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Two centuries of Pachyrhynchus (Curculionidae, Entiminae, Pachyrhynchini) research: a comprehensive annotated checklist of taxonomy, species groups, distribution, and ecological insights in Southeast Asia
expand article infoYun Ho, Shan-Min Chen, Ace Kevin S. Amarga§|, Jing-Fu Tsai, Bin-Hong Ho#, Ming-Luen Jeng, Hui-Yun Tseng
‡ National Taiwan University, Taipei, Taiwan
§ Biodiversity Research Center, Academia Sinica, Taipei, Taiwan
| National Taiwan Normal University, Taipei, Taiwan
¶ National Museum of Natural Science, Taichung, Taiwan
# National Sun Yat-sen University, Kaohsiung, Taiwan
Open Access

Abstract

The genus Pachyrhynchus comprises a group of entimine weevils with diverse body coloration, endemic to the islands of Southeast Asia, with the highest diversity of species found in the Philippines. In the past ten years, many new species have been discovered, necessitating an updated list of Pachyrhynchus weevils for future study. This checklist provides the valid names, synonyms, type depository, type locality, and distribution of each Pachyrhynchus species, with ecological information such as host plants and life history. Species are further classified into defined species groups based on recent revisions using morphology and genetic analyses. A region-based analysis across eight major faunal areas highlights spatial patterns in species richness. Since the first Pachyrhynchus species was described 200 years ago, a total of 179 species and 43 subspecies (excluding nominotypical subspecies) have been recognized, with 45.8% of Pachyrhynchus species distributed in the Greater Luzon Faunal Region and 39.1% in Greater Mindanao. However, most species still lack ecological information, indicating the difficulty in collecting these data, and further investigations are needed.

Key words:

Distribution, diversity, Easter egg weevil, host plant, Pachyrrhynchus, synonym, type

Introduction

The genus Pachyrhynchus Germar, 1824 (Curculionidae, Entiminae, Pachyrhynchini) (Alonso-Zarazaga and Lyal 1999) is a group of insular broad-nosed weevils distributed from Indonesia to the Ishigaki and Iriomote islands of Japan, including the Philippines, and the Lanyu (Orchid Island/Botel Tobago) and Ludao (Green Island) islands in southeastern Taiwan. They are a group of flightless beetles characterized by completely fused elytra and a wide variety of elytral coloration, with the majority of species found in the Philippines.

The genus Pachyrhynchus was established by Germar (1824) based on a specimen of P. moniliferus collected from Manila, Luzon Island (Philippines). Members of this genus are characterized by a prominent short and thick rostrum, short antennae, absence of scutellar shield, and fused elytra, resulting in flightlessness. Four researchers, George Robert Waterhouse (United Kingdom), Louis Alexandre Auguste Chevrolat (French), Wilhelm Schultze (Germany), and Karl Maria Heller (Austria), were the most influential authorities in the development of Pachyrhynchus taxonomy. Since the first species was described, Waterhouse and Chevrolat contributed more than 20 species in 1841. Later, Heller and Schultze described more than 70 species during 1899–1934, bringing the total to over 120 species by 1934 (Fig. 1). Heller (1912) published the first monograph on Pachyrhynchus, and categorized Pachyrhynchus weevils into seven groups. Later, Schultze (1923, 1924) expanded the classification into 15 groups both primarily based on coloration patterns, and provided keys to the species within these groups.

Figure 1. 

Cumulative number of species published since 1824.

The taxonomy of Pachyrhynchus remained poorly studied until 80 years later (Yoshitake 2012). Since 2012, approximately 90 new synonyms and combinations have been proposed, mostly by Hiraku Yoshitake (see Yoshitake 2017a, 2019a, b, 2020; Yoshitake et al. 2019a), Maurizio Bollino (see Bollino et al. 2017), Anita Rukmane-Bārbale and Analyn Anzano Cabras (see Rukmane and Barševskis 2016; Rukmane-Bārbale 2020a). These taxonomic works have enriched knowledge on the diversity of Pachyrhynchus, resulting in 179 described species and 43 subspecies to date. Furthermore, six new species groups have been proposed: the absurdus species group (Rukmane 2017), the amabilis species group (Bollino et al. 2017; Bollino et al. 2020), the atrocyaneus species group (Bollino 2022), the pinorum species group (Rukmane-Bārbale 2020b), the speciosus species group (Rukmane 2019a), and the schoenherri species group (Bollino et al. 2017). These studies used genitalia and the endophallus of the aedeagus as reliable diagnostic characters.

Rukmane (2018a) published a checklist of Pachyrhynchus based on an extensive review of existing literature, and proposed a total of 40 new synonyms. Recently, Pachyrhynchus researchers have not only focused on taxonomic re-evaluation but have also paid more attention to various other topics, including ecology (Hsu et al. 2017; Tseng et al. 2021), color signaling (Tseng et al. 2014; Lee et al. 2018; Hsu et al. 2021), dispersal mechanisms (Ye et al. 2018; Lin et al. 2021), and phylogenetic studies (Tseng et al. 2018; Ye et al. 2022; Van Dam et al. 2023; Chen et al. 2025). Considering the above, new studies and integrated literature reviews have enhanced our understanding of the taxonomy and distribution of Pachyrhynchus species. This study aims to present an updated species checklist of the genus Pachyrhynchus, with synonyms, type localities, depositories, distributions, host plant information, and species group classifications from various approaches.

Materials and methods

Checklist and literature review

The valid names, synonyms, type depositories, type localities, distributions, and species groups of each Pachyrhynchus species are provided. Verification of type depositories is primarily based on the original descriptions, museum collection lists, or personal visits to the depositories. If the type locality cannot be confirmed because the identity of the type specimens (whether they are nominal subspecies or different subspecies) is unverified in the museum collection list; an asterisk (*) is used to indicate uncertainty regarding the depository. Distribution data were obtained by reviewing the literature. Additional information on host plant species, as reported in the literature (citations listed below) or observed by the authors (noted as “This study”) in the wild, is also provided.

The materials we examined are deposited in the following institutions:

NHMUK Natural History Museum, London, United Kingdom

SDEI Senckenberg Deutsches Entomologisches Institut, Müncheberg, Germany

Other materials from the literature or from museum collection lists are also cited, which are deposited in the following institutions and personal collections:

CFS Franco Sandel private collection, Miane, Italy

CMU-MZ Central Mindanao University, University Museum, Zoological Section, Philippines

DHA Derek Helmericks private collection, Anchorage, Alaska, USA

DUBC Daugavpils University, Institute of Life Sciences and Technology, Coleopterological Research Centre, Ilgas, Daugavpils District, Latvia

HUMSHokkaido University Museum, Sapporo, Japan

MBLI Maurizio Bollino private collection, Lecce, Italy

MCKUM Munetoshi Maruyama Collection at the Kyushu University Museum, Fukuoka, Japan

MNHN National Museum of Natural History, Paris, France

MZZC Ming-Zhi Zhao private collection, Zhuhai, China

NHRS Naturhistoriska riksmuseet, Stockholm, Sweden

NIAES Institute for Agro-Environmental Sciences, NARO, Tsukuba, Japan

PCKS Private collection of Kaoru Sakai, Tokyo, Japan

PNMNH Philippine National Museum of Natural History, Manila, Philippines

SMTD Senckenberg Naturhistorische Sammlungen, Museum für Tierkunde, Dresden, Germany

UMCRC University of Mindanao Coleoptera Research Center, Davao City, Philippines

UPLB University of the Philippines Los Baños, Los Baños, Philippines

ZMHB Naturkundemuseum Berlin (former Zoologische Museum der Humboldt-Universität), Berlin, Germany

Distributional analysis

Based on geographical distribution of islands, we divided the species distribution into eight regions, including the four Pleistocene Aggregate Island Complexes (PAICs) faunal regions (Heaney 1986; Brown and Diesmos 2002; Brown et al. 2013), Indonesian archipelago, islands north of mainland Luzon, Lubang Island and Sibuyan Island (Fig. 2). These regions are defined as follows:

Figure 2. 

Species distribution of Pachyrhynchus weevils. Different colors on the map represent the defined regions. The blank islands of the Philippine archipelago indicate areas that do not belong to any of the four PAICs. Some Pachyrhynchini but no Pachyrhynchus species are found in the Palawan region, marked with a red star. The map was generated using the ggplot2 package in R (Wickham 2016). The bar plot shows the percentage of species distribution across different regions. The numbers in the bars represent the number of species or subspecies (excluding nominotypical subspecies).

  1. Greater Luzon PAIC: Luzon Island, Polillo Island, Catanduanes Island, Marinduque Island
  2. Greater Mindoro PAIC: Mindoro Island
  3. Greater Negros-Panay PAIC: Panay Island, Negros Island, Cebu Island, Masbate Island
  4. Greater Mindanao PAIC: Samar Island, Leyte Island, Bohol Island, Biliran Island, Dinagat Island, Siargao Island, Basilan Island, Bucas Grande Island
  5. Lubang Island
  6. Sibuyan Island
  7. Islands north of mainland Luzon: Ishigaki Island, Iriomote Island, Lanyu Island, Ludao Island, Batanes Islands, Babuyan Islands
  8. Indonesian archipelago: Maluku Islands and West Papua Province with its adjacent islands of Indonesian Papua

Results

The genus Pachyrhynchus currently comprises 179 described species and 43 subspecies. The results revealed that Pachyrhynchus is distributed in Japan (1 species, 0.6%), Taiwan (6 species, 3.4%), the Philippines (168 species, 93.9%), and Indonesia (7 species, 3.9%).

Order Coleoptera Linnaeus

Suborder Polyphaga Emery

Infraorder Cucujiformia Lameere

Superfamily Curculionoidea Latreille

Family Curculionidae Latreille

Subfamily Entiminae Schöenherr

Tribe Pachyrhynchini Schöenherr

Genus Pachyrhynchus Germar, 1824

= Sphaerogaster Dejean, 1821

= Somatodes Schoenberr, 1823

= Sphoerogaster Latreille, 1825

= Sphaerogaster Sturm, 1826

= Sphoenogaster Bertbold, 1827

= Pachirhinus Latreille, 1828

= Pochyrhynchus Laporte, [1840]

= Pachyrhinchus Desmarest, [1842]

= Pachyrhincus Desmarest, [1842]

= Pachyrrhynchus Gemminger & Harold, 1871

Pachyrhynchus amabilis Schultze, 1922

Type depository. SMTD (holotype).

Type locality. Philippines (Mindanao Is.: Bukidnon Prov., Lindaban).

Distribution. Philippines: Mindanao Is. (Bukidnon, Lindaban, Intavas; Zamboanga; Davao City; Lanao; Cotabato; Mt. Kalatungan) (Schultze 1922a, 1924; Rukmane 2019b; Rukmane-Bārbale and Cabras 2021; Cabras et al. 2022a).

Remarks. Two individuals were collected from the Bracken fern, Pteridium aquilinum (Dennstaedtiaceae) in Davao City (Cabras et al. 2022a), suggesting it as a potential host plant.

Pachyrhynchus anellifer Heller, 1912

Pachyrhynchus annulatus Behrens, 1887: 256. Synonymized by Schultze (1924: 318).

Type depository. SMTD (holotype).

Type locality. Philippines (Luzon Is.: Benguet Prov., Irisan River).

Distribution. Philippines: Luzon Is. (Benguet, Baguio, Mt. Santo Tomas; Ifugao; Nueva Vizcaya) (Schultze 1924; Rukmane 2019b, c; Rukmane-Bārbale and Cabras 2021).

Remarks. The conspecificity of P. annulatus Behrens, 1887 and P. anellifer Heller, 1912 was recognized by Schultze (1924). Although the former name has priority, it is preoccupied by P. annulatus Chevrolat, 1881, rendering it permanently invalid. Consequently, the valid name of the species is P. anellifer Heller, 1912.

Pachyrhynchus anichtchenkoi Rukmane & Barševskis, 2016

Type depository. DUBC (holotype, paratypes).

Type locality. Philippines (Mindanao Is: Bukidnon Prov., Mt. Kalatungan).

Distribution. Philippines: Mindanao Is. (Agusan; Bukidnon, Mt. Kalatungan, Intavas, Cabanglasan, Mt. Kiamo; Davao de Oro; Compostela; Lanao del Sur; Sarrangani, Kiamba; Zamboanga) (Cabras et al. 2017; Rukmane 2019b; Rukmane-Bārbale and Cabras 2021).

Pachyrhynchus annulatus Chevrolat, 1881

Type depository. Unknown.

Type locality. Philippines (Chevrolat 1881a).

Distribution. Philippines: Luzon Is. (Benguet, Mt. Pulog; Ifugao; Mountain Prov.) (Schultze 1924; Rukmane 2019b).

Remarks. Pachyrhynchus annulatus Behrens, 1887 is a junior primary homonym of P. annulatus Chevrolat, 1881; therefore, it is permanently invalid.

Pachyrhynchus antonkozlovi Rukmane & Barševskis, 2016

Type depository. DUBC (holotype, paratypes).

Type locality. Philippines (Mindanao Is.: Sarangani province, Malungon).

Distribution. Only known from the type locality.

Pachyrhynchus apicatus Schultze, 1922

Type depository. SMTD (syntypes).

Type locality. Philippines (Polillo Is.).

Distribution. Philippines: Luzon Is. (Aurora, Dingalan, Sierra Madre); Marinduque Is.; Mindoro Is. (Mt. Halcon); Polillo Is. (Schultze 1922b; Schultze 1924; Háva and Rukmane 2018; Rukmane 2019b, 2019c; Rukmane-Bārbale and Cabras 2021).

Pachyrhynchus apocyrtoides Schultze, 1922

Type depository. SMTD (holotype).

Type locality. Philippines (Mindanao Is.: Bukidnon Prov., Lindaban).

Distribution. Philippines: Mindanao Is. (Agusan; Bukidnon; Cotabato; Davao del Sur; Surigao del Norte) (Schultze 1922a; Schultze 1923; Rukmane-Bārbale and Cabras 2021).

Pachyrhynchus apoensis Yoshitake, 2012

Pachyrhynchus pseudapoensis Rukmane & Barševskis, 2016: 94. Synonymized by Yoshitake (2016a: 197).

Type depository. NIAES (holotype, paratypes); PCKS (paratypes).

Type locality. Philippines (Mindanao Is.: Mt. Apo).

Distribution. Philippines: Mindanao Is. (Bukidnon; Cotabato; Davao del Sur, Mt. Apo; Sarrangani) (Cabras and Yoshitake 2016; Rukmane-Bārbale and Cabras 2021; Cabras et al. 2022a).

Host plants. Impatiens platypetala (Balsaminaceae), Melastoma sp. (Melastomaceae), Diplazium davaoense (Woodsiaceae) and Saurauia sp. (Actinidiaceae) (Cabras and Yoshitake 2016)

Remarks. Pachyrhynchus apoensis belongs to the P. schoenherri species group and conforms to the original described characters and distribution to the P. schoenherri species group (Bollino et al. 2017). Rukmane and Barševskis (2016) described P. pseudapoensis, based on a single female specimen from Luzon, which was treated as a synonym as P. apoensis by Yoshitake (2016a). The locality “North Luzon” may need to be reevaluated, as P. apoensis and its related species are known only from Mindanao and its adjacent islands (Yoshitake 2016a).

Pachyrhynchus ardentius Schultze, 1919

Type depository. SMTD (holotype).

Type locality. Philippines (Siargao Is.).

Distribution. Philippines: Siargao Is.; Mindanao Is. (Bukidnon, Cotabato, Davao del sur, Sarangani; Mt. Apo) (Schultze 1924; Rukmane 2019b; Rukmane-Bārbale and Cabras 2021).

Pachyrhynchus argus Pascoe, 1873

Type depository. NHMUK (holotype).

Type locality. Philippines.

Distribution. Philippines: Luzon Is. (Benguet, Baguio, Mt. Santo Tomas, Trinidad, Mt. Trail, Pauai, Mt. Polis, Mt. Pulog; Manila) (Schultze 1924; Rukmane 2018a, 2019b, c).

Pachyrhynchus atrocyaneus Schultze, 1922

Type depository. SMTD (syntypes).

Type locality. Mindanao (Zamboanga Prov.: Zamboanga).

Distribution. Philippines: Mindanao Is. (Zamboanga) (Schultze 1922a; Schultze 1924; Rukmane-Bārbale and Cabras 2021).

Pachyrhynchus atronitens Yoshitake, 2019

Type depository. NIAES (holotype, paratypes); KUM (paratypes).

Type locality. Philippines (Luzon Is.: Kalinga Prov., Pinukpuk).

Distribution. Philippines: Luzon Is. (Abra; Kalinga) (Yoshitake 2019c; Rukmane-Bārbale and Cabras 2021).

Pachyrhynchus baluganus Schultze, 1924

Type depository. SMTD (holotype).

Type locality. Philippines (Luzon Is.: Rizal Prov., Mount San Isidro).

Distribution. Philippines: Luzon Is. (Quirino; Rizal) (Rukmane-Bārbale and Cabras 2021).

Pachyrhynchus banglas Bollino, Sandel & Rukmane, 2017

Type depository. SMTD (holotype); MBLI, DUBC, NIAES, KUM, CFS (paratypes).

Type locality. Philippines (Mindanao Is.: Bukidnon Prov., Kabanglasan).

