Nitidulidae, with almost 4500 known species, is a mid-sized family of the order Coleoptera. Within this taxon, the diverse subfamily Meligethinae comprises about 700 pollen-eating species described worldwide [1–4]. Recently, the classification of Meligethinae underwent several changes in light of both molecular and morphological evidence, restricting the concept of several included genera, particularly in the case of the previously polyphyletic genus Meligethes Stephens, 1830 [2, 5–6]. Meligethes, even as presently bounded [2] [Figs 1A (a), (b)], is still a rather species-rich genus, consisting of more than 60 species predominantly occurring in the Eastern Palaearctic [7–10] (Fig. 1 B-C; Table 1). This group includes species which are all associated with flowers of Rosaceae as larvae [2]. Most of Meligethes are oligophagous, although some members appear strictly monophagous [1, 9–10]. Some locally common Meligethes [e.g. M. atratus (Oliver, 1790), M. flavimanus Stephens, 1830, M. violaceus Reitter, 1873)] also represent economically significant potential pests, attacking blossoms of ornamental roses (Rosa spp.) and plum trees (Prunus spp.) in Europe, Japan, and elsewhere [1, 11–13].
The closely related and purported sister genus Brassicogethes Audisio & Cline, 2009 [Figs 1 A (c), D], comprises instead some forty, mostly Western Palaearctic species, all associated with Brassicaceae [2, 5, 14–23], and a few species [e.g. B. aeneus (F.) and B. viridescens (F.)] represent economically important pests, massively attacking blossoms of oilseed rapes, broccoli, cauliflowers, and others.
Table 1. Summary of the information on all 62 [Meligethes s.str. + Odonthogethes] described species.
specific epithet
|
author(s) and year of description
|
subgenus
|
species group
|
host- plant
|
distribution
|
type(s)
|
notes
|
argentithorax
|
Audisio, Sabatelli & Jelínek, 2015
|
Meligethes
|
auripilis/
binotatus
|
Rosa spp.
|
W and SW China (Qinghai, Sichuan)
|
CAR-MZUR
|
|
atratus
|
(Olivier, 1790)
|
Meligethes
|
atratus
|
Rosa spp.
|
W Palearctic Region, N China, Russia
|
MHNP
|
|
aurantirugosus
|
Audisio, Sabatelli & Jelínek, 2015
|
Odonthogethes
|
aurantirugosus
|
Rubus sp.?
|
W Nepal
|
IZAS
|
female unknown
|
aureolineatus
|
Audisio, Sabatelli & Jelínek, 2015
|
Meligethes
|
auripilis/ binotatus
|
Rosa spp.
|
SW China (Sichuan)
|
NMPC
|
female genitalia unknown
|
auricomus
|
Rebmann, 1956
|
Meligethes
|
auripilis/ binotatus
|
Rosa sp.?
|
SE China (Fujian, Jiangxi)
|
SMF
|
|
aurifer
|
Audisio, Sabatelli & Jelínek, 2015
|
Meligethes
|
auripilis/ binotatus
|
Rosa spp.
|
Central China (Shaanxi, Shanxi)
|
NMPC
|
|
auripilis
|
Reitter, 1889
|
Meligethes
|
auripilis/ binotatus
|
Rosa spp.
|
SW and NW China (Sichuan, Yunnan, Gansu, Shanxi, Shaanxi)
|
unknown
|
|
auropilosus
|
Liu, Yang, Huang, Jelínek & Audisio, 2016
|
Meligethes
|
nepalensis
|
Rosa spp.?
|
SW and Central China (Xizang, Sichuan, Hubei, Shaanxi)
|
IZAS
|
|
aurorugosus
|
Liu, Yang, Huang, Jelínek & Audisio, 2016
|
Odonthogethes
|
aurantirugosus
|
Rubus sp.?
|
W China (Xizang)
|
IZAS
|
female unknown
|
binotatus
|
Grouvelle, 1894
|
Meligethes
|
auripilis/ binotatus
|
Rosa spp.
