaghan M.T., Wild R., Elliot M., Fujisawa T., Balke M., Inward D.J.G., Lees D., Ranaivosolo R., Eggleton P., Barraclough T. & Vogler A.P. 2009. Accelerated species inventory of Madagascar using coalescent-based models of species delineation. *Systematic Biology* 58 (3): 298–311. https://doi.org/10.1093/sysbio/syp027

Morimoto K. 1959. Description of a new species of the genus *Derelomus* Schonherr (Col., Curculionidae), with notes on the biology and immature stages. *Entomological Review of Japan* 10: 46–49.

Pentinsaari M., Vos R. & Mutanen M. 2017. Algorithmic single-locus species delimitation: effects of sampling effort, variation and nonmonophyly in four methods and 1870 species of beetles. *Molecular Ecology Resources* 17 (3): 393–404. https://doi.org/10.1111/1755-0998.12557

Piry S. & Gompel N. 2002. Présence en France de *Neoderelomus piriformis* (Hoffmann, 1938) sur le palmier *Phoenix canariensis* Hort. (Coleoptera, Curculionidae, Derelomini). *Bulletin de la Société entomologique de France* 107 (5): 529–534. https://doi.org/10.3406/bsef.2002.16904

Poinar G. & Legalov A.A. 2015. New Curculioninae (Coleoptera: Curculionidae) in Dominican amber. *Palaeontologia Electronica* 18.1.13A: 1–15. https://doi.org/10.26879/480

Pons J., Barraclough T.G., Gomez-Zurita J., Cardoso A., Duran D.P., Hazell S., Kamoun S., Sumlin W.D. & Vogler A.P. 2006. Sequenced-based species delimitation for the DNA taxonomy of undescribed insects. *Systematic Biology* 55: 595–609. https://doi.org/10.1080/10635150600852011

Puillandre N., Lambert A., Brouillet S. & Achaz G. 2012. ABGD, Automatic Barcode Gap Discovery for primary species delimitation. *Molecular Ecology* 21 (8): 1864–1877. https://doi.org/10.1111/j.1365-294X.2011.05239.x
Puillandre N., Brouillet S. & Achaz G. 2021. ASAP: assemble species by automatic partitioning. *Molecular Ecology Resources* 21: 609–620. https://doi.org/10.1111/1755-0998.13281

Reitter E. 1887. Coleopterologische Notizen. XXII. *Wiener entomologische Zeitung* 6: 104–107. https://doi.org/10.5962/bhl.part.17740

Renner M.A.M., Heslewood M.M., Patzak S.D.F., Schäfer-Verwimp A. & Heinrichs J. 2017. By how much do we underestimate species diversity of liverworts using morphological evidence? An example from Australasian *Plagiochila* (Plagiochilaceae: Jungermanniopsida). *Molecular Phylogenetics and Evolution* 107: 576–593. https://doi.org/10.1016/j.ympev.2016.12.018

Rheinheimer J. 2005. New species of the genera *Derelomus* Schonherr and *Leptopius* Oke (Coleoptera: Curculionidae). *Koleopterologische Rundschau* 75: 383–388.

Richard R. 1958. Curculionides nouveaux ou mal connus des Comores, île Glorieuse, île Europa. *Mémoires de l'Institut scientifique de Madagascar, Entomologie* 10: 41–63.

Roudier A. 1957. Résultats de l’expédition zoologique du Professeur Dr. Hakan Lindberg aux îles du Cap Vert durant l’hiver 1953–54. N° 10. Coléoptères Curculionides nouveaux rapportés des îles du Cap-Vert par le Dr. Hakan Lindberg en 1953–54. *Commentationes Biologicae* 16 (7): 1–20.

Sakai S. 2002. A review of brood-site pollination mutualism: plants providing breeding sites for their pollinators. *Journal of Plant Research* 115: 161–168. https://doi.org/10.1007/s102650200021

Schoenherr C.J. 1825. Continuatio tabulae synopticae familiae curculionidum. *Isis von Oken* 17 (5): 581–588.

Schoenherr C.J. 1844. *Genera et species curculionidum, cum synonymia hujus familiae. Species novae aut hactenus minus cognitae, descriptionibus a Dom. L. Gyllenhal, C.H. Boheman, O.J. Fahraeus, et entomologiis aliis illustratae. Tomus octavus. – Pars prima. Supplementum continens.* Roret, Paris and Fleicher, Leipzig.

Smith S.A. & O'Meara B.C. 2012. treePL: divergence time estimation using penalized likelihood for large phylogenies. *Bioinformatics* 28 (20): 2689–2690. https://doi.org/10.1093/bioinformatics/bts492

Tempère G. & Péricart J. 1989. *Faune de France, Coléoptères Curculionides, 4ᵉ partie. Complément aux trois volumes d'Adolphe Hoffmann. Corrections, additions et répertoire.* Fédération française des Sociétés de Sciences naturelles, Paris.