Distribution. Philippines: Mindanao Is. (Bukidnon, Kabanglasan, Cabanglasan, San Fernando, Intavas, Malaybalay; Summit Mountain Prov.) (Bollino et al. 2017).

Pachyrhynchus barsevskisi Rukmane, 2016

Type depository. DUBC (holotype, paratypes).

Type locality. Philippines (Luzon Is.: Aurora Prov., Dingalan).

Distribution. Philippines: Luzon Is. (Aurora, Dingalan, Ditumabao, Casiguran; Sierra Madre; Nueva Ecija, Caranglan; Quirino, Madela; Nueva Vizcaya, Alfonso Castañeda, Belance) (Rukmane-Bārbale 2020b; Rukmane-Bārbale and Cabras 2021).

Pachyrhynchus basilanus Heller, 1923

Type depository. SMTD (holotype).

Type locality. Philippines (Basilan Is.).

Distribution. Philippines: Basilan Is.; Mindanao Is. (Zamboanga City, Labuan) (Heller 1923a; Bollino 2022).

Pachyrhynchus benguetanus Schultze, 1924

Type depository. SMTD (syntypes).

Type locality. Philippines (Luzon Is.: Benguet Prov.).

Distribution. Philippines: Luzon Is. (Benguet, Ifugao) (Rukmane-Bārbale and Cabras 2021).

Pachyrhynchus bollinoi Rukmane-Bārbale, 2020

Type depository. DUBC (holotype, paratypes).

Type locality. Philippines (Luzon Is.: Mountain Prov., Barlig).

Distribution. Philippines: Luzon Is. (Ifugao, Banaue; Mountain Prov., Barlig; Nueva Vizcaya, Kayapa) (Rukmane-Bārbale 2020c).

Pachyrhynchus bucasanus bucasanus Schultze, 1922

Type depository. SMTD (holotype).

Type locality. Philippines (Bucas Grande Is.: Surigao del Norte Prov.).

Distribution. Only known from the type locality (Schultze 1922a).

Pachyrhynchus bucasanus ornatus Schultze, 1934

Type depository. SMTD (syntypes).

Type locality. Philippines (Samar Is.: Borongan).

Distribution. Only known from the type locality.

Pachyrhynchus cabrasae Rukmane & Barševskis, 2016

Type depository. DUBC (holotype, paratypes).

Type locality. Philippines (Mindanao Is.: Bukidnon Prov., Mt. Kalatungan).

Distribution. Philippines: Mindanao Is. (Agusan; Bukidnon, Mt.

Kalatungan, Cabanglasan, Mt. Kiamo; Davao del Sur) (Cabras et al. 2017; Rukmane-Bārbale and Cabras 2021).

Pachyrhynchus caeruleovittatus Yoshitake, 2012

Type depository. NIAES (holotype, paratypes).

Type locality. Philippines (Mindanao Is.: Mt. Parker).

Distribution. Philippines: Mindanao Is. (Sarangani, South Cotabato)

(Rukmane-Bārbale and Cabras 2021).

Pachyrhynchus caeruleus Yoshitake, 2019

Type depository. KUM (holotype, paratypes).

Type locality. Philippines (Luzon Is.: Aurora Prov., central part of Sierra Madre Mountain Range).

Distribution. Only known from the type locality (Yoshitake 2019d).

Pachyrhynchus cagayanus Heller, 1929

Type depository. SMTD (holotype).

Type locality. Philippines (Luzon Is.: Cagayan Prov., Tuao).

Distribution. Philippines: Luzon Is. (Aurora, Dingalan; Cagayan; Isabela; Kalinga; Quirino) (Rukmane-Bārbale and Cabras 2021).

Pachyrhynchus callainimaculatus Yoshitake, 2017

Type depository. NIAES (holotype).

Type locality. Philippines (Luzon Is.: Cagayan Prov., Sierra Madre).

Distribution. Only known from the type locality (Yoshitake 2017b).

Pachyrhynchus cebrem Patano & Rukmane-Bārbale, 2022

Type depository. CMU-MZ (holotype, paratypes).

Type locality. Philippines (Mindanao Is.: Davao de Oro Prov., Maragusan, Mt. Candalaga).

Distribution. Only known from the type locality.

Pachyrhynchus chamissoi Schultze, 1922

Type depository. SMTD (syntypes).

Type locality. Philippines (Mindanao Is.: Bukidnon Prov., Lindaban).

Distribution. Philippines: Mindanao Is. (Bukidnon, Lindabon); Pollilo Is. (Schultze 1922a; Schultze 1924; Rukmane 2019c).

Pachyrhynchus chlorites Chevrolat, 1881

Pachyrhynchus rutilans Behrens, 1887: 247. Synonymized by Schultze (1924: 327).

Type depository. NHRS (syntypes).

Type locality. Philippines (Chevrolat 1881b).

Distribution. Philippines: Babuyan Is.; Calayan Is.; Luzon Is. (Aurora; Cagayan; Ifugao, Banaue; Ilocos Norte; Isabela; Nueva Vizcaya; Quirino; Manila) (Schultze 1924; Rukmane 2019b; Rukmane-Bārbale and Cabras 2021).

Pachyrhynchus chlorites insularis Kano, 1929

Type depository. Unknown.

Type locality. Taiwan (Taitung County, Lanyu Is. (= Kotosho)).

Distribution. Only known from the type locality.

Host plants. According to our field observations, they feed on Bischofia javanica (Phyllanthaceae) in Lanyu (this study).

Remarks. P. insularis Kano, 1929, from Lanyu was originally considered a distinct species. However, Schultze (1937), after examining specimens of P. chlorites from Calayan with Kano’s description, regarded the Lanyu population as a variety of P. chlorites. According to the ICZN (Article 45.6.4.), a variety proposed before 1961 is automatically treated as subspecies. Whether P. chlorites insularis in Lanyu is a distinct species requires further clarification.

Pachyrhynchus chrysocyaneus Rukmane-Bārbale, 2024

Type depository. DUBC (holotype, paratypes).

Type locality. Philippines (Luzon Is.: Nueva Ecija Prov., Gabaldon).

Distribution. Only known from the type locality.

Pachyrhynchus chrysomaculatus Bollino, 2022

Type depository. SMTD (holotype); MBLI, MCKUM, CFS (paratypes).

Type locality. Philippines (Mindanao Is.: Bukidnon Prov., Cabanglasan).

Distribution. Only known from the type locality.

Pachyrhynchus cinereomaculatus Rukmane-Bārbale, 2020

Type depository. DUBC (holotype, paratypes).

Type locality. Philippines (Luzon Is.: Ifugao Prov., Banaue).

Distribution. Philippines: Luzon Is. (Aurora; Ifugao; Nueva Vizcaya, Kasibu) (Rukmane-Bārbale 2020c).

Pachyrhynchus cingulatus Pascoe, 1873

Type depository. Unknown.

Type locality. Indonesia (Morotai Is.).

Distribution. Only known from the type locality.

Remarks. The type specimen named by Pascoe might be deposited in NHMUK. However, JFT did not find it in the collections.

Pachyrhynchus circulatus Heller, 1912

Type depository. NHMUK (syntype), SMTD (syntype).

Type locality. Philippines (Catanduanes Is.).

Distribution. Philippines: Catanduanes Is. (Schultze 1923; Rukmane 2019b).

Pachyrhynchus circulimaculatus Yoshitake, 2019

Type depository. NIAES (holotype).

Type locality. Philippines (Mindanao Is.: Misamis Occidental Prov.).

Distribution. Philippines: Mindanao Is. (Agusan del Norte, Misamis Occidental) (Yoshitake 2019e).

Pachyrhynchus conformis Yoshitake, 2017

Type depository. NIAES (holotype, paratypes); CFS, MBLI (paratypes).

Type locality. Philippines (Samar Is.: Samar Prov.).

Distribution. Philippines: Samar Is. (Samar, Hinabangan) (Yoshitake 2017c, Rukmane-Bārbale and Cabras 2021).

Pachyrhynchus confusus Schultze, 1923

Type depository. NHMUK (syntypes), SMTD (syntypes).

Type locality. Philippines (Luzon Is.: Los Baños).

Distribution. Philippines: Luzon Is. (Benguet, Mt. Makiling; Laguna, Los Baños) (Rukmane 2019c)

Host plants. Acrostichum aureum (Pteridaceae) (Schultze 1924).

Pachyrhynchus congestus congestus Pascoe, 1873

Pachyrhynchus luteoguttatus Chevrolat, 1881b: 360. Synonymized by Schultze (1924: 324).

Type depository. NHMUK (holotype).

Type locality. Philippines.

Distribution. Philippines: Luzon Is. (Aurora; Benguet, Baguio, Atoc, Mt. Santo Tomas, Mt. Trail, P.I. Atok, Mt. Pawai, Mt. Mirador, Trinidad; Manila; Mountain Prov., Bontoc; Nueva Vizcaya, Kasibu; Ifugao; Quirino; Kallinga) (Schultze 1924; Rukmane 2019b, 2019c; Rukmane-Bārbale and Cabras 2021).

Pachyrhynchus congestus aedamlayroni Rukmane, 2019

Type depository. DUBC (holotype, paratypes).

Type locality. Philippines (Luzon Is.: Ilocos Norte Prov., Adams).

Distribution. Only known from the type locality.

Pachyrhynchus congestus coerulans Kraatz, 1888

Type depository. SDEI (*).

Type locality. Philippines.

Distribution. Philippines: Luzon Is. (Mountain Prov., Bontoc, Mt. Polis; Kalinga) (Schultze 1924; Rukmane 2019c).

Pachyrhynchus congestus immarginatus Kraatz, 1888

Type depository. SDEI (*).

Type locality. Philippines.

Distribution. Philippines: Luzon Is. (Benguet, Mt. Data) (Schultze 1924).

Pachyrhynchus congestus mirabilis Yoshitake, 2017

Type depository. NIAES (holotype, paratypes).

Type locality. Philippines (Luzon Is.: Nueva Vizcaya Prov., Lohong).

Distribution. Philippines: Luzon Is. (Cagayan Vally region; Nueva Vizcaya, Belance, Lohong) (Yoshitake 2017b).

Pachyrhynchus congestus ocellatus Schultze, 1924

Type depository. SMTD (*).

Type locality. Philippines (Luzon Is.: Benguet Prov., settlement Bokod near Mount Pandan).

Distribution. Only known from the type locality.

Pachyrhynchus congestus pavonius Heller, 1921

Type depository. SMTD (*).

Type locality. Philippines (Luzon Is., Nueva Vizcaya Prov., Imugan).

Distribution. Philippines: Luzon Is. (Nueva Vizcaya, Imugan) (Schultze 1924).

Pachyrhynchus consobrinus Schultze, 1922

Type depository. SMTD (syntypes).

Type locality. Philippines (Luzon Is., Mountain Prov., Bontoc).

Distribution. Philippines: Luzon Is. (Abra, Malibcong, Tineg; Apayao, Conner; Benguet; Moutain Prov., Barlig, Bontoc, Mt. Polis; Kalinga, Pinukpuk) (Schultze 1922b; Schultze 1924; Rukmane-Bārbale 2020b; Rukmane-Bārbale and Cabras 2021).

Pachyrhynchus corpulentus Schultze, 1922

Type depository. SMTD (syntypes).

Type locality. Philippines (Mindanao Is.: Bukidnon Prov., Lindabon).

Distribution. Philippines: Mindanao Is. (Bukidnon, Lindabon; Zamboanga) (Schultze 1922a; Schultze 1924; Rukmane 2019b).

Pachyrhynchus corpulentus balatukan Patano & Macalaba, 2023

Type depository. CMU-MZ (holotype, paratypes).

Type locality. Philippines (Mindanao Is.: Misamis Oriental Prov., Gingoog City, Mt. Balatukan).

Distribution. Only known from the type locality.

Pachyrhynchus croesus Oberthur, 1879

Type depository. NHRS (holotype).

Type locality. Indonesia (Sangir).

Distribution. Indonesia: Sangihe Isls. (Sangir Is.); Talaud Isls. (Kabaruan Is., Salibabu Is.) (Schultze 1924; Yoshitake et al. 2016; Yoshitake 2018a; Rukmane 2018b, 2019b).

Pachyrhynchus cruciatus Schultze, 1923

Type depository. SMTD (*).

Type locality. Philippines (Luzon Is.: Tayabas Prov., Baler).

Distribution. Philippines: Luzon Is. (Abra; Aurora; Ifugao, Banaue; Sierra Madre; Dingalon; Manila) (Rukmane and Barševskis 2016; Rukmane 2018c; Rukmane-Bārbale and Cabras 2021).

Pachyrhynchus cumingii cumingii Waterhouse, 1841

Type depository. SMTD (*).

Type locality. Philippines.

Distribution. Philippines: Leyte Is. (Leyte); Mindanao Is. (Bukidnon, Mt. Kiamo, Cabanglasan, Intavas, Mt. Kalatungan, Panamokan; Sarangani, Kiamba); Bohol Is.; Samar Is. (Hinabangan, Marabut) (Waterhouse 1841; Cabras et al. 2017).

Remarks. The original spelling of this species is “cumingii,” with a double “i,” following the ICZN (Articles 31.1.1 and 31.1.2); however, many references in the literature have incorrectly used the spelling “cumingi”.

Pachyrhynchus cumingii boholensis Schultze, 1924

Type depository. SMTD (*).

Type locality. Philippines (Bohol Is.: Bilar).

Distribution. Only known from the type locality.

Pachyrhynchus davaoensis Schultze, 1934

Type depository. SMTD (holotype).

Type locality. Philippines (Mindanao Is.: Davao Prov., Apo Volcano).

Distribution. Philippines: Mindanao Is. (Bukidnon, Cabanglasan; Cotabato, Tboli; Surigao del Sur, Esperanza, San Miguel) (Rukmane 2019a; Rukmane-Bārbale and Cabras 2021; Cabras et al. 2022a).

Pachyrhynchus decussatus decussatus Waterhouse, 1841

Type depository. NHMUK (holotype).

Type locality. Philippines.

Distribution. Philippines: Catanduanes Is. (Virac) (Waterhouse 1841; Schultze 1923); Luzon Is. (south Luzon, Mt. Bulusan) (Rukmane-Bārbale 2020d).

Pachyrhynchus decussatus catanduanensis Rukmane-Bārbale, 2020

Type depository. DUBC (holotype, paratype).

Type locality. Philippines (Catanduanes Is.: Catanduanes Prov., Pandan).

Distribution. Only known from the type locality.

Pachyrhynchus disargus Rukmane, 2019

Type depository. DUBC (holotype, paratypes).

Type locality. Philippines (Luzon Is.: Nueva Vizcaya Prov., Kasibu).

Distribution. Philippines: Luzon Is. (Nueva Vizcaya, Kasibu, Kayapa) (Rukmane 2019d).

Pachyrhynchus disgestus Heller, 1929

Type depository. SMTD (holotype).

Type locality. Philippines (Luzon Is.: Nueva Vizcaya Prov., Bayombong).

Distribution. Philippines: Luzon Is. (Ifugao, Moutain Prov., Nueva Vizcaya) (Rukmane 2019b; Rukmane-Bārbale and Cabras 2021).

Pachyrhynchus dohrni Behrens, 1887

Type depository. Unknown.

Type locality. Philippines.

Distribution. Philippines: Luzon Is. (Manila) (Rukmane 2019b).

Pachyrhynchus domino Rukmane, 2016

Type depository. DUBC (holotype).

Type locality. Philippines (Mindoro Is.: Mindoro Oriental Prov., Mt. Halcon).

Distribution. Philippines: Mindoro Is. (Mindoro Oriental Prov., Mt. Halcon, Baco) (Rukmane and Háva 2020; Rukmane-Bārbale and Cabras 2021).

Pachyrhynchus dubiosus Schultze, 1922

Type depository. SMTD (syntypes).

Type locality. Philippines (Luzon Is.: Benguet Prov., Berg Santo Tomas und Haight’s Place).

Distribution. Philippines: Luzon Is. (Benguet, Loo, Mt. Santo Tomas, Pauai; Mountain Prov.) (Schultze 1922b; Schultze 1924; Rukmane 2019b, 2019c).

Pachyrhynchus elegans Waterhouse, 1842

Pachyrhynchus eos Heller, 1924: 171. Synonymized by Bollino, Sandel and Rukmane (2017: 201).

Type depository. NHMUK (holotype).

Type locality. Unknown.

Distribution. Philippines: Samar Is. (Northern Samar, Lope De Vega; Samar, Hinabangan) (Waterhouse 1842; Rukmane-Bārbale and Cabras 2021).

Pachyrhynchus eques Heller, 1912

Type depository. SMTD (holotype).

Type locality. Philippines (Luzon Is.: Cagayan, Abulog River).

Distribution. Philippines: Luzon Is. (Cagayan, Abulog River) (Schultze 1924).

Pachyrhynchus equester Heller, 1929

Type depository. SMTD (holotype).

Type locality. Philippines (Luzon Is.: Benguet Prov., Baguio).

Distribution. Only known from the type locality.

Pachyrhynchus erichsoni erichsoni Waterhouse, 1841

Pachyrhynchus erichsoni var. chrysocompsus Heller, 1912: 307. Synonymized by Schultze (1923: 663).

Type depository. NHMUK (syntype); SMTD (syntype).

Type locality. Philippines.