|
NE India, Nepal, SW China (Yunnan, Sichuan), N Myanmar, Bhutan
|
MHNP
|
|
bourdilloni
|
Easton, 1968
|
Odonthogethes
|
chinensis
|
Rubus sp.?
|
E Nepal
|
BMNH
|
|
brassicogethoides
|
Audisio, Sabatelli & Jelínek, 2015
|
Odonthogethes
|
chinensis
|
Rubus sp.?
|
SW China (Yunnan)
|
NMPC
|
male unknown
|
castanescens
|
Grouvelle, 1903
|
Odonthogethes
|
ferrugineus
|
unknown
|
N India (Darjeeling), SW China (Yunnan)
|
MHNP
|
|
chinensis
|
Kirejtshuk, 1979
|
Odonthogethes
|
chinensis
|
Rubus spp.
|
W and Central China (Xizang, Yunnan, Sichuan, Chongqing, Gansu, Shaanxi, Henan, Hubei)
|
ZIN
|
|
cinereoargenteus
|
Audisio, Sabatelli & Jelínek, 2015
|
Meligethes
|
auripilis/ binotatus
|
Rosa spp.
|
SW China (Sichuan)
|
NMPC
|
|
cinereus
|
Jelínek, 1978
|
Meligethes
|
nepalensis
|
Rosa sp.?
|
Bhutan
|
NHMB
|
|
clinei
|
Audisio, Sabatelli & Jelínek, 2015
|
Meligethes
|
auripilis/ binotatus
|
Rosa sp.?
|
SW China (Yunnan)
|
CAS
|
female unknown
|
cyaneus
|
Easton, 1957
|
Meligethes
|
atratus
|
Rosa sp.?
|
Japan
|
BMNH
|
|
denticulatus
|
(Heer, 1841)
|
Odonthogethes
|
denticulatus
|
Rubus spp.
|
W Palearctic Region, N China, Russia
|
ETHZ
|
|
elytralis
|
Audisio, Sabatelli & Jelínek, 2015
|
Meligethes
|
auripilis/ binotatus
|
Rosa spp.
|
SW China (Sichuan)
|
NMPC
|
|
ferrugineus
|
Reitter, 1873
|
Odonthogethes
|
ferrugineus
|
unknown
|
N India (Sikkim)
|
MHNP
|
|
ferruginoides
|
Audisio, Sabatelli & Jelínek, 2015
|
Odonthogethes
|
ferrugineus
|
Pyracantha sp.
|
Central and SW China (Hubei, Sichuan)
|
NMPC
|
|
flavicollis
|
Reitter, 1873
|
Odonthogethes
|
flavicollis
|
Photinia sp. ?
|
E Russia, Japan, North Korea, SW, SE and Central China (Henan, Zhejiang, Chongqing, Jiangxi, Taiwan)
|
BMNH
|
|
flavimanus
|
Stephens, 1830
|
Meligethes
|
atratus
|
Rosa spp.
|
W Palearctic Region, N China, Russia
|
BMNH
|
|
griseus
|
Jelínek, 1978
|
Meligethes
|
nepalensis
|
Rosa sp.?
|
Bhutan
|
NHMB
|
|
hammondi
|
Kirejtshuk, 1980
|
Meligethes
|
atratus
|
Rosa spp.
|
W and central China (Shaanxi, Sichuan, Shanxi, Henan, Hubei)
|
BMNH
|
|
henan
|
Audisio, Sabatelli & Jelínek, 2015
|
Odonthogethes
|
chinensis
|
Rubus sp.?
|
Central China (Henan)
|
NMPC
|
|
inexpectatus
|
Liu, Huang, Cline, Sabatelli & Audisio, 2017
|
Odonthogethes
|
chinensis
|
Rubus sp.?
|
SW China (Sichuan)
|
NWAU
|
male unknown
|
lloydi
|
Easton, 1968
|
Odonthogethes
|
pectoralis
|
Malus sp.?