Veyret P. 1940. Notes éthologiques sur *Derelomus chamaeropis* L. et *D. subcostatus* Boh. (Col. Curlionidae). *ASSNTV* 23: 56–62.

Voss E. 1932. Weitere Curculioniden aus Yunnan und Szetschwan der Sammlung Hauser (Col. Curc.). *Wiener entomologische Zeitung* 49 (4): 285–302.

Zhang J., Kapli P., Pavlidis P. & Stamatakis A. 2013. A general species delimitation method with applications to phylogenetic placements. *Bioinformatics* 29 (22): 2869–2876. https://doi.org/10.1093/bioinformatics/btt499

Printed versions of all papers are deposited in the libraries of four of the institutes that are members of the EJT consortium: Muséum national d'Histoire naturelle, Paris, France; Meise Botanic Garden, Belgium; Royal Museum for Central Africa, Tervuren, Belgium; Royal Belgian Institute of Natural Sciences, Brussels, Belgium. The other members of the consortium are: Natural History Museum of Denmark, Copenhagen, Denmark; Naturalis Biodiversity Center, Leiden, the Netherlands; Museo Nacional de Ciencias Naturales-CSIC, Madrid, Spain; Leibniz Institute for the Analysis of Biodiversity Change, Bonn – Hamburg, Germany; National Museum of the Czech Republic, Prague, Czech Republic; The Steinhardt Museum of Natural History, Tel Aviv, Israël.
Appendix 1

GenBank accession numbers for the gene fragments used. Numbers in bold refer to newly generated sequences in this study (continued on next page).

