Bulletin of the Buffalo Society of Natural Sciences - Buffalo Museum ...

Bulletin of the Buffalo Society of Natural Sciences, Volume 40 59 

NOTES ON THE BIOGEOGRAPHY AND PHYLOGENY OF 

EASTERN ASIAN GHOST MOTHS (LEPIDOPTERA: HEPIALIDAE) 

John R. Grehan 

Buffalo Museum of Science, 1020 Humboldt Parkway, Buffalo, NY 14211 

jgrehan@sciencebuff.org 

ABSTRACT – The distribution records of eastern Asian Hepialidae are mapped for the genera 

Bipectilus Chu & Wang, 1985, Endoclita Felder, 1874, Hepialiscus Hampson, [1893], Napialus Chu 

& Wang, 1985, Palpifer Hampson, [1893], Parahepialiscus Viette, 1950, Thitarodes Viette, 1968 and 

Xhoaphyrix Viette, 1953. All are distributed across mainland eastern Asia between the Himalayas and 

China, with Endoclita and Thitarodes also extending north to Japan and the Russian Far East. 

Endoclita and Palpifer occur south of the Himalayas, and only Thitarodes is absent from South East 

Asia. The similarity of distribution ranges between the genera, and the presence of vicariant lineages 

in Bipectilus and the Hepialiscus group (with respect to Napialus) is predicted to be the result of their 

having evolved from widespread ancestral distributions that largely encompassed the generic ranges. 

These ancestral distributions overlapped Eurasian and North American genera in the Russian Far East 

and Japan. The distribution of Hepialiscus, Parahepialiscus, and Xhoaphryx may together represent 

the Asian fragment of an ancestral range that also included Australia and New Guinea where it is now 

represented by Oxycanus Walker, 1856. Endoclita also has a very close vicariant relationship with the 

Australasian Aenetus Herrich-Schäffer, 1855 that may have originated from a formerly widespread 

ancestor across eastern Asia and Australasia. The geological history behind the convergence of 

tectonic plates at their common biogeographic boundary may have played a role in the differentiation 

of the two genera. 

KEYWORDS – Hepialidae, ghost moth, biogeography, phylogeny, Asia. 

INTRODUCTION 

The global biogeography and phylogeny of 

ghost moths (Hepialidae) is poorly understood, 

particularly for inter-generic and intercontinental 

relationships (Nielsen et al., 2000; Grehan and 

Rawlins, 2003). The family has a global range, 

although absent from some extensive geographic 

areas such as Madagascar, the Caribbean 

islands, west-central tropical Africa, and the 

lowlands of eastern and central United States, 

even though apparently suitable habitats are 

present. In regions where Hepialidae occur there 

are seven geographic clusters of genera and 

species (based on the classification of Nielsen et 

al., 2000) where geographic overlap is minimal 

or absent. The generic and species diversity of 

these geographic clusters are as follows: 

1) America north of Mexico: 4-16. 

2) Western Eurasia/North Africa: 7-78. 

3) America south of the United States: 14-132. 

4) Southern Africa: 6-78. 

5) Australasia: 9-190. 

6) Fiji/Samoa: 1-1. 

7) Eastern Asia: 10-47. 

Generic diversity is not substantially 

different between the clusters other than for 

Fiji/Samoa with its monotypic genus, and 

America south of the United States which has 

the highest number of genera and almost as 

many documented species as Australasia with 

the highest species diversity. The only generic 

overlap between these clusters involves 

Korscheltellus Börner, 1920, Gazoryctra 

Hübner, [1820], and Phymatopus Wallengren, 

1869 that range across North America, Western 

Eurasia/North Africa, and Eastern Asia. 

In this paper the distribution ranges of the 

endemic eastern Asian genera are examined to 

characterize their geographic limits and how 

these limits do or do not intersect with genera in 

western Asia and Australasia. In the absence of 

a comprehensive revision and synthesis of the 

systematics and taxonomy for most of these 

groups, the level of biogeographic detail is 

limited principally to each genus as a whole. 

The eastern Asian genera (or generic groups) 

considered here are (1) Bipectilus Chu & Wang, 

1985, (2) Hepialiscus Hampson, [1893] (along 

with the closely related Napialus Chu & Wang,


1985, Xhoaphyrix, and Parahepialiscus), (3) 

Thitarodes Viette, 1968, (4) Palpifer Hampson, 

[1893], and (5) Endoclita Felder, 1874. 

The geographic range defined by the total 

range of all of these genera includes South East 

Asia, Ceylon/India and the Russian Far 

East/Japan. Specimens that were dissected and 

illustrated here are from the following 

collections: JRG (John R. Grehan collection); 

CMNH (Carnegie Museum of Natural History); 

NZAC (New Zealand Arthropod Collection). 

1) Bipectilus Chu & Wang, 1985 

Characterized as one of the most distinctive and 

peculiar hepialid genera, Bipectilus differs from 

all other Eurasian and Oriental genera in having 

bipectinate antennae and highly specialized male 

genitalia. The genus may represent a basal 

lineage within the Hepialidae (Nielsen, 1988). 

Bipectilus ranges between eastern China (south 

of 33° N), eastern Nepal, and southern Burma. 

All eight species are vicariant, and there are two 

vicariant lineages, an eastern group comprising 

two species, and the remaining four species 

between Nepal, central and southern China, and 

Thailand (Fig. 1). 

Fig. 1. Distribution records of Bipectilus. Each symbol represents different species. Vicariant subgroups 

represented by blue and red symbols (Nielsen, 1988). 