|
Nepal, SW China (Yunnan)
|
BMNH
|
|
luteomaculatus
|
Liu, Huang, Cline & Audisio, 2018
|
Odonthogethes
|
chinensis
|
Rubus sp.?
|
Central China (Hubei)
|
NWAU
|
|
luteoornatus
|
Audisio, Sabatelli & Jelínek, 2015
|
Odonthogethes
|
chinensis
|
Rubus sp.?
|
SW China (Yunnan)
|
CAS
|
|
lutra
|
Solsky, 1876
|
Meligethes
|
vulpes
|
Rosa spp.
|
Uzbekistan
|
ZMUM
|
|
macrofemoratus
|
Liu, Yang, Huang, Jelínek & Audisio, 2016
|
Meligethes
|
auripilis/ binotatus
|
Rosa spp.
|
Central China (Ningxia, Shaanxi, Hubei)
|
MHBU
|
|
marmota
|
Audisio, Sabatelli & Jelínek, 2015
|
Meligethes
|
auripilis/ binotatus
|
Rosa sp.?
|
Nepal
|
MHNG
|
|
martes
|
Audisio, Sabatelli & Jelínek, 2015
|
Meligethes
|
vulpes
|
Rosa sp.?
|
SW and N China (Shaanxi, Shanxi, Sichuan)
|
NMPC
|
|
melleus
|
Grouvelle, 1908
|
Meligethes
|
vulpes
|
Rosa spp.
|
Myanmar, N India, N Pakistan, Afghanistan, S Tajikistan, Nepal
|
MNHN
|
|
nepalensis
|
Easton, 1968
|
Meligethes
|
nepalensis
|
Rosa spp.
|
Nepal, N India
|
BMNH
|
|
nigroaeneus
|
Audisio, Sabatelli & Jelínek, 2015
|
Odonthogethes
|
chinensis
|
Rubus sp.?
|
SW China (Yunnan)
|
CAS
|
|
nivalis
|
Audisio, Sabatelli & Jelínek, 2015
|
Meligethes
|
auripilis/ binotatus
|
Rosa sp.?
|
SW China (Xizang, Yunnan, Chongqing)
|
NMPC
|
|
occultus
|
Audisio, Sabatelli & Jelínek, 2015
|
Odonthogethes
|
chinensis
|
Rubus sp.?
|
SW China (Yunnan)
|
NMPC
|
male unknown
|
pallidoelytrorum
|
Chen & Kirejtshuk, 2013
|
Odonthogethes
|
chinensis
|
Rubus sp.
|
SW China (Sichuan)
|
IZAS
|
|
pectoralis
|
Rebmann, 1956
|
Odonthogethes
|
pectoralis
|
Malus sp.
|
S Japan, SW, SE and Central China (Guizhou, Hubei, Fujian, Zhejiang, Taiwan)
|
SMF
|
|
pseudochinensis
|
Audisio, Sabatelli & Jelínek, 2015
|
Odonthogethes
|
chinensis
|
Pyracantha sp.
|
Central China (Chongqing, Shaanxi, Hubei)
|
NMPC
|
|
pseudopectoralis
|
Audisio, Sabatelli & Jelínek, 2015
|
Odonthogethes
|
pectoralis
|
Malus sp.?
|
SW China (Yunnan, Sichuan)
|
NMPC
|
|
sadanarii
|
S.-T. Hisamatsu, 2009
|
Odonthogethes
|
pectoralis
|
Malus sp.?
|
SE China (Taiwan)
|
MNST
|
|
schuelkei
|
Audisio, Sabatelli & Jelínek, 2015
|
Odonthogethes
|
chinensis
|
Rubus sp.?
|
W China (Sichuan, Shaanxi?)
|
NMPC
|
|
scrobescens
|
Chen, Lin, Huang & Yang, 2015
|
Odonthogethes
|
chinensis
|
Rubus sp.?