<table><thead><tr><th>Sequence_ID</th><th>Organism</th><th>ArgK</th><th>COI</th><th>COII</th><th>EF1</th></tr></thead><tbody><tr><td>JHAR03280_0101</td><td>Derelomus antonioui</td><td></td><td>OK188822</td><td></td><td></td></tr><tr><td>JHAR02041_0103</td><td>Derelomus bivirgatus</td><td></td><td><b>PV698441</b></td><td></td><td></td></tr><tr><td>JHAR02041_0105</td><td>Derelomus bivirgatus</td><td></td><td><b>PV698442</b></td><td></td><td></td></tr><tr><td>JHAR02044_0108</td><td>Derelomus bivirgatus</td><td></td><td><b>PV698443</b></td><td></td><td></td></tr><tr><td>JHAR03278_0101</td><td>Derelomus brevis</td><td>ON551420</td><td>ON553436</td><td>ON551428</td><td>ON551455</td></tr><tr><td>JHAR05867_0101</td><td>Derelomus caldarai</td><td></td><td><b>PV698444</b></td><td></td><td></td></tr><tr><td>JHAR03281_0102</td><td>Derelomus cervicalis</td><td></td><td><b>PV698445</b></td><td></td><td>ON551478</td></tr><tr><td>JHAR01504_0101</td><td>Derelomus chamaeropis</td><td>OK181237</td><td>OK188812</td><td>OK283558</td><td>OK283501</td></tr><tr><td>JHAR03220_0102</td><td>Derelomus chamaeropis</td><td></td><td>OK188813</td><td></td><td></td></tr><tr><td>JHAR02984_0101</td><td>Derelomus chamaeropis</td><td></td><td><b>PV698473</b></td><td></td><td></td></tr><tr><td>JHAR04643_0104</td><td>Derelomus chamaeropis</td><td></td><td><b>PV698446</b></td><td></td><td></td></tr><tr><td>JHAR04643_0105</td><td>Derelomus chamaeropis</td><td></td><td><b>PV698447</b></td><td></td><td></td></tr><tr><td>JHAR04643_0106</td><td>Derelomus chamaeropis</td><td></td><td><b>PV698448</b></td><td></td><td></td></tr><tr><td>JHAR04762_0101</td><td>Derelomus chamaeropis</td><td></td><td><b>PV698449</b></td><td></td><td></td></tr><tr><td>JHAR04764_0106</td><td>Derelomus chamaeropis</td><td></td><td><b>PV698450</b></td><td></td><td></td></tr><tr><td>JHAR04764_0107</td><td>Derelomus chamaeropis</td><td></td><td><b>PV698451</b></td><td></td><td></td></tr><tr><td>JHAR04764_0108</td><td>Derelomus chamaeropis</td><td></td><td><b>PV698452</b></td><td></td><td></td></tr><tr><td>JHAR04784_0103</td><td>Derelomus chamaeropis</td><td></td><td><b>PV698453</b></td><td></td><td></td></tr><tr><td>JHAR04784_0104</td><td>Derelomus chamaeropis</td><td></td><td><b>PV698454</b></td><td></td><td></td></tr><tr><td>JHAR04786_0102</td><td>Derelomus chamaeropis</td><td></td><td><b>PV698455</b></td><td></td><td></td></tr><tr><td>JHAR04886_0105</td><td>Derelomus chamaeropis</td><td></td><td><b>PV698456</b></td><td></td><td></td></tr><tr><td>JHAR04886_0106</td><td>Derelomus chamaeropis</td><td></td><td><b>PV698457</b></td><td></td><td></td></tr><tr><td>JHAR04886_0107</td><td>Derelomus chamaeropis</td><td></td><td><b>PV698458</b></td><td></td><td></td></tr><tr><td>JHAR04886_0108</td><td>Derelomus chamaeropis</td><td></td><td><b>PV698459</b></td><td></td><td></td></tr><tr><td>JHAR05699_0101</td><td>Derelomus chamaeropis</td><td></td><td><b>PV698460</b></td><td></td><td></td></tr><tr><td>JHAR05737_0101</td><td>Derelomus chamaeropis</td><td></td><td><b>PV698461</b></td><td></td><td></td></tr><tr><td>JHAR05737_0102</td><td>Derelomus chamaeropis</td><td></td><td><b>PV698462</b></td><td></td><td></td></tr><tr><td>JHAR07851_0101</td><td>Derelomus chamaeropis</td><td></td><td><b>PV698463</b></td><td></td><td></td></tr><tr><td>JHAR07851_0102</td><td>Derelomus chamaeropis</td><td></td><td><b>PV698464</b></td><td></td><td></td></tr><tr><td>JHAR07851_0103</td><td>Derelomus chamaeropis</td><td></td><td><b>PV698465</b></td><td></td><td></td></tr><tr><td>JHAR07851_0104</td><td>Derelomus chamaeropis</td><td></td><td><b>PV698466</b></td><td></td><td></td></tr><tr><td>JHAR07851_0105</td><td>Derelomus chamaeropis</td><td></td><td><b>PV698467</b></td><td></td><td></td></tr><tr><td>JHAR07980_0101</td><td>Derelomus chamaeropis</td><td></td><td><b>PV698468</b></td><td></td><td></td></tr><tr><td>JHAR07980_0102</td><td>Derelomus chamaeropis</td><td></td><td><b>PV698469</b></td><td></td><td></td></tr><tr><td>JHAR07980_0103</td><td>Derelomus chamaeropis</td><td></td><td><b>PV698470</b></td><td></td><td></td></tr><tr><td>JHAR07980_0104</td><td>Derelomus chamaeropis</td><td></td><td><b>PV698471</b></td><td></td><td></td></tr><tr><td>JHAR07980_0105</td><td>Derelomus chamaeropis</td><td></td><td><b>PV698472</b></td><td></td><td></td></tr><tr><td>JHAR01181_0101</td><td>Derelomus costiger</td><td>ON551404</td><td>ON553418</td><td>ON551445</td><td>ON551460</td></tr><tr><td>JHAR03078_0104</td><td>Derelomus costiger</td><td>ON551415</td><td>ON553419</td><td>ON551435</td><td>ON551465</td></tr><tr><td>JHAR03078_0101</td><td>Derelomus costiger</td><td>ON551414</td><td>ON553420</td><td>ON551434</td><td>ON551464</td></tr><tr><td>JHAR02042_0101</td><td>Derelomus costiger</td><td>ON551413</td><td>ON553421</td><td>ON551449</td><td>ON551459</td></tr><tr><td>JHAR01182_0101</td><td>Derelomus costiger</td><td>ON551405</td><td>ON553422</td><td>ON551446</td><td>ON551461</td></tr></tbody></table>
GenBank accession numbers for the gene fragments used. Numbers in bold refer to newly generated sequences in this study (continued).