Distribution. Philippines: Dinagat Is.; Leyte Is. (Sogod; Cabalian); Samar Is. (Hinabanga); Luzon Is. (Sorsogon; Cagayan; Ifuga, Banaue; Sierra Madre; Aurora, Dingalan, Labulo; Nueva Vizcaya, Belance; Quirino, Disimungal, Madela); Marinduque Is. (Boac; Buenavista); Mindoro Is. (Baco); Mindanao Is. (Bukidnon, Mt. Kiamo, Cabanglasan, Intavas, Panamokan; Cotobato, Kidapawan, Mt. Apo; Surigao del Sur, Esperanza; Davao City, Marilog District, Catigan, Toril, Tamayong, Calinan) (Waterhouse 1841; Schultze 1923; Cabras et al. 2017; Rukmane 2019b; Rukmane-Bārbale and Cabras 2021; Cabras et al. 2022a).

Host plants. The P. erichsoni complex has been documented on the following plants: Melastoma malabathricum (Melastomataceae), Lithocarpus boholensis (Fagaceae), Dendrocnide sp. (Urticaceae), Callicarpa sp. (Lamiaceae), Philodendron sp. (Araceae), and Amaranthus sp. (Amaranthaceae) (Cabras et al. 2022a).

Pachyrhynchus erichsoni eschscholtzii Waterhouse, 1841

Type depository. NHMUK (syntype); SMTD (syntype).

Type locality. Philippines.

Distribution. Philippines: Luzon Is. (Laguna Prov., Paete, Mt. Banahao; Ilocos Norte Prov., Bangui); Polillo Is.; Marinduque Is. (Waterhouse 1841; Schultze 1923, 1924; Rukmane-Bārbale and Cabras 2021).

Pachyrhynchus erosus Schultze, 1920

Type depository. SMTD (syntypes).

Type locality. Philippines (Luzon Is.: Benguet Prov., mountain trail near Atoc).

Distribution. Philippines: Luzon Is. (Benguet, Abatan, La Trinidad) (Rukmane-Bārbale and Cabras 2021).

Pachyrhynchus esperanza Bollino, Sandel & Rukmane, 2017

Type depository. SMTD (holotype); MBLI, DUBC, NIAES, KUM, CFS (paratypes).

Type locality. Philippines (Mindanao Is.: Agusan del Sur Prov.).

Distribution. Philippines: Mindanao Is. (Agusan del Sur, Esperanza, Sibagat, Talacogon; Caraga region; Bukidnon, Cabanglasan, Valencia; Surigao del Sur).

Pachyrhynchus faisali Bollino, 2023

Type depository. SMTD (holotype).

Type locality. Indonesia (New Guinea Is.: West Papua, South Sorong Regency, Teminabuan).

Distribution. Only known from the type locality.

Pachyrhynchus felipeae Rukmane & Cabras, 2018

Type depository. DUBC (holotype, paratypes).

Type locality. Philippines (Panay Is.: Antique Prov., Culasi, Aklan).

Distribution. Philippines: Panay Is. (Antique, Culasi, Aklan).

Remarks. This species is very similar and sympatric with P. franciscoi Rukmane and Cabras (2018b) from Panay Island, Philippines.

Pachyrhynchus florulentus Yoshitake, 2019

Type depository. KUM (holotype); NIAES (paratype).

Type locality. Philippines (Luzon Is.: Ilocos Norte Prov., Pagudpud).

Distribution. Philippines: Luzon Is. (Cagayan, Ilocos Norte, Nueva Vizcaya) (Yoshitake 2019c; Rukmane-Bārbale and Cabras 2021).

Pachyrhynchus forsteni Vollenhoven, 1864

Type depository. Unknown.

Type locality. Indonesia (Molucas Is.: North Maluku Prov., Ternate).

Distribution. Indonesia: Maluku Isls. (Halmahera Is., Makian Is., Molucas Is., Ternate Is., Morotai Is.) (Yoshitake 2018b; Rukmane 2019b; Bollino 2023).

Remarks. According to the original description of P. forsteni by Snellen van Vollenhoven (1864), the distribution of this species included Ternate, Halmahera, and Sumatra. However, the record from Sumatra is now questioned due to geographical discrepancies (Yoshitake 2018b).

Pachyrhynchus franciscoi Rukmane & Cabras, 2018

Type depository. DUBC (holotype, paratype).

Type locality. Philippines (Panay Is.: Antique Prov., Culasi, Aklan).

Distribution. Philippines: Panay Is. (Antique, Culasi, Aklan).

Remarks. This species is very similar and sympatric with P. felipeae Rukmane and Cabras (2018b) from Panay Island, Philippines.

Pachyrhynchus galeraensis Schultze, 1934

Type depository. SMTD (holotype).

Type locality. Philippines (Mindoro Is.: Puetro Galera).

Distribution. Philippines: Mindoro Is. (Puetro Galera).

Pachyrhynchus gemmatus gemmatus Waterhouse, 1841

Pachyrhynchus gemmatus var. atratus Heller, 1912: 308. Synonymized by Schultze (1924: 329).

Type depository. NHMUK (syntype), SMTD (*).

Type locality. Philippines.

Distribution. Philippines: Luzon Is. (Aurora; Calayan, Sanchez Mira, Santa Ana.; Moutain Prov.; Manila; Nueva Vizcaya; Isabela; Quirino) (Waterhouse 1841; Schultze 1924; Rukmane 2018c, 2019b; Rukmane-Bārbale and Cabras 2021).

Pachyrhynchus gemmatus purpureus Kraatz, 1888

Type depository. SMTD (*).

Type locality. Philippines (Luzon Is.).

Distribution. Philippines: Luzon Is. (Cagayan, Sanchez Mira) (Schultze 1924).

Pachyrhynchus gilvomaculatus Yoshitake, 2017

Type depository. NIAES (holotype); KUM, MBLI (paratypes).

Type locality. Philippines (Mindanao Is.: South Cotabato Prov., Mt. Matutum).

Distribution. Philippines: Mindanao Is. (Soccsksargen region, South Cotabato Prov.) (Yoshitake 2017c).

Pachyrhynchus gloriosus gloriosus Faust, 1895

Type depository. SMTD (*).

Type locality. Philippines (Luzon Is.).

Distribution. Philippines: Luzon Is. (Aurora; Ifugao; Isabela; Kalinga; Laguna, Mt. Banahao; Nueva Vizcaya; Rizal) (Faust 1895a; Schultze 1924; Rukmane-Bārbale and Cabras 2021).

Remarks. A specimen examined by Rukmane (2019b) was recorded from Negros Island, a locality that differs from previously known distribution records, which have been confined to Luzon. While this may be a special record, its validity as a confirmed observation requires further verification.

Pachyrhynchus gloriosus abbreviatus Schultze, 1922

Type depository. SMTD (*).

Type locality. Philippines (Luzon Is.: Mountain Prov., Bontoc).

Distribution. Philippines: Luzon Is. (Mountain Prov., Bontoc; Kalinga, Lubuagan) (Schultze 1922b; Schultze 1924).

Pachyrhynchus halconensis Schultze, 1922

Type depository. SMTD (syntypes).

Type locality. Philippines (Mindoro Is.: Mt. Halcon).

Distribution. Philippines: Mindoro Is. (Mt. Halcon) (Schultze 1922b; Schultze 1923; Rukmane 2019b; Rukmane-Bārbale and Cabras 2021).

Pachyrhynchus helenperrinae Rukmane, 2018

Type depository. MNHN (holotype, paratypes).

Type locality. Philippines (Ch. Semper).

Distribution. Philippines: Mindanao Is. or Mindanao PAIC.

Remarks. Rukmane (2018d) mentioned that species on a particular island share common features, which are clearly distinct from those of species found on another island. Therefore, based on their general appearance, this species might be distributed across Mindanao Island or the Mindanao PAIC (Rukmane 2018d).

Pachyrhynchus helleri Kuntzen, 1914

Type depository. ZMHB (holotype).

Type locality. Philippines (Luzon Is.).

Distribution. Philippines: Luzon Is. (Schultze 1924).

Pachyrhynchus hirokii Yoshitake, 2012

Type depository. NIAES (holotype, paratypes); PCKS (paratypes).

Type locality. Philippines (Mindanao Is.: Mt. Apo).

Distribution. Philippines: Mindanao Is. (Bukidnon; Cotabato; Davao del Sur; Lanao del Sur) (Rukmane 2019b; Rukmane-Bārbale and Cabras 2021).

Pachyrhynchus igorota Schultze, 1917

Type depository. SMTD (syntypes)

Type locality. Philippines (Luzon Is.: Benguet Prov., Haight’s Place).

Distribution. Philippines: Luzon Is. (Benguet, Mt. Pawai, Mt. Pulog; Ifugao; Isabela, Mt. Moises; Mountain Prov., Mt. Polis; Nueva Vizcaya) (Schultze 1924; Rukmane 2019c; Rukmane-Bārbale and Cabras 2021).

Pachyrhynchus ilgas Rukmane, 2017

Type depository. DUBC (holotype).

Type locality. Philippines (Samar Is.: Northern Samar Prov., Lope De Vega).

Distribution. Philippines: Samar Is. (Northern Samar) (Rukmane 2017).

Pachyrhynchus imitans Rukmane & Bollino, 2020

Type depository. SMTD (holotype); DUBC, MZZC, MBLI (paratypes).

Type locality. Philippines (Mindanao Is.: Zamboanga City, Labuan).

Distribution. Philippines: Mindanao Is. (Zamboanga City, Zamboanga del Norte, Zamboanga Sibugay).

Pachyrhynchus inclytus Pascoe, 1873

Pachyrhynchus ignipes Chevrolat, 1881b: 359. Synonymized by Schultze (1924: 339).

Pachyrhynchus modestior Behrens, 1887: 240. Synonymized by Schultze (1924: 339).

P. modestior var. apicalis Kraatz, 1888: 26. Synonymized by Schultze (1924: 340).

P. modestior var. transversatus Heller, 1921: 544. Synonymized by Schultze (1924: 340).

Type depository. NHMUK (syntype), SMTD (*).

Type locality. Philippines.

Distribution. Philippines: Luzon Is. (Benguet, Mt. Trail, Baguio, Mt. Mirador, Mt. Santo Tomas, Trinidad; Ifugao; Manila; Mountain Prov.; Nueva Vizcaya, Imugan) (Schultze 1924; Rukmane 2019b, 2019c; Rukmane-Bārbale and Cabras 2021).

Pachyrhynchus infernalis Fairmaire, 1897

Pachyrhynchus niger Sakaguchi, 1927: 24. Synonymized by Rukmane (2018a: 65).

Type depository. Unknown.

Type locality. Japan (Ryukyu, Ishigaki Is.).

Distribution. Japan: Ryukyu Isls. (Iriomote Is., Ishigaki Is.) (Schultze 1923; Starr and Wang 1992).

Host plants. Mangifera indica (Anacardiaceae) and Glochidion sp. (Euphorbiaceae) (Oshiro 1991).

Remarks. Pachyrhynchus infernalis is known from Ryukyu Islands. However, the identification of a specimen from “Bangsamoro” as P. infernalis by Rukmane-Bārbale and Cabras (2021) from the DUBC collection might need to be reevaluated.

Pachyrhynchus jitanasaius Chen & Lin, 2017

Type depository. NMNS (holotype, paratypes).

Type locality. Taiwan (Taitung County, Ludao Is.).

Distribution. Taiwan: Ludao Is.

Host plants. Among all the individuals found on the host plants, 50% were found on Shoebutton ardisia (Ardisia elliptica, Primulaceae), 48% on beef wood (Casuarina equisetifolia, Casuarinaceae), and 2% on litsea (Litsea acutivens, Lauraceae) (Chen et al. 2017).

Pachyrhynchus jugifer Waterhouse, 1841

Pachyrhynchus rhodopterus Chevrolat, 1841: 224. Synonymized by Schultze (1923: 653).

Type depository. NHMUK (holotype).

Type locality. Philippines.

Distribution. Philippines: Panay Is. (Capiz, Aklan, Jamindan, Mt. Macosolon) (Waterhouse 1841; Schultze 1923; Rukmane 2019b; Rukmane-Bārbale and Cabras 2021).

Remarks. Four specimens examined by Rukmane (2018c) were recorded from Luzon Island (Manila). However, other specimens were found on Panay Island.

Pachyrhynchus kirklayroni Rukmane, 2019

Type depository. DUBC (holotype).

Type locality. Philippines (Luzon Is.: Ilocos Norte Prov., Adams).

Distribution. Philippines: Luzon Is. (Ilocos Norte) (Rukmane 2019d).

Pachyrhynchus kraslavae Rukmane & Barševskis, 2016

Type depository. DUBC (holotype).

Type locality. Philippines (Mindanao Is.: Davao de Oro Prov., Mabini).

Distribution. Philippines: Mindanao Is. (Agusan del Sur, San Francisco; Surigao del Norte) (Rukmane 2019a).

Pachyrhynchus lacunosus Heller, 1912

Type depository. NHMUK (syntype); SDEI (syntype).

Type locality. Philippines.

Distribution. Philippines: Luzon Is. (Benguet, Mt. Pulogloko; Ifugao, Asipulo, Banaue, Hungduan, Tinok; Isabela; Nueva Viscaya, Ambaguio, Belance, Dupax Del Sur, Kasibu, Kayapa, Malico, Sta. Fe.; Quezon) (Schultze 1924; Rukmane-Bārbale 2020b; Rukmane-Bārbale and Cabras 2021).

Pachyrhynchus latifasciatus Waterhouse, 1842

Type depository. NHMUK (holotype).

Type locality. Philippines.

Distribution. Philippines: Samar Is. (Northern Samar, Lope de Vega) (Waterhouse 1842; Yoshitake 2017c; Rukmane-Bārbale and Cabras 2021).

Pachyrhynchus layroni Rukmane & Cabras, 2018

Type depository. DUBC (holotype, paratypes).

Type locality. Philippines (Panay Is.: Antique Prov., Madja-as).

Distribution. Philippines: Panay Is. (Antique) (Rukmane and Cabras 2018b).

Pachyrhynchus libucanus Schultze, 1923

Type depository. SMTD (holotype).

Type locality. Philippines (Libucan Is.: near Samar).

Distribution. Philippines: Libucan Is.

Pachyrhynchus loheri Schultze, 1917

Type depository. SMTD (holotype).

Type locality. Philippines (Luzon Is.: Bulacan Prov., Mount Guinuisan).

Distribution. Philippines: Luzon Is. (Bulacan, Mount Guinisan) (Schultze 1924; Rukmane-Bārbale 2020b).

Pachyrhynchus lorquini Chevrolat, 1881

Pachyrhynchus flavopunctatus Kraatz, 1888: 30. Synonymized by Schultze (1924: 334).

Pachyrhynchus flavomaculatus Kraatz, 1888: 32. Synonymized by Schultze (1924: 334).

Type depository. NHRS (syntypes).

Type locality. Philippines.

Distribution. Philippines: Luzon Is. (Laguna, Mt. Maquiling); Marinduque Is. (Mt. Malindig) (Schultze 1924; Rukmane 2018c; Rukmane 2019b).

Remarks. In the original description of P. lorquini, the type specimen was recorded from Maldonado (Chevrolat 1881b), but subsequent findings were in Luzon. The actual distribution range of the species needs to be re-evaluated.

Pachyrhynchus lubanganus Bollino & Sandel, 2015

Type depository. SMTD (holotype); MBLI, CFS (paratypes).

Type locality. Philippines (Lubang Is.: south-west of Tilik).

Distribution. Philippines: Lubang Is.

Pachyrhynchus marinduquensis Rukmane & Barševskis, 2016

Type depository. DUBC (holotype, paratype).

Type locality. Philippines (Marinduque Is.: Marinduque Prov., Buenavista).

Distribution. Philippines: Marinduque Is. (Marinduque, Buenavista, Boac; Mt. Malindig) (Rukmane-Bārbale and Cabras 2021).

Pachyrhynchus maruyamai Yoshitake, 2019

Type depository. NIAES (holotype); MCKUM (paratypes).

Type locality. Philippines (Catanduanes Is.: San Miguel, Kilikilihan).

Distribution. Philippines: Catanduanes Is. (Yoshitake 2019f).

Pachyrhynchus masatoshii Yoshitake & Yap, 2017

Type depository. UPLB (holotype).

Type locality. Philippines (Luzon Is.: Quezon Prov., Dolores, Brgy. Kinabuhayan, Mt. Banahaw).

Distribution. Philippines: Luzon Is. (Calabarzon regio) (Yoshitake and Yap 2017).

Pachyrhynchus masatoshiensis Rukmane-Bārbale, 2024

Type depository. DUBC (holotype, paratypes).

Type locality. Philippines (Luzon Is.: Albay Prov., Tabaco, Mayon Volcano).

Distribution. Philippines: Luzon Is. (Quezon, Gen Nakar) (Rukmane-Bārbale 2024).

Pachyrhynchus miltoni Cabras & Rukmane, 2016

Type depository. CMU-MZ (holotype, paratype).

Type locality. Philippines (Mindanao Is.: Davao del Sur Prov., Davao City, Marilog District).

Distribution. Philippines: Mindanao Is. (Davao del Sur) (Mohagan et al. 2018; Rukmane-Bārbale and Cabras 2021; Cabras et al. 2022a).

Host plants. This species has been documented on Melastoma malabathricum (Melastomataceae), Piper aduncum (Piperaceae), Lithocarpus boholensis (Fagaceae), Theobroma cacao (Malvaceae), Atuna racemosa (Chrysobalanaceae), and Helianthus sp. (Asteraceae) (Cabras et al. 2022a).