|
SW China (Sichuan, Hubei, Chongqing)
|
IZAS
|
|
semenovi
|
Kirejtshuk, 1979
|
Meligethes
|
auripilis/ binotatus
|
Rosa spp.
|
SE Russia (Ussuri), Central, SW and NW China (Sichuan, Hubei, Shaanxi)
|
ZIN
|
|
shirakii
|
S. Hisamatsu, 1956
|
Odonthogethes
|
ferrugineus
|
Prunus sp.?
|
S Japan (Kyū-Shū), SE China (Guizhou, Zhejiang, Taiwan)
|
EUMJ
|
|
simulator
|
Audisio, Sabatelli & Jelínek, 2015
|
Odonthogethes
|
Chinensis
|
Rubus sp.?
|
Central China (Gansu, Shaanxi)
|
NMPC
|
|
stenotarsus
|
Audisio, Sabatelli & Jelínek, 2015
|
Meligethes
|
auripilis/ binotatus
|
Rosa sp.?
|
SW China (N Yunnan, Xizang)
|
NKMS
|
|
torquatus
|
Jelínek, 1997
|
Meligethes
|
atratus
|
Rosa spp.
|
SE China (Taiwan)
|
NMPC
|
|
transmissus
|
Kirejtshuk, 1988
|
Meligethes
|
auripilis/ binotatus
|
Rosa spp.
|
SW China (Sichuan, Yunnan)
|
ZIN
|
|
trapezithorax
|
Liu, Huang, Cline & Audisio, 2018
|
Odonthogethes
|
chinensis
|
Rubus sp.?
|
Central China (Hubei)
|
NWAU
|
|
tricuspidatus
|
Liu, Huang, Cline & Audisio, 2018
|
Odonthogethes
|
chinensis
|
Rubus sp.?
|
Central China (Hubei)
|
NWAU
|
male unknown
|
tryznai
|
Audisio, Sabatelli & Jelínek, 2015
|
Meligethes
|
auripilis/ binotatus
|
Rosa spp.
|
SW China (Yunnan/Xizang border)
|
NMPC
|
|
violaceus
|
Reitter, 1873
|
Meligethes
|
atratus
|
Rosa spp.
|
China (Anhui, Shaanxi, Hubei, Zhejiang, Fujian, Guizhou, Jiangxi, Yunnan, Sichuan), SE Russia (Ussuri), Japan
|
BMNH
|
|
volkovichi
|
Audisio, Sabatelli & Jelínek, 2015
|
Meligethes
|
nepalensis
|
Rosa sp.?
|
SW China (Yunnan)
|
CAS
|
|
vulpes
|
Solsky, 1876
|
Meligethes
|
vulpes
|
Rosa spp.
|
Uzbekistan, Kyrgyzstan, Tajikistan, Turkmenistan, NW China (Xinjiang)
|
ZMUM
|
|
wagneri
|
Rebmann, 1956
|
Odonthogethes
|
denticulatus
|
Sorbaria sp.?
|
SE and Central China (Fujian, Zhejiang, Taiwan, E Shaanxi)
|
SMF
|
|
xenogynus
|
Audisio, Sabatelli & Jelínek, 2015
|
Odonthogethes
|
ferrugineus
|
Rubus sp.?
|
SW and central China (Sichuan, Shaanxi)
|
NMPC
|
|
yak
|
Liu, Yang, Huang, Jelínek & Audisio, 2016
|
Meligethes
|
auripilis/ binotatus
|
Rosa sp.?
|
SW China (Sichuan)
|
IZAS
|
female unknown
|
Most species of Meligethes (and several within the related genus Brassicogethes) have been recently analyzed through an integrated approach combining morphological, molecular, and bionomical data on larval ecology (2, 5, 14–25). These contributions were based on morphological and molecular data on adults and on bionomical data of larvae. The thus far available molecular data set for Meligethes and Brassicogethes, including sequences of three mitochondrial and nuclear genes for 23 species (Table 2), allowed us to depict a first phylogenetic scenario of Meligethes and some other related genera, and also provide a framework for understanding the origin of this group of Meligethinae, its evolution on different subfamilies of Rosaceae, and, possibly, the shift of ancestral Brassicogethes to Brassicaceae.