<table><thead><tr><th>Sequence_ID</th><th>Organism</th><th>ArgK</th><th>COI</th><th>COII</th><th>EF1</th></tr></thead><tbody><tr><td>JHAR02041_0101</td><td>Derelomus costiger</td><td>ON551412</td><td>ON553423</td><td>MZ275338</td><td>MZ275409</td></tr><tr><td>JHAR02041_0106</td><td>Derelomus costiger</td><td></td><td><strong>PV698474</strong></td><td></td><td></td></tr><tr><td>JHAR02041_0107</td><td>Derelomus costiger</td><td></td><td><strong>PV698475</strong></td><td></td><td></td></tr><tr><td>JHAR02335_0101</td><td>Derelomus costiger</td><td></td><td></td><td></td><td>ON551462</td></tr><tr><td>JHAR02335_0102</td><td>Derelomus costiger</td><td>ON551410</td><td></td><td>ON551436</td><td>ON551463</td></tr><tr><td>JHAR03078_0103</td><td>Derelomus crypticus</td><td>ON551416</td><td>ON553424</td><td>ON551432</td><td>ON551466</td></tr><tr><td>JHAR01252_0101</td><td>Derelomus fasciatus</td><td>OK181243</td><td>OK188823</td><td>OK283564</td><td>OK283506</td></tr><tr><td>JHAR01476_0101</td><td>Derelomus fasciatus</td><td>OK181244</td><td>OK283507</td><td></td><td>OK283507</td></tr><tr><td>JHAR01476_0103</td><td>Derelomus fasciatus</td><td></td><td>ON553434</td><td></td><td>ON551475</td></tr><tr><td>JHAR01476_0102</td><td>Derelomus fasciatus</td><td>ON551421</td><td>ON553435</td><td></td><td>ON551476</td></tr><tr><td>JHAR02095_0101</td><td>Derelomus karooensis</td><td>ON551422</td><td>ON553427</td><td>ON551429</td><td>ON551477</td></tr><tr><td>JHAR00920_0101</td><td>Derelomus pallidus</td><td>ON551400</td><td>ON553416</td><td></td><td>ON551474</td></tr><tr><td>JHAR01361_0101</td><td>Derelomus pallidus</td><td>ON551403</td><td>ON553430</td><td>ON551448</td><td>ON551474</td></tr><tr><td>JHAR00820_0102</td><td>Derelomus pallidus</td><td></td><td><strong>PV698476</strong></td><td></td><td></td></tr><tr><td>JHAR00917_0102</td><td>Derelomus pallidus</td><td></td><td><strong>PV698477</strong></td><td></td><td></td></tr><tr><td>JHAR05784_0103</td><td>Derelomus pallidus</td><td></td><td><strong>PV698478</strong></td><td></td><td></td></tr><tr><td>JHAR05784_0104</td><td>Derelomus pallidus</td><td></td><td><strong>PV698479</strong></td><td></td><td></td></tr><tr><td>JHAR02380_0101</td><td>Derelomus pallidus</td><td></td><td></td><td>ON551430</td><td></td></tr><tr><td>JHAR02043_0101</td><td>Derelomus peglerae</td><td>ON551419</td><td>ON553433</td><td>ON551450</td><td>ON551469</td></tr><tr><td>JHAR02043_0102</td><td>Derelomus peglerae</td><td></td><td><strong>PV698480</strong></td><td></td><td></td></tr><tr><td>JHAR02043_0104</td><td>Derelomus peglerae</td><td></td><td><strong>PV698481</strong></td><td></td><td></td></tr><tr><td>JHAR00695_0101</td><td>Derelomus piriformis</td><td>OK181235</td><td>OK188814</td><td>OK283559</td><td>OK283503</td></tr><tr><td>JHAR01180_0101</td><td>Derelomus postfasciatus</td><td>ON551406</td><td>ON553425</td><td>ON551444</td><td>ON551467</td></tr><tr><td>JHAR03078_0102</td><td>Derelomus postfasciatus</td><td>ON551418</td><td>ON553426</td><td>ON551433</td><td>ON551468</td></tr><tr><td>JHAR01174_0101</td><td>Derelomus prochesi</td><td>ON551402</td><td>ON553431</td><td>ON551447</td><td>ON551471</td></tr><tr><td>JHAR01174_0102</td><td>Derelomus prochesi</td><td>ON551417</td><td>ON553432</td><td>ON551431</td><td>ON551472</td></tr><tr><td>JHAR05978_0101</td><td>Derelomus prochesi</td><td></td><td><strong>PV698482</strong></td><td></td><td></td></tr><tr><td>JHAR01036_0101</td><td>Derelomus schoedli</td><td></td><td>OK188815</td><td>OK283555</td><td></td></tr><tr><td>JHAR01036_0102</td><td>Derelomus schoedli</td><td></td><td>OK188816</td><td>OK283556</td><td></td></tr><tr><td>JHAR03077_0103</td><td>Derelomus strangulatus</td><td></td><td>ON553437</td><td>ON551437</td><td>ON551482</td></tr><tr><td>JHAR01561_0101</td><td>Derelomus strelitziae</td><td>ON551407</td><td>ON553428</td><td>ON551443</td><td></td></tr><tr><td>JHAR01572_0101</td><td>Derelomus strelitziae</td><td>ON551408</td><td>ON553429</td><td>ON551442</td><td>ON551479</td></tr><tr><td>JHAR06422_0101</td><td>Derelomus trinotatus</td><td></td><td><strong>PV698483</strong></td><td></td><td></td></tr><tr><td>JHAR06423_0101</td><td>Derelomus trinotatus</td><td></td><td><strong>PV698484</strong></td><td></td><td></td></tr></tbody></table>