Pachyrhynchus mindoroensis Rukmane & Háva, 2020

Type depository. DUBC (holotype, paratypes); JHAC (paratype).

Type locality. Philippines (Mindoro Is.: Oriental Mindoro Prov., Puerto Galera).

Distribution. Philippines: Mindoro Is. (Oriental Mindoro, Baco, Puerto Galera, Bacon, Mt. Halcon).

Pachyrhynchus mohagani Bollino & Sandel, 2015

Type depository. SMTD (holotype); MBLI, CFS (paratypes).

Type locality. Philippines (Lubang Is.: south-west of Tilik).

Distribution. Philippines: Lubang Is.

Pachyrhynchus mollendorffi mollendorffi Heller, 1898

Type depository. SMTD (holotype).

Type locality. Philippines.

Distribution. Philippines: Panay Is. (Antique) (Rukmane and Cabras 2018a).

Pachyrhynchus mollendorffi marinduquanus Rukmane & Cabras, 2018

Type depository. DUBC (holotype, paratypes).

Type locality. Philippines (Marinduque Is.: Marinduque Prov., Buenavista).

Distribution. Philippines: Marinduque Is. (Marinduque, Buenavista, Mt. Malinding) (Rukmane and Cabras 2018a).

Pachyrhynchus moniliferus moniliferus Germar, 1824

Pachyrhynchus confinis Chevrolat, 1841: 226. Synonymized by Schultze (1923: 634).

Type depository. SMTD (*).

Type locality. Philippines (Luzon Is.: Manila).

Distribution. Philippines: Luzon Is. (Aurora, Dingalan; Cagayan; Ifugao, Banaue; Quirino; Nueva Vizcaya, Belance; Isabela; Mountain Prov.; Kalinga, Pinukpuk; Benguet; Camarines; Batangas; Laguna, Los Baños, Mt. Makiling, Mt. Banahao; Rizal, Bosoboso, Montalban, Taytay, Antipolo; Manila; Paete, Bataan, Limay); Mindoro Is. (Mt. Halcon); Marinduque Is.; Bohol Is.; Negros Is.; Camiguin Is.; Calayan Is.; Catanduanes Is. (Schultze 1923; Rukmane 2018c, 2018e, 2019b, 2019c; Rukmane-Bārbale and Cabras 2021).

Host plant. Cocoa, Theobroma cacao (Kayashima 1940); Bishop Wood, Bischofia javanica (Phyllanthaceae) (this study).

Remarks. The larvae of this species were found to feed on cacao fruit imported from the Philippines to Taiwan (Kayashima 1940). In addition, it is considered as one of the insect pests of cocoa plantations in the Philippines, especially in Luzon (Domingo et al. 2023).

Pachyrhynchus moniliferus abranus Heller, 1934

Type depository. SMTD (*).

Type locality. Philippines (Luzon Is.: Abra Prov.).

Distribution. Philippines: Luzon Is. (Abra, Nueva Vizcaya); Mindoro Is. (Baco; Mt. Halcon) (Rukmane 2018e).

Remarks. Heller (1934) treated P. stellulifer as a distinct species, and described abranus as its subspecies simultaneously. However, Heller described stellulifer as a variety (subspecies) of P. moniliferus in 1912. This classification has been followed by Schultze (1924, 1934) and Rukmane (2018a, 2018e). There has been no further related study since then until Rukmane (2018a, e) listed abranus as a subspecies of P. moniliferus, which is followed herein. Rukmane (2018e) incorrectly spelled “abranus” as “eburnus”, and the specimens from Mindoro Island might need to be evaluated in the future.

Pachyrhynchus moniliferus babuyanensis Rukmane, 2018

Type depository. DUBC (holotype, paratypes).

Type locality. Philippines (Babuyan Is.).

Distribution. Only known from the type locality.

Pachyrhynchus moniliferus chevrolati Eydoux & Souleyet, 1839

Pachyrhynchus chlorolineatus Waterhouse, 1841: 25. Synonymized by Schultze (1923: 644).

Pachyrhynchus mandarinus Chevrolat, 1841: 226. Synonymized by Schultze (1923: 644).

Pachyrhynchus concinnus Waterhouse, 1842: 70. Synonymized by Schultze (1923: 644).

Type depository. SMTD (*).

Type locality. Philippines.

Distribution. Philippines: Catanduanes Is. (Catanduanes Prov., Virac); Luzon Is. (Aurora, Dingalan; Isabela; Quirino; Manila; Tayabas; Albay; Polillo); Samar Is. (Catarman, Borongan); Mindanao Is. (Misamis Occidental, Labuyo) (Schultze 1923; Rukmane 2018c, 2018d, 2018e).

Pachyrhynchus moniliferus herbidus Rukmane, 2018

Type depository. DUBC (holotype); SMTD (paratypes).

Type locality. Philippines (Samar Is.: Northern Samar, Lope De Vega).

Distribution. Only known from the type locality.

Pachyrhynchus moniliferus jagori Heller, 1912

Type depository. SMTD (*).

Type locality. Philippines (Samar Is.).

Distribution. Philippines: Samar Is.; Catanduanes Is. (Rukmane 2018e).

Remarks. The type locality of this subspecies is Samar Island, Philippines (Heller 1912). However, Rukmane (2018e) examined a specimen from Catanduanes Island, which requires further clarification.

Pachyrhynchus moniliferus stellulifer Heller, 1912

Type depository. SMTD (*).

Type locality. Philippines.

Distribution. Philippines: Luzon Is. (Benguet, Ilocos Sur, Pangasinan); ​​Mindoro Is. (Naujan; Mangaran; San Jose; Mt. Halcon; Puerto Galera) (Schultze 1923, 1934).

Pachyrhynchus morio Heller, 1912

Type depository. NHMUK (syntype); SMTD (syntypes).

Type locality. Philippines (Luzon Is.).

Distribution. Philippines: Luzon Is. (Benguet, Mt. Lusong) (Schultze 1924).

Pachyrhynchus morotaiensis Vollenhoven, 1864

Pachyrhynchus waterhousei Faust, 1895b: 95. Synonymized by Schultze (1923: 661).

Type depository. SMTD (syntypes).

Type locality Indonesia (Morotai).

Distribution. Indonesia: Maluku Isls. (Morotai, Moluccas) (Faust 1895b; Schultze 1923; Rukmane 2018c).

Pachyrhynchus multipunctatus multipunctatus Waterhouse, 1841

Pachyrhynchus auroguttatus Chevrolat, 1881: 348. Synonymized by Schultze (1924: 316).

Type depository. Unknown.

Type locality. Philippines.

Distribution. Philippines: Bohol Is. (Bohol, Sevilla); Luzon Is. (Nueva Vizcaya; Manila) (Waterhouse 1841; Schultze 1924; Yoshitake 2018c; Rukmane 2019b; Rukmane-Bārbale and Cabras 2021).

Pachyrhynchus multipunctatus endoi Yoshitake, 2018

Type depository. NIAES (holotype, paratypes); MCKUM (paratypes).

Type locality. Philippines (Cebu Is.: Cebu city, Barangay Busay, Sitio Rosvelt).

Distribution. Only known from the type locality (Yoshitake 2018c).

Pachyrhynchus naokii Yoshitake, 2012

Type depository. NIAES (holotype).

Type locality. Philippines (Mindanao Is.: Mt. Syniop).

Distribution. Philippines: Mindanao Is. (Cotabato, Sultan Kudarat; Lanao del Sur) (Rukmane-Bārbale and Cabras 2021).

Pachyrhynchus naokoae Yoshitake, 2019

Pachyrhynchus sibuyanensis Rukmane 2019b: 42. Synonymized by Yoshitake (2019a: 408).

Type depository. NIAES (holotype, paratypes); KUM (paratypes).

Type locality. Philippines (Sibuyan Is.: Mimaropa region, Romblon Prov.).

Distribution. Only known from the type locality.

Pachyrhynchus negrosensis Schultze, 1924

Type depository. SMTD (holotype).

Type locality. Philippines (Negros Is.: Occidental Negros Prov., Cuernos Mountains).

Distribution. Philippines: Negros Is. (Occidental Negros Prov., Mt. Canlaon, Benedicto) (Rukmane 2019b; Rukmane-Bārbale and Cabras 2021).

Pachyrhynchus neoabsurdus Rukmane, 2017

Type depository. DUBC (holotype, paratype).

Type locality. Philippines (Mindanao Is.: Bukidnon Prov., Mt. Kalatungan).

Distribution. Philippines: Mindanao Is. (Bukidnon, Intavas) (Rukmane 2017).

Pachyrhynchus niisatoi Yoshitake, 2017

Type depository. KUM (holotype); KUM, MBLI, NIAES (paratypes).

Type locality. Philippines (Luzon Is.: Quirino Prov., Nagutipunan).

Distribution. Philippines: Luzon Is. (Central Luzon; Cagayan Valley region; Aurora; Nueva Vizcaya, Kayapa, Kasibu, Dupax Del Sur; Quirino, Madela, Nagtipunan; Isabela; Aurora; Rizal) (Yoshitake 2017d; Rukmane-Bārbale 2020b; Rukmane-Bārbale and Cabras 2021).

Pachyrhynchus nitcisi Rukmane & Barševskis, 2016

Type depository. DUBC (holotype, paratypes).

Type locality. Philippines (Mindanao Is.: Sarangani Prov., Malungon).

Distribution. Philippines: Mindanao Is. (Agusan; Agusan del Sur, Sibagat; Bukidnon; Cotabato; Sarangani, Malungon) (Rukmane-Bārbale and Cabras 2021).

Pachyrhynchus nobilis nobilis Heller, 1912

Type depository. NHMUK (syntype); SMTD (syntype); SDEI (syntype).

Type locality. Philippines.

Distribution. Philippines: Babuyan Islands (Babuyan Claro; Calayan Is.); Luzon Is. (Ifugao; Isabela; Moutain Prov.; Nueva Vizcaya; Quirino) (Rukmane 2019b; Rukmane-Bārbale and Cabras 2021; Tseng et al. 2021).

Pachyrhynchus nobilis yamianus Kano, 1929

Type depository. Unknown.

Type locality. Taiwan (Taitung County, Lanyu Is. (= Kotosho)).

Distribution. Taiwan: Lanyu Is.; Ludao Is.

Host plants. Epipremnum pinnatum (Araceae) (this study).

Remarks. Pachyrhynchus nobilis yamianus was initially considered a distinct species as P. yamianus Kano, 1929. However, Schultze (1937) examined the specimens and regard it as a subspecies of P. nobilis. Whether P. nobilis yamianus is a distinct species requires re-examination.

Pachyrhynchus noeli Yoshitake, 2019

Type depository. KUM (holotype).

Type locality. Philippines (Sibuyan Is.: Mimaropa region, Romblon Prov.).

Distribution. Only known from the type locality (Yoshitake 2019e).

Pachyrhynchus notocruciatus Yoshitake, 2017

Type depository. NIAES (holotype).

Type locality. Philippines (Mindanao Is.: Mt. Apo).

Distribution. Philippines: Mindanao Is. (Cotabato, Alamada; Mt. Apo) (Yoshitake 2017c; Rukmane-Bārbale and Cabras 2021).

Pachyrhynchus obumanuvu Cabras, Donato, Medina, & Van Dam, 2021

Type depository. PNMNH (holotype); UMCRC, CASENT, MBLI (paratypes).

Type locality. Philippines (Mindanao Is.: Davao City).

Distribution. Philippines: Mindanao Is. (Bukidnon, Davao city) (Cabras et al. 2021, 2022a).

Host plants. Procris urdanetensis (Urticaceae) and Elatostema sp. (Urticaceae). (Cabras et al. 2021, 2022a)

Pachyrhynchus occidentalis Rukmane, 2017

Type depository. DUBC (holotype, paratypes).

Type locality. Philippines (Mindanao Is.: Davao Del Sur Prov., Kapatagan).

Distribution. Philippines: Mindanao Is. (Cotabato, Mt. Parke; Davao Del Sur, Mt. Apo; Sarangani, Kiamba) (Rukmane 2017; Háva and Rukmane 2018; Rukmane 2019b; Yoshitake and Bollino 2019; Rukmane-Bārbale and Cabras 2021).

Pachyrhynchus ochroplagiatus ochroplagiatus Heller, 1912

Type depository. SMTD (holotype).

Type locality. Philippines (Luzon Is.: Benguet Prov., Mt. Pulog).

Distribution. Philippines: Luzon Is. (Aurora; Benguet, Mt. Pulog; Nueva Vizcaya) (Schultze 1924; Rukmane-Bārbale and Cabras 2021).

Pachyrhynchus ochroplagiatus multiplagiatus Schultze, 1924

Type depository. SMTD (*).

Type locality. Philippines.

Distribution. Philippines: Luzon Is. (Rukmane 2018a).

Pachyrhynchus octoannulatus Yoshitake, Bollino & Sandel, 2019

Type depository. NIAES (holotype, paratypes); KUM, CFS, MBLI (paratypes).

Type locality. Philippines (Mindanao Is.: Lanao del Sur Prov., Wao).

Distribution. Philippines: Mindanao Is. (Bukidnon; Bangsamoro Autonomous Region ni Muslim Mindanao; Soccksargen region; Lanao del Sur, Wao; Cotabato) (Rukmane 2019a; Yoshitake et al. 2019b).

Pachyrhynchus ohbayashii Yoshitake, 2017

Type depository. NIAES (holotype).

Type locality. Indonesia (Biak Is.: Irian Jaya, North of Biak).

Distribution. Only known from the type locality (Yoshitake 2017a).

Pachyrhynchus orbifer orbifer Waterhouse, 1841

Pachyrhynchus alboguttatus Chevrolat, 1841: 226. Synonymized by Schultze (1923: 638).

Pachyrhynchus fahrei Schoenherr, 1845: 388. Synonymized by Schultze (1923: 638).

Pachyrhynchus fimbriatus Chevrolat, 1841: 224. Synonymized by Schultze (1923: 637).

Pachyrhynchus globulipennis Chevrolat, 1841: 225. Synonymized by Schultze (1923: 637).

Pachyrhynchus pretiosus Chevrolat, 1841: 225. Synonymized by Schultze (1923: 637).

Pachyrhynchus scintillans Chevrolat, 1841: 225. Synonymized by Schultze (1923: 637).

Type depository. NHRS (syntypes); SMTD (*).

Type locality. Philippines.

Distribution. Philippines: Luzon Is. (Aurora; Benguet, Baguio to Bontoc, Bued River Valley, Baguio, Mt. Santo Tomas, Mount Mirador, Trinidad, Mountain Trail; Ifugao; Ilocos Norte, Bangui, Burgos, Mount Nagapatan; Isabela; Mountain Prov.; Nueva Viscaya; Quirino; Kalinga, Lubuagan; Cagayan, Sanchez Mira; Manila) (Waterhouse 1841; Schultze 1923; Rukmane and Barševskis 2017; Rukmane 2018c, 2019b, 2019c).

Remarks. One female was found in Ako (Pingtung, Taiwan) (Kôno 1930); however, no record was found after that.

Pachyrhynchus orbifer ardens Chevrolat, 1841

Type depository. SMTD (*).

Type locality. Philippines.

Distribution. Philippines: Luzon Is. (Cagayan, Sanchez Mira) (Schultze 1923; Rukmane 2018a).

Pachyrhynchus orbifer azureus Schultze, 1922

Type depository. SMTD (*).

Type locality. Philippines (Luzon Is.: Benguet Prov., Kabayan).

Distribution. Philippines: Luzon Is. (Benguet, Kabayan; Cagayan; Isabela; Kalinga) (Schultze 1922b; Schultze 1923; Rukmane and Barševskis 2017).

Pachyrhynchus orbifer callainus Yoshitake, 2017

Type depository. NIAES (holotype, paratypes).

Type locality. Philippines (Luzon Is.: Cagayan Prov., Sierra Madre).

Distribution. Philippines: Luzon Is. (Cagayan Valley region) (Yoshitake 2017b).

Pachyrhynchus orbifer circulifer Chevrolat, 1841

Type depository. SMTD (*).

Type locality. Philippines.

Distribution. Philippines: Luzon Is. (Apayao; Cagayan; Mountain Prov.; Ilocos Norte, Bangui; Isabela; Quirino; Nueva Vizcaya) (Schultze 1923; Rukmane and Barševskis 2017).

Pachyrhynchus orbifer gemmans Chevrolat, 1841

Type depository. NHRS (syntypes); SMTD (*).

Type locality. Philippines (Luzon Is.: Manila).

Distribution. Philippines: Luzon Is. (Apoyao; Cagayan, Tuao, Rio Chico; Isabela, Mt. Moises; Ifugao; Mountain Prov.; Quirino; Laguna; Manila) (Rukmane and Barševskis 2017; Rukmane 2018c, 2019c).

Pachyrhynchus orbifer inornatus Waterhouse, 1841

Type depository. SMTD (*).

Type locality. Philippines.

Distribution. Philippines: Calayan Is.; Luzon Is. (Cagayan, Ilocos Norte) (Waterhouse 1841; Schultze 1923).

Pachyrhynchus orbifer murinus Heller, 1934

Type depository. SMTD (*).

Type locality. Philippines (Luzon Is.: Ilocos Sur Prov., Cabugao).