Table 2 Species and specimens of [Meligethes s.str. and Odonthogethes], Brassicogethes and Meligethinus for molecular analysis.
Species
|
Sample ID
|
Localities
|
Meligethes auripilis
|
MAU1_1
|
China- Sichuan, Kangding co
|
Meligethes auripilis
|
MAU1_2
|
China- Sichuan, Kangding co
|
Meligethes auropilosus
|
MAURO1_1
|
China- Hubei, Shennongjia Forest, Shennong Peak area
|
Meligethes binotatus
|
MBI1_1
|
China- Sichuan-Xiangcheng co.
|
Meligethes chinensis
|
4C2
|
China- Chongqing, Shizhu, Huangshui
|
Meligethes elytralis
|
MEL1_1
|
China- Sichuan, Xiangcheng co
|
Meligethes ferruginoides
|
MFE1_1
|
China- Sichuan-Moxi-Yanzigou
|
Meligethes hammondi
|
MHA 1_1
|
China- Hubei, Shennongjia forest, Muyu
|
Meligethes luteomaculatus
|
MLU1_1
|
China- Hubei, Shennongjia Forest, Shennong Peak area
|
Meligethes macrofemoratus
|
MMA1_1
|
China- Hubei, Shennongjia forest
|
Meligethes pallidoelytrorum
|
MPA2_1
|
China-Sichuan-Ganzi-Moxi Town
|
Meligethes pectoralis
|
MPEC1_1
|
China- Guizhou-Tongzi-Louguan-Mts-Wanmuhuahai
|
Meligethes pectoralis
|
MPEC1_2
|
China- Guizhou-Tongzi-Louguan-Mts-Wanmuhuahai
|
Meligethes pseudochinensis
|
4A10
|
China- Hubei, Shennongjia forest, Muyu
|
Meligethes pseudochinensis
|
4C5
|
China- Chongqing, Shizhu, Huangshui
|
Meligethes semenovi
|
2C41
|
China- Tibet, Shannan, Cuona, Gongri
|
Meligethes semenovi
|
2C42
|
China- Tibet, Shannan, Cuona, Gongri
|
Meligethes scrobescens
|
4B7
|
China- Chongqing, Shizhu, Huangshui
|
Meligethes transmissus
|
MTR1_1
|
China- Sichuan, Kangding co
|
Meligethes violaceus
|
MVI1_1
|
China- Zhejiang-Quzhou City-Jiangshan City
|
Meligethes wagneri
|
4C4
|
China- Chongqing, Shizhu, Huangshui
|
Meligethes xenogynus
|
3A31
|
China- Shaanxi, Meixian, Haoping temple
|
Meligethes xenogynus
|
3A32
|
China- Shaanxi, Meixian, Haoping temple
|
Brassicogethes coracinus
|
CR8_1
|
Turkey-Ardahan-road between Göle and Susuz
|
Brassicogethes aeneus
|
BAE13_3
|
Italy- Lazio- Pomezia - Borgo di pratica di mare
|
Brassicogethes aeneus
|
BAE13_4
|
Italy- Lazio- Pomezia - Borgo di pratica di mare
|
Brassicogethes salvan
|
BSA1.1
|
Italy- Piemonte- Mt.Palafrè- Lago inf. Del Frisson
|
Meligethinus peringueyi
|
MEP1_1
|
Mozambique- Maputo,Reserva Especial
|
Meligethinus dolosus
|
MED1_2
|
Mozambique- Maputo,Reserva Especial
|
Although the genus Meligethes was included in previous phylogenetic studies, those were aimed to resolve suprageneric classification using molecular approaches [24–26] and only a few widespread W-Palaearctic species were represented. Herein, we analyzed 69 morphological characters (with 157 character states) of adults (Fig. 5) for all 63 known Meligethes (s.l.) species (including a thus far undescribed species from S China), as well as 11 outgroup species belonging to 7 different related Meligethinae genera, including 3 representative species of Brassicogethes. Five additional bionomical characters (with 12 character states) were also analyzed combining available (published and unpublished) data on Meligethes larval ecology. The present work, then, constitutes the first comprehensive phylogenetic analysis at the species level of this large group, with a focus on E Palaearctic lineages.
The genus Meligethes Stephens, 1830
As recently summarized [7], the genus Meligethes [Figs 1 A (a), (b)] was formally established by Stephens in 1830 [27], based on the type species Nitidula rufipes Marsham, 1802 [present day valid name: Meligethes atratus (Olivier, 1790)]. A new preliminary phylogenetic scenario for Meligethes s.l., previously including a heterogeneous and polyphyletic mixture of taxa, formally comprising more than 500 species worldwide, was recently presented [2, 24–26]. In the former paper [2], which was based on morphology of adults and preliminary molecular data, 22 genera were described as new, and 6 previously recognized subgenera of Meligethes were elevated to generic rank, delimiting Meligethes to some thirty Palaearctic species that utilized Rosaceae as larval host-plants. Following the original descriptions of the few European species [27–30], several new or presumed new species were separately added from the Eastern Palaearctic and northern Oriental Regions by a long series of authors in the time span from 1845 to 1997, including two important (although very preliminary) revisions [31–32] of both purported subgenera. Two Chinese species were added [33–34], before the revision of the whole genus Meligethes [7], as currently delimited [2], and 23 new species (21 from China) have been described. Following this revision [7], a few other new species from China were also added [8–10], and other potential new species are presently under scrutiny based on recently collected Chinese material. The genus, comprising a new species still pending formal description, but considered in the present analysis (M. sp. cfr. pectoralis from S China; Tables 1, and 3-4 in additional files), now includes 63 species in two purported subgenera (Meligethes s.str. and Odonthogethes Reitter, 1871) [7–10] (Table 1).
Most Meligethes (s.l.) diversity occurs in the southern countries of the Eastern Palearctic and in the northern portions of the Oriental Region (Middle Asia, China, Japan, N Indian subcontinent); but China certainly represents the main hot-spot of the genus, ca. 85% of the known species being at least marginally distributed in this country (Figs 1 B–C, and Table 1) [7, 10].
Species belonging to this supposed monophyletic genus are characterized by the following unequivocal autoapomorphic morphological and bionomical characters:
1) temples behind eyes (posterolateral view): with distinct, deep elliptical pit, positioned inside the posterior terminal portion of the antennal grooves (Fig. 5a); or with distinct, shallow, subcircular pit, placed more dorsad, outside antennal grooves, close to the posterior-lateral edge of the eye (Fig. 5b; pits on temples or inside antennal grooves are absent in all other Meligethinae exhibiting not raised notosternal sutures). 2) distal posterior portion of ventral antennal grooves (observed in ventral view): abruptly sloping, markedly delimited, deep, and distinctly wider than median portion (differently shaped in all other Meligethinae exhibiting not raised notosternal sutures). 3) larval development: on Rosaceae.