Distribution. Philippines: Luzon Is. (Apayao; Ilocos Sur; Mountain Prov.; Quirino) (Rukmane and Barševskis 2017).

Pachyrhynchus orbifer striatomaculatus Yoshitake, 2017

Type depository. NIAES (holotype, paratypes); KUM (paratypes).

Type locality. Philippines (Luzon Is.: Nueva Vizcaya Prov., Aritao).

Distribution. Philippines: Luzon Is. (Cagayan Valley region) (Yoshitake 2017c).

Pachyrhynchus orientalis Rukmane, 2017

Type depository. DUBC (holotype, paratypes).

Type locality. Philippines (Mindanao Is.: Bukidnon Prov., Cabanglasan).

Distribution. Philippines: Mindanao Is. (Bukidnon, Cabanglasan; Surigao Del Sur, San Miguel) (Rukmane 2017).

Pachyrhynchus ottomerkli Rukmane, 2019

Type depository. HUF (holotype, paratypes).

Type locality. Philippines (Mindanao Is.: Bukidnon Prov., Maluko).

Distribution. Only known from the type locality (Rukmane 2019c).

Pachyrhynchus panumanon Cabras & Medina, 2022

Type depository. PNMNH (holotype); UMCRC (paratypes).

Type locality. Philippines (Mindanao Is.: Misamis Oriental Prov., Gingoog City).

Distribution. Only known from the type locality.

Host plants. Glochidion sp. (Phyllanthaceae) and Clethra sp. (Clethraceae) (Cabras et al. 2022b).

Pachyrhynchus perpulcher Waterhouse, 1841

Type depository. NHMUK (holotype).

Type locality. Philippines.

Distribution. Philippines: Babuyan Is.; Batanes Is.; Calayan Is.; Camiguin Is.; Luzon Is. (Cagayan; Ilocos Norte; Nueva Vizcaya; Kalinga, Pinukpuk; Quirino) (Waterhouse 1841; Schultze 1924; Rukmane 2019b; Rukmane-Bārbale and Cabras 2021; Tseng et al. 2021).

Pachyrhynchus phaleratus phaleratus Waterhouse, 1841

Pachyrhynchus elenae Rukmane, 2016: 82. Synonymized by Yoshitake, Bollino and Sandel (2019a: 197).

Type depository. NHMUK (holotype).

Type locality. Philippines (Catanduanes Is.: Virac).

Distribution. Philippines: Catanduanes Is. (Virac); Luzon Is. (Aurora; Isabela; Quirino) (Waterhouse 1841; Schultze 1923; Yoshitake et al. 2019a).

Pachyrhynchus phaleratus badiovittatus Yoshitake, 2019

Type depository. KUM (holotype).

Type locality. Philippines (Marinduque Is.: Marinduque Prov., Gasan, Tiguion).

Distribution. Only known from the type locality.

Pachyrhynchus phaleratus dannylayroni Rukmane, 2019

Type depository. DUBC (holotype).

Type locality. Philippines (Luzon Is.: Aurora Prov., Dingalan).

Distribution. Only known from the type locality (Rukmane 2019d).

Pachyrhynchus pinorum pinorum Pascoe, 1873

Pachyrhynchus pinorum var. dimidiatus Heller, 1912: 306. Synonymized by Schultze (1924: 351).

Type depository. NHMUK (syntype), SMTD (*).

Type locality. Philippines (Luzon Is.).

Distribution. Philippines: Luzon Is. (Mt. Makiling; Benguet, Mt. Trail, Baguio, Atoc, Tublay, La Trinidad; Ifugao, Banaue, Hungduan; Moutain Prov., Barlig; Nueva Vizcaya, Kayapa, Sta. Fe., Dupax) (Schultze, 1920; Rukmane 2019b, 2019c; Rukmane-Bārbale 2020b; Rukmane-Bārbale and Cabras 2021).

Pachyrhynchus pinorum transversalis Heller, 1912

Type depository. SMTD (*).

Type locality. Philippines.

Distribution. Philippines: Luzon Is.

Pachyrhynchus postpubescens postpubescens Schultze, 1922

Type depository. SMTD (syntypes).

Type locality. Philippines (Mindanao Is.: Bukidnon Prov., Lindabon).

Distribution. Philippines: Bohol Is.; Luzon Is. (Bukidnon, Mt. Kiamo, Cabanglasan, Intavas, Kalatungan, Panamokan, San Fernando); Mindanao Is. (Davao del Sur, Mt. Apo) (Schultze 1922b; Cabras et al. 2017; Rukmane 2019a, 2019b; Rukmane-Bārbale and Cabras 2021).

Pachyrhynchus postpubescens confluens Janczyk, 1959

Type depository. Unknown.

Type locality. Philippines (Mindanao Is.).

Distribution. Only known from the type locality.

Pachyrhynchus pseudamabilis Yoshitake, 2012

Type depository. NIAES (holotype, paratypes).

Type locality. Philippines (Mindanao Is.: Mt. Apo).

Distribution. Philippines: Mindanao Is. (Mt. Apo; Agusan del Sur, Sibagat; Bukidnon, Cabanglasan, Panamokan, Mt. Kalatungan; Cotabato, Mt. Parke, Kidapawan; Davao del Sur, Davao City, Marilog District, Catigan, Toril, Calinan, Kapatagan; Lanao del Sur, Wao; Misamis Oriental, Mat-i, Claveria; Sarangani, Kiamba) (Cabras 2020; Rukmane-Bārbale and Cabras 2021; Cabras et al. 2022a).

Host plants. Theobroma cacao (Malvaceae), Croton leiophyllus (Euphorbiaceae), Piper aduncum (Piperaceae), and Lithocarpus boholensis (Fagaceae) (Cabras et al. 2022a).

Pachyrhynchus pseudhalconensis Rukmane, 2016

Type depository. DUBC (holotype, paratypes).

Type locality. Philippines (Mindoro Is.: Mindoro Oriental Prov., Puerto Galera).

Distribution. Philippines: Mindoro Is. (Mindoro Oriental Prov., Puerto Galera, Mt. Halcon, Baco).

Pachyrhynchus pseudoproteus Schultze, 1922

Type depository. SMTD (syntypes).

Type locality. Philippines (Luzon Is.: Laguna Prov.).

Distribution. Philippines: Luzon Is. (Laguna) (Schultze 1922b; Schultze 1924).

Remarks. Two specimens examined by Rukmane (2019c) were recorded from Leyte Island (Baybey). However, the distribution of the other specimens was in Luzon and so requires further verification.

Pachyrhynchus psittacinus Heller, 1912

Type depository. SMTD (holotype).

Type locality. Philippines (Luzon Is.: Bataan Prov., Lamao).

Distribution. Philippines: Luzon Is. (Bataan, Lamao; Mountain) (Schultze 1924; Rukmane-Bārbale and Cabras 2021).

Pachyrhynchus psittaculus Heller, 1921

Type depository. Unknown.

Type locality. Philippines (Luzon Is.: Quezon Prov., Mt. Banahaw).

Distribution. Philippines: Luzon Is. (Laguna; Quezon) (Schultze 1924).

Pachyrhynchus pulchellus Behrens, 1887

Pachyrhynchus bakeri Heller, 1921: 542. Synonymized by Schultze (1924: 338).

Pachyrhynchus pulchellus var. modestioroides Schultze, 1922b: 576. Synonymized by Schultze (1924: 338).

Type depository. SMTD (*).

Type locality. Philippines.

Distribution. Philippines: Luzon Is. (Benguet, Baguio, Mt. Polis, Mt. Santo Tomas, Trinidad, Mt. Trail, Atoc; Bulacan, Guinnisan; Ifugao; Mountain Prov.; Nueva Vizcaya; Manila) (Heller 1921; Schultze 1922b, 1924; Rukmane 2019b, 2019c; Rukmane-Bārbale and Cabras 2021).

Pachyrhynchus rebus Rukmane, 2016

Type depository. DUBC (holotype).

Type locality. Philippines (Luzon Is.: Quirino Prov., Nagtipunan).

Distribution. Only known from the type locality.

Pachyrhynchus regius regius Schultze, 1922

Type depository. SMTD (syntypes).

Type locality. Philippines (Leyte Is.: Cabalian).

Distribution. Philippines: Leyte Is. (mountains near Cabalian, Sogod); Mindanao Is. (Agusan del Norte, Sibagat; Bukidnon, Mt. Kiamo, Cabanglasan, San Fernando); Samar Is. (Marabot) (Schultze 1922b; Schultze 1923; Cabras et al. 2017; Rukmane-Bārbale and Cabras 2021).

Pachyrhynchus regius boronganus Schultze, 1934

Type depository. SMTD (syntypes).

Type locality. Philippines (Samar Is.: Eastern Samar, Borongan).

Distribution. Philippines: Samar Is. (​​Northern Samar; Samar; Eastern Samar, Borongan).

Pachyrhynchus reicherti Schultze, 1929

Type depository. SMTD (holotype).

Type locality. Philippines (Mindanao Is.: Davao Prov., Mt. Apo).

Distribution. Philippines: Mindanao Is. (Bukidnon; Cotabato; Davao del Sur, Mt. Apo); Mindoro Is. (Schultze 1929; Cabras and Medina 2018; Rukmane 2019b; Rukmane-Bārbale and Cabras 2021).

Pachyrhynchus reticulatus Waterhouse, 1841

Type depository. NHRS (syntype); SMTD (*).

Type locality. Philippines.

Distribution. Philippines: Catanduanes Is.; Luzon Is. (Abra; Aurora; Camarines Sur; Laguna, Lilio, Paete, Mt. Bonahao; Manila); Marinduque Is. (Boac) (Waterhouse 1841; Schultze 1923; Rukmane 2018c, 2019b, 2019c; Rukmane-Bārbale and Cabras 2021).

Pachyrhynchus rizali Schultze, 1934

Type depository. SMTD (syntypes).

Type locality. Philippines (Luzon Is.: Aurora Prov., Casiguran; Nueva Vizcaya Prov., Mount Dibago).

Distribution. Philippines: Luzon Is. (Aurora, Casiguran; Cagayan, Gonzaga, Sta. Ana; Nueva Vizcaya, Mount Dibago; Quirino) (Rukmane-Bārbale and Cabras 2021; Villegas et al. 2024).

Pachyrhynchus rochaorum Yoshitake, 2020

Type depository. NIAES (holotype, paratypes); KUM (paratypes).

Type locality. Philippines (Luzon Is.: Calabarzon region, Quezon Prov., Dolores, Brgy. Kinabuhayan, Mt. Banahaw).

Distribution. Only known from the type locality.

Pachyrhynchus roseomaculatus Waterhouse, 1841

Pachyrhynchus striatus Waterhouse, 1841: 25. Synonymized by Yoshitake (2019b: 193).

Type depository. NHMUK (holotype).

Type locality. Philippines.

Distribution. Philippines: Masbate Is. (Yoshitake 2019g).

Remarks. The specimen received by Schultze was obtained through an exchange, and he suggested that its location was Luzon Island (Schultze 1923). The only confirmed locality is Masbate Island, as examined by Yoshitake (2019g). Further research is needed to determine the distribution of this species.

Pachyrhynchus rufopunctatus Waterhouse, 1842

Type depository. NHMUK (holotype).

Type locality. Philippines.

Distribution. Philippines: Luzon Is. (Manila); Mindoro Is. (Mt. Halcon); Polillo Is. (Waterhouse 1842; Schultze 1924; Rukmane 2018c; Rukmane-Bārbale and Cabras 2021).

Pachyrhynchus rugicollis rugicollis Waterhouse, 1841

Type depository. Unknown.

Type locality. Philippines.

Distribution. Philippines: Luzon Is. (Aurora; Ifugao; Nueva Vizcaya, Sorsogon, Bulusan; Zambales Prov.; Battan Prov.; mountains of Mariveles; Manila) (Waterhouse 1841; Schultze 1923; Rukmane 2018c; Rukmane-Bārbale and Cabras 2021).

Pachyrhynchus rugicollis aurinius Heller, 1921

Type depository. Unknown.

Type locality. Philippines (Luzon Is.: Zambales Prov., Iba).

Distribution. Only known from the type locality.

Pachyrhynchus rugicollis crucifer Heller, 1912

Type depository. Unknown.

Type locality. Philippines (Luzon Is.).

Distribution. Only known from the type locality.

Pachyrhynchus rukmaneae rukmaneae Barševskis, 2016

Pachyrhynchus takakuwai Yoshitake, 2016b: 36. Synonymized by Yoshitake (2017c: 262).

Type depository. DUBC (holotype, paratypes).

Type locality. Philippines (Marinduque Is.: Boac).

Distribution. Philippines: Marinduque Is. (Buenavista, Mt. Malindig) (Barševskis 2016; Rukmane 2019b).

Remarks. Barševskis (2016) described this species based on a series of specimens from Marinduque Island, Philippines, including several other localities (e.g., Boac, Buenavista, and Mt. Malindig). However, Yoshitake and Yap (2017) considered that all materials of the species were collected from mountain areas, such as Sihi or Mt. Malindig, and suggested that the holotype locality, Boac, was mislabeled due to commercial processes. Yoshitake and Yap (2017) recommended that further studies were needed to clarify the distribution range of this species.

Pachyrhynchus rukmaneae paucisignatus Yoshitake, 2017

Type depository. KUM (holotype, paratypes); NIAES (paratypes).

Type locality. Philippines (Marinduque Is.: Boac).

Distribution. Philippines: Marinduque Is. (Boac) (Yoshitake and Yap 2017; Rukmane-Bārbale and Cabras 2021).

Pachyrhynchus sagittatus Rukmane, 2019

Type depository. DUBC (holotype).

Type locality. Philippines (Luzon Is.: Quirino Prov., Sierra Madre).

Distribution. Philippines: Luzon Is. (Quirino) (Rukmane 2019d, Rukmane-Bārbale and Cabras 2021).

Pachyrhynchus sakaii Yoshitake, 2017

Type depository. NIAES (holotype).

Type locality. Philippines (Samar Is.).

Distribution. Only known from the type locality (Yoshitake 2017c).

Pachyrhynchus samarensis Schultze, 1923

Type depository. Unknown.

Type locality. Philippines (Samar Is.: Catarman).

Distribution. Philippines: Samar Is. (Northern Samar, Catarman, Lope De Vega; Samar) (Rukmane 2019a).

Pachyrhynchus sanchezi Heller, 1912

Type depository. SMTD (holotype).

Type locality. Philippines (Luzon Is.: Benguet Prov., Baguio).

Distribution. Philippines: Luzon Is. (Auror; Benguet, Baguio, Mt. Mirador, Mt. Santo Tomas, Mt. Trail; Ifugao; Isabela; Mountain Prov.; Nueva Vizcaya) (Schultze 1924; Heller 1912; Rukmane-Bārbale and Cabras 2021).

Remarks. Two specimens examined by Rukmane (2019b) were recorded from Mindanao Island (Surigao). However, other specimens were found in Luzon. It is likely that the two Mindanao specimens were mislabeled.

Pachyrhynchus sarcitis sarcitis Behrens, 1887

Type depository. Unknown.

Type locality. Philippines.

Distribution. Philippines: Babuyan Islands. (Calayan Is.; Camiguin Is.; Fuga Is; Babuyan Is.); Batanes Islands (Batan Is.; Sabtang Is.; Itbayat Is.); Luzon Is. (Cagayan; Isabela; Quirino) (Schultze 1924; Rukmane 2019b; Rukmane-Bārbale and Cabras 2021; Tseng et al. 2021).

Host plants. Philippine Leea, Leea philippinensis (Leeaceae) and Manila Leea, Leea guineensis (Leeaceae) (Tseng et al. 2021).

Pachyrhynchus sarcitis kotoensis Kôno, 1930

Type depository. HUMS (syntypes).

Type locality. Taiwan (Taitung County, Lanyu Is. (= Kotosho)).

Distribution. Taiwan: Lanyu Is.; Ludao Is.

Host plants. Philippine Leea, Leea philippinensis (Leeaceae) and Manila Leea, Leea guineensis (Leeaceae) (this study).

Remarks. In a captive environment, the number of larval stages ranged from four to six, and the duration from eggs to adults varied between 133–169 days (Huang et al. 2018). A larva was found inside the stem of L. guineensis in Lanyu (this study). This species is relatively rarer on Ludao Is. than on Lanyu Is. (this study).

Pachyrhynchus schoenherri Waterhouse, 1841

Type depository. NHMUK (holotype).

Type locality. Philippines.

Distribution. Philippine: Leyte Is. (Mahaplag) (Waterhouse 1841; Rukmane-Bārbale and Cabras 2021).

Pachyrhynchus schuetzei Schultze, 1917

Type depository. SMTD (syntypes).

Type locality. Philippines (Luzon Is.: Benguet Prov., Haight’s Place).

Distribution. Philippines: Luzon Is. (Benguet, Loo, Mt. Pawai; Ifugao; Mountain Prov.; Nueva Vizcaya) (Schultze 1924; Rukmane 2019c; Rukmane-Bārbale and Cabras 2021).

Pachyrhynchus semiignitus Schultze, 1922

Type depository. SMTD (holotype).

Type locality. Philippines (Mindanao Is.: Cotabato Prov., Pikit).

Distribution. Only known from the type locality (Schultze 1922b).

Pachyrhynchus semperi Heller, 1912

Type depository. SMTD (syntypes).

Type locality. Philippines.

Distribution. Philippines: Babuyan Is. (Camiguin Norte; Calayan Is.; Babuyan Is.); Batanes Is. (Batan Is.) (Rukmane 2019b; Rukmane-Bārbale 2020b; Rukmane-Bārbale and Cabras 2021; Tseng et al. 2021).

Host plants. Bishop Wood Tree, Bischofia javanica (Euphorbiaceae) (Tseng et al. 2021).

Pachyrhynchus septentrionalis Yoshitake, 2017

Type depository. NIAES (holotype, paratypes); CFS, KUM, MBLI (paratypes).

Type locality. Philippines (Luzon Is.: Cagayan Prov., Sierra Madre Mountains).

Distribution. Philippines: Luzon Is. (Cagayan Vally region, Aurora, Cagayan) (Yoshitake 2017d; Rukmane-Bārbale and Cabras 2021).

Pachyrhynchus sergejevae Rukmane, 2018

Type depository. DUBC (holotype, paratypes).

Type locality. Philippines (Mindanao Is.: Bukidnon Prov., Cabanglasan).

Distribution. Philippines: Mindanao Is. (Bukidnon, Mt. Kalatungan, Intavas) (Rukmane 2018f).

Pachyrhynchus shavrini Rukmane & Barševskis, 2016

Type depository. DUBC (holotype, paratypes).

Type locality. Philippines (Samar Is.: Northern Samar Prov., Lope De Vega).

Distribution. Philippines: Samar Is. (Northern Samar, Lope De Vega; Samar, Hinabangan) (Rukmane-Bārbale and Cabras 2021).

Remarks. One specimen examined by Rukmane (2019b) was recorded from Luzon Island. However, the distribution of other specimens was in Samar. The distribution of P. shavrini requires further verification.

Pachyrhynchus signaticollis Schultze, 1922

Pachyrhynchus transversarius Heller, 1923b: 8. Synonymized by Schultze (1923: 666).

Type depository. SMTD (syntypes).

Type locality. Philippines (Mindanao Is.: Agusan Prov., Butuan).

Distribution. Philippines: Mindanao Is. (Agusan Prov., Butuan; Surigao) (Schultze 1922a; Schultze 1923; Rukmane-Bārbale and Cabras 2021).

Pachyrhynchus signatus Schultze, 1919

Type depository. SMTD (holotype).

Type locality. Philippines (Siargao Is.).

Distribution. Only known from the type locality.

Pachyrhynchus similis Bollino, 2022

Type depository. SMTD (holotype); MBLI, DUBC, CFS (paratypes).

Type locality. Philippines (Mindanao Is.: Lanao del Sur Prov., Near Wao).

Distribution. Philippines: Mindanao Is. (south eastern Lanao del Sur; north eastern North Cotabato; Bukidnon, west of Mount Kitanglad).

Pachyrhynchus smaragdinus Behrens, 1887

Pachyrhynchus smaragdinus var. carnosus Kraatz, 1888: 32. Synonymized by Schultze (1924: 313).

Pachyrhynchus smaragdinus var. purpurascens Kraatz, 1888: 32. Synonymized by Schultze (1924: 313).

Type depository. SMTD (syntype).

Type locality. Philippines.

Distribution. Philippines: Bohol Is.; Mindanao Is. (Bukidnon, Misamis, Surigao); Samar Is. (Northern Samar, Lope De Vega; Samar, Hinabangan) (Rukmane 2019b; Rukmane-Bārbale and Cabras 2021).

Pachyrhynchus sonani Kôno, 1930

Type depository. HUMS (holotype).

Type locality. Taiwan (Taitung County, Lanyu Is. (= Kotosho)).

Distribution. Taiwan: Lanyu Is.

Host plants. Among all the individuals found on the host plants, 93.75% were found on Sea poison tree (Barringtonia asiatica, Lecythidaceae), 3.13% on beef wood (Casuarina equisetifolia, Casuarinaceae) and 3.13% on Indian almond (Terminalia catappa, Combretaceae) (Chen et al. 2017).

Remarks. One female was observed to lay eggs in the xylem of B. asiatica, and the larvae were also found in the decayed xylem of the same host plant (Hsu et al. 2017).

Pachyrhynchus speciosus Waterhouse, 1841

Pachyrhynchus absurdus Schultze, 1919: 550. Synonymized by Schultze (1923: 657) and Rukmane-Bārbale (2020a: 35).

Type depository. NHMUK (syntype), NHRS (syntype), SMTD (syntypes).

Type locality. Philippines.

Distribution. Philippines: Bohol Is. (Bilar); Bucas Grande Is.; Dinagat Is.; Leyte Is.; Mindanao Is. (Bukidnon, Cabanglasan; Cotabato, Saob; Davao del Sur, Mt. Apo, Marilog District, Catigan, Toril, Davao City; Sarangani, Kiamba; Surigao Prov., Surigao, Tandag); Samar Is. (Northern Samar; Samar, Hinabangan); Siargao Is. (Waterhouse 1841; Schultze 1919, 1923; Háva and Rukmane 2018; Rukmane 2019a, b; Rukmane-Bārbale and Cabras 2021; Cabras et al. 2022a).

Host plants. This species was collected on the leaves of Melastoma malabathricum (Melastomataceae) and Medinilla cumingii (Melastomataceae) (Cabras et al. 2022a).

Pachyrhynchus sphaericollaris Schultze, 1923

Type depository. SMTD (holotype).

Type locality. Philippines (Luzon Is.: Kalinga Prov., Pinukpuk).

Distribution. Philippines: Luzon Is. (Cagayan; Isabela, Pinablanca; Kalinga) (Rukmane 2019c; Rukmane-Bārbale and Cabras 2021).

Pachyrhynchus sphenomorphoides Yoshitake, 2012

Type depository. NIAES (holotype, paratypes).

Type locality. Philippines (Basilan Is.: near Zamboanga Peninsula of West Mindanao).

Distribution. Philippines: Basilan Is.; Mindanao Is. (Zamboanga del Norte Prov.) (Bollino 2022).

Pachyrhynchus stellio Heller, 1912

Type depository. SMTD (syntypes).

Type locality. Philippines (Luzon Is.: Bataan Prov., Lamao).

Distribution. Philippines: Luzon Is. (Aurora, Bataan Prov., Lamao, Mt. Mariveles; Benguet, Isabela, Mountain, Nueva Vizcaya, Rizal) (Schultze 1923; Rukmane-Bārbale and Cabras 2021).

Pachyrhynchus subamabilis Yoshitake, 2012

Type depository. NIAES (holotype).

Type locality. Philippines (Mindanao Is.: Mt. Apo).

Distribution. Philippines: Mindanao Is. (Bukidnon; Cotabato; Lanao del Sur) (Rukmane 2019b; Rukmane-Bārbale and Cabras 2021).

Pachyrhynchus subanon Bollino, 2022

Type depository. SMTD (holotype); MBLI (paratypes).

Type locality. Philippines (Mindanao Is.: Misamis Occidental Prov., Brgy. Gandawan, Mt. Malindang Range, Lake Duminagat-Mt. Ginlajan).

Distribution. Only known from the type locality.

Pachyrhynchus subgo Bollino, 2025

Type depository. SMTD (holotype); MBLI, DHA (paratypes).

Type locality. Philippines (Cebu Is.: Cebu Prov., Mt. Manunggal).

Distribution. Philippines: Cebu Is. (Cebu Prov., Balamban, Mt. Manunggal, Brgy. Sunog, Mt. Mauyog, hills north-east of Kaluangan) (Bollino and Ersoy 2025).

Pachyrhynchus subpalidius Rukmane-Bārbale, 2022

Type depository. DUBC (holotype, paratypes).

Type locality. Philippines (Luzon Is.: Mountain Prov., Barlig, Bontoc).

Distribution. Only known from the type locality.

Pachyrhynchus sulphureomaculatus Schultze, 1922

Type depository. SMTD (holotype).

Type locality. Philippines (Mindanao Is.: Cotabato Prov., Cotabato).

Distribution. Philippines: Mindanao Is. (Agusan; Bukidnon, Mt. Kiamo; Cotabato, Kidapawan, Mt. Parker; Davao City, Marilog District, Mt. Apo; Sarrangani, Kiamba; Surigao del Sur, Esperanza, San Miguel, Tandag); Leyte Is. (Sogod); Samar Is. (Hinabangan) (Schultze 1922b; Cabras et al. 2017, 2022a; Rukmane-Bārbale and Cabras 2021).

Host plant. Lithocarpus boholensis (Fagaceae) (Cabras et al. 2022a).

Pachyrhynchus sumptuosoides Yoshitake, 2017

Type depository. KUM (holotype, paratypes), NIAES (paratypes).

Type locality. Philippines (Luzon Is.: Apayao Prov., Conner).

Distribution. Philippines: Luzon Is. (Cordillera Administrative Region; Apayao, Conner; Cagayan, Santa Ana; Kalinga, Pinukpuk, Laguna) (Yoshitake 2017c; Rukmane-Bārbale 2020b; Rukmane-Bārbale and Cabras 2021).

Pachyrhynchus sumptuosus Schultze, 1917

Type depository. SMTD (syntypes).

Type locality. Philippines (Luzon Is.: Mountain Prov., Bontoc).

Distribution. Philippines: Luzon Is. (Apayao; Ilocos Sur, Cabagao; Kalinga, Lubuagan, Balbalan; Mountain Prov., Bontoc; Nueva Vizcaya) (Schultze 1924; Rukmane 2019c; Rukmane-Bārbale and Cabras 2021).

Pachyrhynchus tadauchii Yoshitake, 2012

Type depository. NIAES (holotype, paratypes).

Type locality. Philippines (Mindanao Is.: Surigao del Sur Prov., Bislig).

Distribution. Philippines: Mindanao Is. (Agusan; Agusan del Sur, San Francisco; Bukidnon, Cabanglasan; Davao del Sur, Mt. Apo, Mt. Matutum; Sarangani, Kiamba, Malungon, Maitum; Surigao del Sur, San Miguel, Tandag) (Rukmane 2019a; Rukmane-Bārbale and Cabras 2021).

Pachyrhynchus taylori taylori Schultze, 1922

Type depository. SMTD (syntypes).

Type locality. Philippines (Luzon Is.: Kalinga Prov., Balbalan).

Distribution. Philippines: Luzon Is. (Kalinga; Nueva Vizcaya; Ifugao) (Schultze 1922a; Schultze 1924; Rukmane-Bārbale and Cabras 2021).

Pachyrhynchus taylori metalescens Schultze, 1924

Type depository. SMTD (syntypes).

Type locality. Philippines (Luzon Is.: Ifugao Prov., Polis Pass).

Distribution. Only known from the type locality.

Pachyrhynchus tetramaculatus Rukmane, 2019

Type depository. DUBC (holotype, paratypes).

Type locality. Philippines (Luzon Is.: Nueva Vizcaya Prov., Quezon).

Distribution. Only known from the type locality (Rukmane 2019d).

Pachyrhynchus tikoi Rukmane, 2016

Type depository. DUBC (holotype, paratypes).

Type locality. Philippines (Mindanao Is.: Bukidnon Prov., Cabanglasan).

Distribution. Philippines: Mindanao Is. (Bukidnon, Mt. Apo, Cabanglasan) (Rukmane-Bārbale and Cabras 2021).

Pachyrhynchus tilikensis Bollino & Sandel, 2015

Type depository. SMTD (holotype); MBLI, CFS (paratypes).

Type locality. Philippines (Lubang Is.: south-west of Tilik).

Distribution. Only known from the type locality.

Pachyrhynchus tobafolius Kano, 1929

Type depository. Unknown.

Type locality. Taiwan (Taitung County, Lanyu Is. (= Kotosho)).

Distribution. Taiwan: Lanyu Is., Laudao Is.

Host plants. Pipturus arborescens (Urticaceae).

Remarks. Pachyrhynchus tobafolius is only distributed on Lanyu and Ludao islands. However, according to Rukmane (2019b), examination of specimens from NHRS revealed that two specimens were mislabeled with the collection site listed as Japan. Kano (1929) mentioned that P. tobafolius feeds on the leaves of Barringtonia asiatica on the seashore. However, during our field observations, we found that this species mainly feeds on plants of the Urticaceae family (this study).

Pachyrhynchus torresi Rukmane, 2018

Type depository. DUBC (holotype, paratypes).

Type locality. Philippines (Mindanao Is.: Zamboanga Prov., Labuan).

Distribution. Philippines: Mindanao Is. (Zamboanga, Labuan) (Rukmane 2018f, 2019b; Rukmane-Bārbale and Cabras 2021).

Pachyrhynchus tristis Heller, 1912

Type depository. NHMUK (syntypes), SDEI (syntypes), SMTD (syntypes).

Type locality. Philippines (Luzon Is.).

Distribution. Philippines: Luzon Is. (Benguet; Moutain Prov.) (Schultze 1924; Rukmane 2019b; Rukmane-Bārbale 2020b).

Pachyrhynchus valainisi Rukmane & Barševskis, 2016

Type depository. DUBC (holotype, paratype).

Type locality. Philippines (Mindoro Is.: Mindoro Oriental Prov., Puerto Galera).

Distribution. Philippines: Mindoro Is. (Mindoro Oriental Prov., Puerto Galera, Baco, Mt. Halcon) (Rukmane-Bārbale and Cabras 2021).

Pachyrhynchus venustus venustus Waterhouse, 1841

Pachyrhynchus venustus var. aureomaculatus Kraatz, 1888: 27. Synonymized by Schultze (1924: 310).

Pachyrhynchus virgatus Schultze, 1919: 549. Synonymized by Schultze (1924: 310).

Type depository. NHMUK (syntype), SMTD (syntypes).

Type locality. Philippines.

Distribution. Philippines: Mindanao Is. (Cotabato; Surigao Prov., Surigao; Sarangani); Samar Is. (Northern Samar, Lope De Vega; Samar, Hinabangan, Marabot); Leyte Is. (Sogod) (Waterhouse 1841; Schultze 1919; Rukmane-Bārbale and Cabras 2021).

Remarks. Six specimens examined by Rukmane (2019b) were recorded from Luzon Island (Manila). However, other specimens were found from Mindanao PAIC and the distribution of P. venustus requires further verification.

Pachyrhynchus venustus insulanus Schultze, 1919

Type depository. SMTD (syntypes).

Type locality. Philippines (Siargao Is.; Bucas Grande Is.).

Distribution. Only known from the type locality.

Pachyrhynchus viridans Heller, 1912

Type depository. SMTD (holotype).

Type locality. Philippines (Calayan).

Distribution. Philippines: Calayan Is.; Negros Is.; Sibuyan Is. (Heller 1912; Schultze 1924; Rukmane 2019b).

Pachyrhynchus viridis (Chevrolat, 1879)

Apocyrtus viridis Chevrolat, 1879: 134.

Sphenomorpha viridis: Dalla Torre et al., 1931: 19.

Pachyrhynchus viridis: Yoshitake, 2017a: 243.

Type depository. NHRS (holotype).

Type locality. Indonesia (Dorey).

Distribution. Indonesia: Dorey Is.; West Papua (Manokwari Prov., Ransiki District, Neney village) (Yoshitake 2017a; Bollino 2023).

Pachyrhynchus yoshitakei Bollino & Rukmane, 2020

Type depository. SMTD (holotype); MBLI (paratypes).

Type locality. Philippines (Mindanao Is.: Misamis Occidental Prov., Oroquieta City, Mt. Malindang Range, Mt. Capole-Sebucal).

Distribution. Philippines: Mindanao Is. (Misamis Occidental, Mt. Malindang Range).

Pachyrhynchus yoshitakeorum Yoshitake, Bollino & Sandel, 2019

Type depository. NIAES (holotype, paratypes); KUM, CFS, MBLI (paratypes).

Type locality. Philippines (Bohol Is.: Bohol Prov., Duero, Brgy. Payao, Sitio Pangpang).

Distribution. Philippines: Bohol Is.; Dinagat Is.; Cebu Is. (Rukmane 2019a; Yoshitake et al. 2019b; Rukmane-Bārbale and Cabras 2021; Bollino and Ersoy 2025).

Host plants. This species is often found on cacao (Theobroma cacao, Malvaceae) and coffee (Coffea arabica, Rubiaceae) trees (Yoshitake et al. 2019b).

Pachyrhynchus yukae Yoshitake, 2019

Type depository. NIAES (holotype, paratypes); MCKUM (paratypes).

Type locality. Philippines (Luzon Is.: Quezon Prov., Dolores, Barangay Kinabuhayan, Sitio Bangkong, Kahoy).

Distribution. Only known from the type locality.

Pachyrhynchus yuukae Yoshitake, 2019

Type depository. NIAES (holotype, paratypes).

Type locality. Philippines (Luzon Is.: Nueva Ecija Prov., Carranglan, Lohong).

Distribution. Only known from the type locality (Yoshitake 2019d).

Pachyrhynchus zamboanganus Yoshitake, 2012

Type depository. NIAES (holotype).

Type locality. Philippines (Mindanao Is.: Zamboanga del Norte Prov.).

Distribution. Only known from the type locality.

Pachyrhynchus zebra Schultze, 1917

Type depository. SMTD (holotype).

Type locality. Philippines (Luzon Is.: Benguet Prov., Mt. Santo Tomas).

Distribution. Philippines: Luzon Is. (Aurora; Benguet, Baguio, Mt. Santo Tomas; Rizal) (Schultze 1923; Rukmane 2019b; Rukmane-Bārbale and Cabras 2021).