Several other morphological characters are shared with its supposed sister genus Brassicogethes Audisio & Cline, 2009 (Fig. 1, A (c)) [2, 5, 24, 26], whose included species all develop as larvae on the unrelated plant family Brassicaceae. These characters include, but are not limited to: 1) terminal tarsomere simple, not toothed at base (in all Meligethes s.str.; slightly to strongly toothed at base in Odonthogethes) (Fig. 5k-m); 2) terminal maxillary palpomera long and slender, ca. 3× longer than wide (Fig. 5e); 3) notosternal sutures usually not distinct, even in anterior portion (Fig. 5f), except in 3 species of Meligethes s.str.: M. violaceus, M. torquatus and M. cyaneus (Tables 1 and 3-4 in additional files); 4) protibiae bearing only small, minute and subequal cuticular teeth along outer edge (Figs 1 A (a), (b), (c), 2g); 5) protibiae usually long and slender, up to 4–4.5× longer than wide (Figs 1 A (a), (b), (c), 2g); 6) pronotum scarcely convex, at least partially flattened at sides, with posterior angles almost right or (in most Meligethes s.str.) slightly turned posteriad (Figs 1 A (a), (b), (c); 7) complete absence of circum-ocular furrows ("occipital sulci") when viewed dorsally (Figs 2 (i), (j)); 8) semi-circular arched impressions on both sides of the proximal basal portion of the last abdominal ventrite large and markedly distinct (Fig. 2 (d)); 9) semi-circular arched impressions on both sides of the proximal basal portion of the pygidium arcuately and regularly convergent distad (Fig. 2 (n)); 10) male genitalia with plesiotypic tegminal shape, tegmen characterized in most species by a deep, V-shaped incision (Figs 2 (o)–(r)), similarly exhibited by several other basal Meligethinae genera, e.g., Meligethinus Grouvelle, Micropria Grouvelle, Pria Stephens, Microporum Waterhouse, and Cryptarchopria Jelínek.
Trophic relationships of Meligethes with Rosaceae host-plants
Rosaceae is a middle-sized plant family that includes some 3000 species and a little less than 100 genera in 3 recognized subfamilies (Rosoideae, Spiraeoideae, and Dryadoideae) [35–36]. Although exhibiting a worldwide distribution, Rosaceae are particularly diverse in northern Hemisphere temperate forests, where several genera and species of woody shrubs and small trees are important components of local forest communities. The family is peculiar in producing several different and highly distinctive types of fruits, including economically important edible fruits such as apples, pears, peaches, apricots, prunes, strawberries, cherries, raspberries, and blackberries. Meligethes s.l. (i.e., Meligethes s.str. + Odonthogethes) specialize on several genera and species of Rosaceae, although, differently from the majority of other Meligethinae lineages [2, 37–38], only large shrubs and small trees of the two main subfamilies Rosoideae and Spiraeoideae are utilized [10]. No Meligethes (nor other Meligethinae genera) are, in fact, known to develop as larvae on herbaceous Rosaceae (e.g. widespread and species-rich Potentilla, Fragaria, and Geum), despite species of these same plant genera being commonly used by adults of several different genera and species of Meligethinae as occasional food-plants [1].
Despite limited or incomplete information on the larval host-plants of some species, Meligethes s.l. appear to all be associated with Rosaceae species belonging to the genera Rosa, Rubus, Malus, Prunus, Crataegus, Pyracantha, Sorbaria, and Photynia [1, 7, 10]. A couple of these (Rosa, Rubus) belong to the subfamily Rosoideae [35], whereas others (Malus, Prunus, Crataegus, Pyracantha, Sorbaria, and Photynia) to the subfamily Spiraeoideae. The two most commonly used larval hosts of Meligethes s.l. species are Rosa L. and Rubus L. Rosa, representing the only known larval host of species in the subgenus Meligethes s.str., includes some 200 species worldwide; Rubus, representing the main larval hosts of species in the purported subgenus Odonthogethes, includes at least between 400 and 1000 species [35, 39–41]. Both genera exhibit biodiversity hot-spots in China and neighboring areas, where ca. 100 and more than 200 species respectively are known to occur [42]. A similar pattern occurs in Meligethes s.l., and in its constituent subgenera, species-groups and complexes [1–2, 7]. Due to these above mentioned botanical phylogenetic and biogeographic scenarios, we comprehensively explored the evolutionary trajectories within Meligethes s.l. to elucidate forms of ancient coevolutionary relationships between this group of pollen beetles and their larval hosts.