Discussion

As the number of described Pachyrhynchus species has increased in recent years, it is vital to provide an updated checklist of species. Such compilations are integral to future faunistic studies as these lists can provide an overview of the species richness of a particular taxa with their natural history and distribution patterns. The present work provides an updated checklist of Pachyrhynchus species, including their host plants, geographic records, synonyms, type repositories, and species groups.

Compared to earlier works on Pachyrhynchus (e.g., Heller 1912; Schultze 1922a, 1923), recent publications incorporate host plant information (see Chen et al. 2017; Cabras et al. 2021, 2022a; Tseng et al. 2021). This information is crucial for understanding the feeding ecology of Pachyrhynchus species. In this account, 19 Pachyrhynchus species are provided with host plant records. Despite the limited number of records, they reflect the diversity of host plant taxa utilized by Pachyrhynchus species. A total of 26 families, 32 genera and 35 host plant species were recorded, including species from the genera Ardisia (Primulaceae), Barringtonia (Lecythidaceae), Bischofia (Euphorbiaceae), Lithocarpus (Fagaceae), Melastoma (Melastomataceae), Philodendron (Araceae), and Saurauia (Actinidiaceae) (Cabras and Yoshitake 2016; Chen et al. 2017; Hsu et al. 2017; Tseng et al. 2021; Cabras et al. 2022a). These relatively few records of host plant associations highlight significant knowledge gaps in Pachyrhynchus faunistic studies that need to be addressed.

We compared species group classifications of Pachyrhynchus weevils based on both morphological and molecular studies (Table 2). Morphology-based groupings were derived from three key sources: Heller (1912), Schultze (1923, 1924), and more recent taxonomic works published between 2017 and 2025. Molecular clades were based on Van Dam et al. (2023) and the phylogenetic framework from Chen et al. (2025). The comparison revealed that several morphological groups are consistent with molecular clades, suggesting concordance between traditional taxonomy and modern phylogenetic inference. For instance, P. amabilis was assigned to Group XIV by Schultze (1924) and to the amabilis species group in recent morphological taxonomy (Bollino et al. 2017, 2020), which aligns with the amabilis clade in Chen et al. (2025). Similarly, P. anellifer and P. annulatus consistently fall under Group VII (Heller 1912; Schultze 1924), and both are placed in the erichsoni clade based on molecular data (Van Dam et al. 2023; Chen et al. 2025). The orbifer clade, as defined in Chen et al. (2025), includes at least 19 species, such as P. orbifer, P. cruciatus, P. jugifer, P. moniliferus, P. speciosus, and P. regius. Most of these species were assigned to Group I or Group V based on the morphological classifications by Heller (1912) and Schultze (1923, 1924). This indicates a moderate level of consistency between traditional morphological groupings and recent molecular phylogenetic results. However, discrepancies were also noted. Some recently described species lack morphological group assignments in earlier literature but have been placed into molecular clades. Future studies focusing on a broader-scale comparison between morphological characters and molecular data are needed to clarify their taxonomic placement.

Distribution records are a vital element in Pachyrhynchus research, particularly regarding biogeography, insular speciation, and the evolution of mimicry systems. These records reveal spatial patterns such as co-occurrences and endemism. The Philippine archipelago is considered the center of diversity for the genus Pachyrhynchus, as the majority of described species are endemic to the country. Zoological demarcations of island groups in the Philippines are based on the Pleistocene Aggregate Island Complex (PAIC) model (Heaney 1986; Brown et al. 2013). Described Pachyrhynchus species in the Philippines primarily occur in the Greater Luzon Faunal Region and the Greater Mindanao Faunal Region. In the Greater Luzon Faunal Region, which encompasses mainland Luzon, Catanduanes, Polillo, Marinduque, and small surrounding islands, there are 82 known species. This region is the most species-rich, accounting for 45.8% of the described Pachyrhynchus species. Among these 82 species, 70 are endemic to this area. In the Greater Mindanao Faunal Region, which includes mainland Mindanao, Samar, Leyte, Biliran, Bohol, Dinagat, Siargao, Basilan, and small surrounding islands, there are 70 reported species, accounting for 39.1% of the described species (Fig. 2, Table 1), and 65 of them being endemic to this region.

Table 1.

List of Pachyrhynchus species and their distribution regions. The letter ‘v’ indicates potential distribution areas mentioned in the literature. The abbreviations for distribution regions are as followed: (1) Greater Luzon PAIC (GL), (2) Greater Mindoro PAIC (GMDO), (3) Greater Negros-Panay PAIC (GNP), (4) Greater Mindanao PAIC (GMDA), (5) Lubang Island (LUB), (6) Sibuyan Island (SBY), (7) Islands north of mainland Luzon (INL), and (8) Indonesia archipelago (IND).

Species Distribution regions
(1) GL (2) GMDO (3) GNP (4) GMDA (5) LUB (6) SBY (7) INL (8) IND
1 P. amabilis Schultze, 1922 v
2 P. anellifer Heller, 1912 v
3 P. anichtchenkoi Rukmane & Barševskis, 2016 v
4 P. annulatus Chevrolat, 1881 v
5 P. antonkozlovi Rukmane & Barševskis, 2016 v
6 P. apicatus Schultze, 1922 v v
7 P. apocyrtoides Schultze, 1922 v
8 P. apoensis Yoshitake, 2012 v
9 P. ardentius Schultze, 1919 v
10 P. argus Pascoe, 1873 v
11 P. atrocyaneus Schultze, 1922 v
12 P. atronitens Yoshitake, 2019 v
13 P. baluganus Schultze, 1924 v
14 P. banglas Bollino, Sandel & Rukmane, 2017 v
15 P. barsevskisi Rukmane, 2016 v
16 P. basilanus Heller, 1923 v
17 P. benguetanus Schultze, 1924 v
18 P. bollinoi Rukmane-Bārbale, 2020 v
19 P. bucasanus Schultze, 1922 v
ssp. ornatus Schultze, 1934 v
20 P. cabrasae Rukmane & Barševskis, 2016 v
21 P. caeruleovittatus Yoshitake, 2012 v
22 P. caeruleus Yoshitake, 2019 v
23 P. cagayanus Heller, 1929 v
24 P. callainimaculatus Yoshitake, 2017 v
25 P. cebrem Patano & Rukmane-Bārbale, 2022 v
26 P. chamissoi Schultze, 1922 v v
27 P. chlorites Chevrolat, 1881 v v
P. chlorites ssp. insularis Kano, 1929 v
28 P. chrysocyaneus Rukmane-Bārbale, 2024 v
29 P. chrysomaculatus Bollino, 2022 v
30 P. cinereomaculatus Rukmane-Bārbale, 2020 v
31 P. cingulatus Pascoe, 1873 v
32 P. circulatus Heller, 1912 v
33 P. circulimaculatus Yoshitake, 2019 v
34 P. conformis Yoshitake, 2017 v
35 P. confusus Schultze, 1923 v
36 P. congestus Pascoe, 1873 v
P. congestus ssp. aedamlayroni Rukmane, 2019 v
P. congestus ssp. coerulans Kraatz, 1888 v
P. congestus ssp. immarginatus Kraatz, 1888 v
P. congestus ssp. mirabilis Yoshitake, 2017 v
P. congestus ssp. ocellatus Schultze, 1924 v
P. congestus ssp. pavonius Heller, 1921 v
37 P. consobrinus Schultze, 1922 v
38 P. corpulentus Schultze, 1922 v
P. corpulentus ssp. balatukan Patano & Macalaba, 2023 v
39 P. croesus Oberthur, 1879 v
40 P. cruciatus Schultze, 1923 v
41 P. cumingii Waterhouse, 1841 v
P. cumingii ssp. boholensis Schultze, 1924 v
42 P. davaoensis Schultze, 1934 v
43 P. decussatus Waterhouse, 1841 v
P. decussatus ssp. catanduanensis Rukmane-Bārbale, 2020 v
44 P. disargus Rukmane, 2019 v
45 P. disgestus Heller, 1929 v
46 P. dohrni Behrens, 1887 v
47 P. domino Rukmane, 2016 v
48 P. dubiosus Schultze, 1922 v
49 P. elegans Waterhouse, 1842 v
50 P. eques Heller, 1912 v
51 P. equester Heller, 1929 v
52 P. erichsoni Waterhouse, 1841 v v v
P. erichsoni ssp. eschscholtzii Waterhouse, 1841 v
53 P. erosus Schultze, 1920 v
54 P. esperanza Bollino, Sandel & Rukmane, 2017 v
55 P. faisali Bollino, 2023 v
56 P. felipeae Rukmane & Cabras, 2018 v
57 P. florulentus Yoshitake, 2019 v
58 P. forsteni Vollenhoven, 1864 v
59 P. franciscoi Rukmane & Cabras, 2018 v
60 P. galeraensis Schultze, 1934 v
61 P. gemmatus Waterhouse, 1841 v
P. gemmatus ssp. purpureus Kraatz, 1888 v
62 P. gilvomaculatus Yoshitake, 2017 v
63 P. gloriosus Faust, 1895 v
P. gloriosus ssp. abbreviatus Schultze, 1922 v
64 P. halconensis Schultze, 1922 v
65 P. helenperrinae Rukmane, 2018 v
66 P. helleri Kuntzen, 1914 v
67 P. hirokii Yoshitake, 2012 v
68 P. igorota Schultze, 1917 v
69 P. ilgas Rukmane, 2017 v
70 P. imitans Rukmane & Bollino, 2020 v
71 P. inclytus Pascoe, 1873 v
72 P. infernalis Fairmaire, 1897 v
73 P. jitanasaius Chen & Lin, 2017 v
74 P. jugifer Waterhouse, 1841 v
75 P. kirklayroni Rukmane, 2019 v
76 P. kraslavae Rukmane & Barševskis, 2016 v
77 P. lacunosus Heller, 1912 v
78 P. latifasciatus Waterhouse, 1842 v
79 P. layroni Rukmane & Cabras, 2018 v
80 P. libucanus Schultze, 1923 v
81 P. loheri Schultze, 1917 v
82 P. lorquini Chevrolat, 1881 v
83 P. lubanganus Bollino & Sandel, 2015 v
84 P. marinduquensis Rukmane & Barševskis, 2016 v
85 P. maruyamai Yoshitake, 2019 v
86 P. masatoshii Yoshitake & Yap, 2017 v
87 P. masatoshiensis Rukmane-Bārbale, 2024 v
88 P. miltoni Cabras & Rukmane, 2016 v
89 P. mindoroensis Rukmane & Háva, 2020 v
90 P. mohagani Bollino & Sandel, 2015 v
91 P. möllendorffi Heller, 1899 v
P. möllendorffi ssp. marinduquanus Rukmane & Cabras, 2018 v
92 P. moniliferus Germar, 1824 v v v v v
P. moniliferus ssp. abranus Heller, 1934 v v
P. moniliferus ssp. babuyanensis Rukmane, 2018 v
P. moniliferus ssp. chevrolati Eydoux & Souleyet, 1839 v v
P. moniliferus ssp. herbidus Rukmane, 2018 v
P. moniliferus ssp. jagori Heller, 1912 v
P. moniliferus ssp. stellulifer Heller, 1912 v v
93 P. morio Heller, 1912 v
94 P. morotaiensis Vollenhoven, 1864 v
95 P. multipunctatus Waterhouse, 1841 v v
P. multipunctatus ssp. endoi Yoshitake, 2018 v
96 P. naokii Yoshitake, 2012 v
97 P. naokoae Yoshitake, 2019 v
98 P. negrosensis Schultze, 1924 v
99 P. neoabsurdus Rukmane, 2017 v
100 P. niisatoi Yoshitake, 2017 v
101 P. nitcisi Rukmane & Barševskis, 2016 v
102 P. nobilis Heller, 1912 v v
P. nobilis ssp. yamianus Kano, 1929 v
103 P. noeli Yoshitake, 2019 v
104 P. notocruciatus Yoshitake, 2017 v
105 P. obumanuvu Cabras, Donato, Medina, & Van Dam, 2021 v
106 P. occidentalis Rukmane, 2017 v
107 P. ochroplagiatus Heller, 1912 v
P. ochroplagiatus ssp. multiplagiatus Schultze, 1924 v
108 P. octoannulatus Yoshitake, Bollino & Sandel, 2019 v
109 P. ohbayashii Yoshitake, 2017 v
110 P. orbifer Waterhouse, 1841 v
P. orbifer ssp. ardens Chevrolat, 1841 v
P. orbifer ssp. azureus Schultze, 1922 v
P. orbifer ssp. callainus Yoshitake, 2017 v
P. orbifer ssp. circulifer Chevrolat, 1841 v
P. orbifer ssp. gemmans Chevrolat, 1841 v
P. orbifer ssp. inornatus Waterhouse, 1841 v v
P. orbifer ssp. murinus Heller, 1934 v
P. orbifer ssp. striatomaculatus Yoshitake, 2017 v
111 P. orientalis Rukmane, 2017 v
112 P. ottomerkli Rukmane, 2019 v
113 P. panumanon Cabras & Medina, 2022 v
114 P. perpulcher Waterhouse, 1841 v v
115 P. phaleratus Waterhouse, 1841 v
P. phaleratus ssp. badiovittatus Yoshitake, 2019 v
P. phaleratus ssp. dannylayroni Rukmane, 2019 v
116 P. pinorum Pascoe, 1873 v
P. pinorum ssp. transversalis Heller, 1912 v
117 P. postpubescens Schultze, 1922 v v
P. postpubescens ssp. confluens Janczyk, 1959 v
118 P. pseudamabilis Yoshitake, 2012 v
119 P. pseudhalconensis Rukmane, 2016 v
120 P. pseudoproteus Schultze, 1922 v
121 P. psittacinus Heller, 1912 v
122 P. psittaculus Heller, 1921 v
123 P. pulchellus Behrens, 1887 v
124 P. rebus Rukmane, 2016 v
125 P. regius Schultze, 1922 v
P. regius ssp. boronganus Schultze, 1934 v
126 P. reicherti Schultze, 1929 v
127 P. reticulatus Waterhouse, 1841 v
128 P. rizali Schultze, 1934 v
129 P. rochaorum Yoshitake, 2020 v
130 P. roseomaculatus Waterhouse, 1841 v
131 P. rufopunctatus Waterhouse, 1842 v v
132 P. rugicollis Waterhouse, 1841 v
P. rugicollis ssp. aurinius Heller, 1921 v
P. rugicollis ssp. crucifer Heller, 1912 v
133 P. rukmaneae Barševskis, 2016 v
P. rukmaneae ssp. paucisignatus Yoshitake, 2017 v
134 P. sagittatus Rukmane, 2019 v
135 P. sakaii Yoshitake, 2017 v
136 P. samarensis Schultze, 1923 v
137 P. sanchezi Heller, 1912 v
138 P. sarcitis Behrens, 1887 v v
P. sarcitis ssp. kotoensis Kôno, 1930 v
139 P. schoenherri Waterhouse, 1841 v
140 P. schuetzei Schultze, 1917 v
141 P. semiignitus Schultze, 1922 v
142 P. semperi Heller, 1912 v
143 P. septentrionalis Yoshitake, 2017 v
144 P. sergejevae Rukmane, 2018 v
145 P. shavrini Rukmane & Barševskis, 2016 v
146 P. signaticollis Schultze, 1922 v
147 P. signatus Schultze, 1919 v
148 P. similis Bollino, 2022 v
149 P. smaragdinus Behrens, 1887 v
150 P. sonani Kôno, 1930 v
151 P. speciosus Waterhouse, 1841 v
152 P. sphaericollaris Schultze, 1923 v
153 P. sphenomorphoides Yoshitake, 2012 v
154 P. stellio Heller, 1912 v
155 P. subamabilis Yoshitake, 2012 v
156 P. subanon Bollino, 2022 v
157 P. subpalidius Rukmane-Bārbale, 2022 v
158 P. sugbo Bollino, 2025 v
159 P. sulphureomaculatus Schultze, 1922 v
160 P. sumptuosoides Yoshitake, 2017 v
161 P. sumptuosus Schultze, 1917 v
162 P. tadauchii Yoshitake, 2012 v
163 P. taylori Schultze, 1922 v
P. taylori ssp. metalescens Schultze, 1924 v
164 P. tetramaculatus Rukmane, 2019 v
165 P. tikoi Rukmane, 2016 v
166 P. tilikensis Bollino & Sandel, 2015 v
167 P. tobafolius Kano, 1929 v
168 P. torresi Rukmane, 2018 v
169 P. tristis Heller, 1912 v
170 P. valainisi Rukmane & Barševskis, 2016 v
171 P. venustus Waterhouse, 1841 v
P. venustus ssp. insulanus Schultze, 1919 v
172 P. viridans Heller, 1912 v v v
173 P. viridis (Chevrolat, 1879) v
174 P. yoshitakei Bollino & Rukmane, 2020 v
175 P. yoshitakeorum Yoshitake, Bollino & Sandel, 2019 v v
176 P. yukae Yoshitake, 2019 v
177 P. yuukae Yoshitake, 2019 v
178 P. zamboanganus Yoshitake, 2012 v
179 P. zebra Schultze, 1917 v
Species 82 10 11 70 3 3 11 7
Subspecies 30 2 1 9 0 0 5 0
Total 112 12 12 79 3 3 16 7
Table 2.

List of Pachyrhynchus species groups, including both morphology-based groups and molecular clades. Pachyrhynchus absurdus was synonymized with P. speciosus by Rukmane-Bārbale (2020); however, the term “absurdus species group” is retained here in accordance with the original classification proposed in Rukmane 2017.

Species Morphology-based group Molecular clade
Heller 1912 Schultze 1923, 1924 Various studies (2017–2022) Van Dam et al. 2023 Chen et al. 2025
1 P. amabilis Schultze, 1922 Group XIV amabilis species group amabilis clade
2 P. anellifer Heller, 1912 Group VII Group VII erichsoni clade
3 P. anichtchenkoi Rukmane & Barševskis, 2016 decussatus clade
4 P. annulatus Chevrolat, 1881 Group VII Group VII erichsoni clade erichsoni clade
5 P. antonkozlovi Rukmane & Barševskis, 2016 erichsoni clade
6 P. apicatus Schultze, 1922 Group V apicatus clade gemmatus clade
7 P. apocyrtoides Schultze, 1922 Group IV
8 P. apoensis Yoshitake, 2012 schoenherri species group gemmatus clade
9 P. ardentius Schultze, 1919 Group XII schoenherri species group
10 P. argus Pascoe, 1873 Group VII Group XV argus clade congestus clade
11 P. atrocyaneus Schultze, 1922 Group XV atrocyaneus species group
12 P. atronitens Yoshitake, 2019 pinorum clade
13 P. baluganus Schultze, 1924 Group VII erichsoni clade
14 P. banglas Bollino, Sandel & Rukmane, 2017 amabilis species group
15 P. barsevskisi Rukmane, 2016 pinorum species group lacunosus clade pinorum clade
16 P. basilanus Heller, 1923 Group XV atrocyaneus species group
17 P. benguetanus Schultze, 1924 Group VIII
18 P. bollinoi Rukmane-Bārbale, 2020 pinorum clade
19 P. bucasanus Schultze, 1922 Group IV
20 P. cabrasae Rukmane & Barševskis, 2016 speciosus species group
21 P. caeruleovittatus Yoshitake, 2012 amabilis clade amabilis clade
22 P. caeruleus Yoshitake, 2019 congestus clade
23 P. cagayanus Heller, 1929 decussatus clade
24 P. callainimaculatus Yoshitake, 2017 pinorum species group
25 P. cebrem Patano & Rukmane-Bārbale, 2022
26 P. chamissoi Schultze, 1922 Group XIV amabilis species group
27 P. chlorites Chevrolat, 1881 Group IV Group VIII congestus clade
28 P. chrysocyaneus Rukmane-Bārbale, 2024
29 P. chrysomaculatus Bollino, 2022 atrocyaneus species group
30 P. cinereomaculatus Rukmane-Bārbale, 2020 pinorum clade
31 P. cingulatus Pascoe, 1873
32 P. circulatus Heller, 1912 Group V Group I orbifer clade
33 P. circulimaculatus Yoshitake, 2019 speciosus species group
34 P. conformis Yoshitake, 2017
35 P. confusus Schultze, 1923 Group V
36 P. congestus Pascoe, 1873 Group IV Group VIII congestus clade congestus clade
37 P. consobrinus Schultze, 1922 Group XIII pinorum species group lacunosus clade pinorum clade
38 P. corpulentus Schultze, 1922 Group XII schoenherri species group phaleratus clade gemmatus clade
39 P. croesus Oberthur, 1879 Group III Group XV gemmatus clade
40 P. cruciatus Schultze, 1923 Group I orbifer clade orbifer clade
41 P. cumingii Waterhouse, 1841 Group V Group XI decussatus clade
42 P. davaoensis Schultze, 1934 speciosus species group orbifer clade
43 P. decussatus Waterhouse, 1841 Group V Group I decussatus clade
44 P. disargus Rukmane, 2019 argus clade
45 P. disgestus Heller, 1929
46 P. dohrni Behrens, 1887 Group II Group VIII congestus clade congestus clade
47 P. domino Rukmane, 2016
48 P. dubiosus Schultze, 1922 Group XIII pinorum species group lacunosus clade
49 P. elegans Waterhouse, 1842 Group VI Group VI schoenherri species group phaleratus clade gemmatus clade
50 P. eques Heller, 1912 Group I Group IX congestus clade
51 P. equester Heller, 1929
52 P. erichsoni Waterhouse, 1841 Group III Group IV erichsoni clade erichsoni clade
53 P. erosus Schultze, 1920 Group VII erichsoni clade erichsoni clade
54 P. esperanza Bollino, Sandel & Rukmane, 2017 schoenherri species group
55 P. faisali Bollino, 2023
56 P. felipeae Rukmane & Cabras, 2018 erichsoni clade
57 P. florulentus Yoshitake, 2019 pinorum clade
58 P. forsteni Vollenhoven, 1864 Group I Group III
59 P. franciscoi Rukmane & Cabras, 2018 erichsoni clade
60 P. galeraensis Schultze, 1934 erichsoni clade
61 P. gemmatus Waterhouse, 1841 Group IV Group VIII congestus clade gemmatus clade
62 P. gilvomaculatus Yoshitake, 2017 amabilis clade
63 P. gloriosus Faust, 1895 Group II Group XI gloriosus clade decussatus clade
64 P. halconensis Schultze, 1922 Group I gemmatus clade
65 P. helenperrinae Rukmane, 2018
66 P. helleri Kuntzen, 1914 Group V gemmatus clade
67 P. hirokii Yoshitake, 2012 atrocyaneus species group amabilis clade
68 P. igorota Schultze, 1917 Group X decussatus clade
69 P. ilgas Rukmane, 2017 absurdus species group
70 P. imitans Rukmane & Bollino, 2020 amabilis species group amabilis clade
71 P. inclytus Pascoe, 1873 Group II Group X inclytus clade decussatus clade
72 P. infernalis Fairmaire, 1897 Group I Group I orbifer clade
73 P. jitanasaius Chen & Lin, 2017 orbifer clade
74 P. jugifer Waterhouse, 1841 Group V Group I orbifer clade
75 P. kirklayroni Rukmane, 2019 decussatus clade
76 P. kraslavae Rukmane & Barševskis, 2016 speciosus species group miltoni clade orbifer clade
77 P. lacunosus Heller, 1912 Group III Group XIII pinorum species group lacunosus clade pinorum clade
78 P. latifasciatus Waterhouse, 1842 Group V Group II erichsoni clade erichsoni clade
79 P. layroni Rukmane & Cabras, 2018 orbifer clade
80 P. libucanus Schultze, 1923 Group I
81 P. loheri Schultze, 1917 Group X pinorum species group pinorum clade
82 P. lorquini Chevrolat, 1881 Group IV Group VIII congestus clade
83 P. lubanganus Bollino & Sandel, 2015
84 P. marinduquensis Rukmane & Barševskis, 2016 orbifer clade
85 P. maruyamai Yoshitake, 2019
86 P. masatoshii Yoshitake & Yap, 2017
87 P. masatoshiensis Rukmane-Bārbale, 2024
88 P. miltoni Cabras & Rukmane, 2016 speciosus species group miltoni clade
89 P. mindoroensis Rukmane & Háva, 2020 congestus clade congestus clade
90 P. mohagani Bollino & Sandel, 2015
91 P. möllendorffi Heller, 1899 Group II Group X gloriosus clade decussatus clade
92 P. moniliferus Germar, 1824 Group V Group I orbifer clade orbifer clade
93 P. morio Heller, 1912 Group IV Group VIII
94 P. morotaiensis Vollenhoven, 1864 Group I Group III morotaiensis clade
95 P. multipunctatus Waterhouse, 1841 Group VI Group VI
96 P. naokii Yoshitake, 2012 atrocyaneus species group
97 P. naokoae Yoshitake, 2019
98 P. negrosensis Schultze, 1924
99 P. neoabsurdus Rukmane, 2017 absurdus species group
100 P. niisatoi Yoshitake, 2017 pinorum species group lacunosus clade pinorum clade
101 P. nitcisi Rukmane & Barševskis, 2016 schoenherri species group
102 P. nobilis Heller, 1912 Group II Group XI erichsoni clade
103 P. noeli Yoshitake, 2019
104 P. notocruciatus Yoshitake, 2017 speciosus species group orbifer clade
105 P. obumanuvu Cabras, Donato, Medina, & Van Dam, 2021
106 P. occidentalis Rukmane, 2017 absurdus species group miltoni clade orbifer clade
107 P. ochroplagiatus Heller, 1912 Group I Group XV argus clade congestus clade
108 P. octoannulatus Yoshitake, Bollino & Sandel, 2019 speciosus species group orbifer clade
109 P. ohbayashii Yoshitake, 2017
110 P. orbifer Waterhouse, 1841 Group V Group I orbifer clade orbifer clade
111 P. orientalis Rukmane, 2017 absurdus species group
112 P. ottomerkli Rukmane, 2019
113 P. panumanon Cabras & Medina, 2022
114 P. perpulcher Waterhouse, 1841 Group III Group XV gloriosus clade decussatus clade
115 P. phaleratus Waterhouse, 1841 Group V Group I phaleratus clade gemmatus clade
116 P. pinorum Pascoe, 1873 Group III Group XIII pinorum species group lacunosus clade pinorum clade
117 P. postpubescens Schultze, 1922 Group II speciosus species group
118 P. pseudamabilis Yoshitake, 2012 amabilis species group amabilis clade amabilis clade
119 P. pseudhalconensis Rukmane, 2016 gloriosus clade decussatus clade
120 P. pseudoproteus Schultze, 1922 Group VI
121 P. psittacinus Heller, 1912 Group III Group X pinorum clade
122 P. psittaculus Heller, 1921 Group X pinorum clade
123 P. pulchellus Behrens, 1887 Group II Group X gloriosus clade decussatus clade
124 P. rebus Rukmane, 2016
125 P. regius Schultze, 1922 Group II speciosus species group orbifer clade
126 P. reicherti Schultze, 1929 orbifer clade
127 P. reticulatus Waterhouse, 1841 Group V Group I miltoni clade orbifer clade
128 P. rizali Schultze, 1934 erichsoni clade
129 P. rochaorum Yoshitake, 2020 decussatus clade
130 P. roseomaculatus Waterhouse, 1841 Group IV Group I
131 P. rufopunctatus Waterhouse, 1842 Group V
132 P. rugicollis Waterhouse, 1841 Group V Group I orbifer clade orbifer clade
133 P. rukmaneae Barševskis, 2016 congestus clade congestus clade
134 P. sagittatus Rukmane, 2019
135 P. sakaii Yoshitake, 2017
136 P. samarensis Schultze, 1923 Group II speciosus species group
137 P. sanchezi Heller, 1912 Group IV Group VIII phaleratus clade gemmatus clade
138 P. sarcitis Behrens, 1887 Group IV Group VIII gemmatus clade
139 P. schoenherri Waterhouse, 1841 Group III Group IV schoenherri species group gemmatus clade
140 P. schuetzei Schultze, 1917 Group VII
141 P. semiignitus Schultze, 1922 Group IV
142 P. semperi Heller, 1912 Group II Group X pinorum species group pinorum clade
143 P. septentrionalis Yoshitake, 2017 pinorum species group lacunosus clade pinorum clade
144 P. sergejevae Rukmane, 2018 atrocyaneus species group
145 P. shavrini Rukmane & Barševskis, 2016 inclytus clade decussatus clade
146 P. signaticollis Schultze, 1922 Group IV
147 P. signatus Schultze, 1919 Group IV
148 P. similis Bollino, 2022 atrocyaneus species group amabilis clade
149 P. smaragdinus Behrens, 1887 Group III Group V apicatus clade
150 P. sonani Kôno, 1930 orbifer clade
151 P. speciosus Waterhouse, 1841 Group V Group II speciosus species group miltoni clade orbifer clade
152 P. sphaericollaris Schultze, 1923 Group I
153 P. sphenomorphoides Yoshitake, 2012 atrocyaneus species group amabilis clade
154 P. stellio Heller, 1912 Group V Group I
155 P. subamabilis Yoshitake, 2012 amabilis species group amabilis clade
156 P. subanon Bollino, 2022 atrocyaneus species group
157 P. subpalidius Rukmane-Bārbale, 2022
158 P. sugbo Bollino, 2025 moniliferus species group
159 P. sulphureomaculatus Schultze, 1922 Group V apicatus clade
160 P. sumptuosoides Yoshitake, 2017 pinorum species group
161 P. sumptuosus Schultze, 1917 Group IX congestus clade congestus clade
162 P. tadauchii Yoshitake, 2012 speciosus species group
163 P. taylori Schultze, 1922 Group VIII gemmatus clade
164 P. tetramaculatus Rukmane, 2019 pinorum clade
165 P. tikoi Rukmane, 2016 amabilis species group
166 P. tilikensis Bollino & Sandel, 2015 gemmatus clade
167 P. tobafolius Kano, 1929 decussatus clade
168 P. torresi Rukmane, 2018 atrocyaneus species group
169 P. tristis Heller, 1912 Group III Group XIII pinorum species group
170 P. valainisi Rukmane & Barševskis, 2016
171 P. venustus Waterhouse, 1841 Group III Group V gemmatus clade
172 P. viridans Heller, 1912 Group IV Group VIII congestus clade
173 P. viridis (Chevrolat, 1879)
174 P. yoshitakei Bollino & Rukmane, 2020 amabilis species group
175 P. yoshitakeorum Yoshitake, Bollino & Sandel, 2019 speciosus species group
176 P. yukae Yoshitake, 2019
177 P. yuukae Yoshitake, 2019 congestus clade
178 P. zamboanganus Yoshitake, 2012 amabilis species group amabilis clade
179 P. zebra Schultze, 1917 Group I

The islands north of mainland Luzon, although consisting of only a small group of islands, harbor 11 species (6.2%), of which 4 are endemic. The island of Mindoro has ten species, representing 5.6% of the described species, with six being endemic. The Greater Negros-Panay Faunal Region has 11 species (8 of which are endemic), representing 6.1% of the described species (Fig. 2, Table 1). Additionally, oceanic islands such as Lubang and Sibuyan contain three species each, P. lubanganus, P. mohagani, and P. tilikinesis on Lubang; P. noeli, P. naokoae, and P. viridans on Sibuyan. Among them, only P. viridans is also distributed in the Greater Negros-Panay PAIC and the islands north of mainland Luzon. Pachyrhynchus species found in the Indonesia archipelago do not occur in the Philippines. Furthermore, 17 Pachyrhynchus species and subspecies are known to occur in more than one faunal region (Table 1).

Despite these records, many areas in the Philippines remain underexplored for Pachyrhynchus, such as the Bicol Peninsula in southern Luzon and the Greater Sulu Faunal Region, which encompasses the archipelagos of Sulu and Tawi-Tawi. Among the 179 Pachyrhynchus species, 165 species are endemic to specific regions or islands, indicating the high level of endemism (92.2%) in these weevils. The high endemism also underscores the importance of biodiversity conservation in these areas. In Taiwan, for example, three endemic species and three endemic subspecies are distributed on two small islands. Due to their endemism and high collection pressure, all the Pachyrhynchus species were listed as rare and valuable species protected under the Wildlife Conservation Act in 2009. In 2019, Pachyrhynchus species in the Philippines were listed as vulnerable in the national list of threatened Philippine fauna and their categories. Deforestation for agricultural purposes poses a threat to Pachyrhynchus weevils in the Philippines, potentially causing habitat loss, especially for locally endemic species (Cabras and Rukmane 2016). Continuously updating records of Pachyrhynchus will not only enhance our understanding of its patterns of diversity but also aid in identifying priority conservation sites for this genus, especially for threatened species.

Acknowledgements

We thank Dr. Mei-Ling Chan (National Museum of Natural Science, Taichung, Taiwan) and Dr. Dávid Rédei (National Chung Hsing University, Taichung, Taiwan) for providing valuable suggestions, and Olaf Jäger for providing type information in Senckenberg Naturhistorische Sammlungen, Museum für Tierkunde, Dresden (SMTD). We also thank the Senckenberg Deutsches Entomologisches Institut (SDEI) and the Natural Hitory Museum, London (NHMUK) for loaning specimens. We deeply mourn the loss of Dr. Analyn Anzano Cabras, whose contributions to the study of Pachyrhynchus weevils and Philippine biodiversity were significant. Dr. Cabras was not only a generous and kind-hearted researcher but also a pivotal force in advancing our understanding of these remarkable insects.

Additional information

Conflict of interest

The authors have declared that no competing interests exist.

Ethical statement

No ethical statement was reported.

Funding

This work was supported by National Science and Technology Council, Taiwan (NSTC 112-2621-B-002-003-MY3) to Hui-Yun Tseng.

Author contributions

Conceptualization: YH, HYT. Funding acquisition: HYT. Investigation: YH, SMC, BHH. Supervision: HYT. Writing – original draft: YH, SMC, BHH. Review and editing: JFT, AKSA, MLJ, HYT.

Author ORCIDs

Shan-Min Chen https://orcid.org/0009-0009-7346-0909

Ace Kevin S. Amarga https://orcid.org/0000-0002-9776-6576

Jing-Fu Tsai https://orcid.org/0000-0003-3076-3976

Bin-Hong Ho https://orcid.org/0000-0001-6008-9076

Hui-Yun Tseng https://orcid.org/0000-0002-1166-4756

Data availability

All of the data that support the findings of this study are available in the main text.

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Yun Ho and Shan-Min Chen contributed equally.
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