2) Hepialiscus Hampson, [1893] group 

Ueda (1988) suggested that Hepialiscus, and the 

genera Napialus (three species), Parahepialiscus 

(monotypic, Fig. 2a), and Xhoaphyrix 

(monotypic, Fig. 2b) may be monophyletic or at 

least represent closely related species. This 

tentative classification was based on their 

sharing a pattern of wing venation where R4 

(=Rs3) and R5 (=Rs4) arise separately from 

R2+3 (=Rs1+Rs2) (see Kristensen, 1999 (Fig. 

3). 

The separate origin of R4 and R5 also 

applies to the New Guinea/Australian Elhamma 

Walker, 1856, Oxycanus Walker, 1856, Jeana 

Tindale, 1935, the New Zealand genera 

Cladoxycanus Dumbleton, 1966, Dioxycanus 

Dumbleton, 1966, Dumbletonius Dugdale, 1988, 

Heloxycanus Dugdale, 1994, and Wiseana 

Viette, 1961, and the North African 

Neohepialiscus Viette, 1948 (Viette, 1948). 

This ‘oxycanine’ pattern (as 

characterized by Dumbleton, 1966) appears to 

be a problematic indicator of relationship 

because it also occurs in the South American 

Roseala Viette, 1950, Cibyra (Aepytus) Herrich- 

Schäffer, [1858], C. (Vietteogorgopis), and C.


(Lamelliformia) (Nielsen and Robinson, 1983; 

Ueda, 1988; Brown et al., 2000). These South 

American genera have abdominal features in the 

tergal lobe (Grehan, 2010) and genitalic 

similarities that appear to be closer to other 

‘cibyrine’ genera rather than Asian and 

Australasian ‘oxycanine’ genera (JRG, personal 

observation), although their future taxonomic 

placement will also be contingent upon a 

comprehensive phylogenetic analysis. 

Fig. 2. Hepialiscus group. (a) Parahepialiscus borneensis (BMNH(E) #953897); (b) Xhoaphryx sp. (BMNH(E) 

#953898). Scale = 5 mm. Images courtesy of Carlos Mielke. Copyright: Trustees of the Natural History Museum, 

London, used with permission 

Fig. 3. Forewing venation. (a) ‘oxcanine’, Dioxycanus oreas (Hudson, 1920); (b) ‘hepialine’, Aoraia insularis 

Dugdale, 1994. Reproduced from Dugdale (1994 figs 75, 77) by permission. 

None of the characters listed for 

Hepialiscus by Ueda (1988) were characterized 

as unique for the genus. Presence of a 

terberculate plate (pinnaculm) anterior to the 

spiracle on the 2 nd to 6 th abdominal segments 

may distinguish Hepialiscus from Bipectilus, but 

not Endoclita (personal observation). 

Napialus was created by Chu and 

Wang (1985) for a new species, and a further 

two species were added by Wu (1992) and 

Nielsen et al. (2000) respectively. Three of the 

four features found in Napialus are also shared 

by Hepialiscus (Ueda, 1988). Napialus may 

represent largely northeastern vicariant of 

Hepialiscus ranging mostly further south in 

China. The monotypic Xhoaphryx of Vietnam 

differs from Hepialiscus in having an oval 

pseudotegumen and it also lacks an epiphysis, 

but Ueda (1988) thought the genera were closely 

related because of the shared oxycanine venation 

and the presence of a subanal sclerite in the male 

genitalia. 

Ueda (1988) concurred with Viette’s 

(1950a) view that Parahepialiscus was also 

closely related to Hepialiscus because of the 

‘oxycanine’ venation and similarity of genitalia. 

Viette (1950a) distinguished Parahepialiscus by 

the presence of lateral sclerification around the 

adeagus, but Ueda (1988) regarded this structure 

to be homologous with the subanal sclerite 

found in Hepialiscus and Xhoaphyrx. 

Viette (1948) also suggested a close 

relationship between Hepialiscus and the North 

African Neohepialiscus algeriensis (de Joannis, 

1903) on the basis of their both sharing 

‘oxycanine’ venation, but he did not include a 

comparison with male Hepialiscus genitalia or


refer to the presence or absence of a subanal 

sclerite. The pseudoteguminal arms of 

Neohepialiscus are long and narrow, and in this 

respect are similar to those of Napialus 

hunanensis (Chu & Wang, 1985: fig. 36). 

Hepialiscus ranges between Nepal, Taiwan, and 

northern Borneo (Fig. 4), but is mostly 

represented by a few scattered records. Within 

this range Xhoaphryx is represented by a single 

published record in northern Vietnam while 

Parahepialiscus is confined to northern Borneo. 

Fig. 4. Distribution of Hepialiscus and closely related genera. Hepialiscus (red circles), Napialus (blue circles), 

Parahepialiscus borneensis (Pfitzner, 1933) (yellow circle), Xhoaphryx lemeei Viette, 1953 (purple circle) 

(Daniel, 1940; Viette, 1953; Chu & Wang, 1985; Ueda, 1988; Wu, 1992; Robinson et al., 1995). 

5) Thitarodes Viette, 1968 

Thitarodes (Fig. 5) was originally proposed with 

four species, but a further 53 were later added 

(Ueda, 1996, 2000; Nielsen et al., 2000; see also 

Zhu et al., 2004). Viette (1968) drew attention to 

the presence of a basal spine on the male valve, 

but at least six species added by Nielsen et al. 

(2000) lack the spine (as illustrated by Chu & 

Wang, 1985). A basal spine occurs in some 

other genera such as Schausiana Viette, 1950 of 

Mexico-Central America (JRG, personal 

observation). Thitarodes is distinguished from 

the Hepialiscus group by vein R4 branching 

from R4+R5, and from other Asian genera by 

the male genitalia. Larvae are subterranean root 

feeders in alpine meadows or pastures (Yang et 

al., 1996; Maczey et al., 2010). 

Fig. 5. (a) Thitarodes namnai Maczey, 2010 (Bhutan), (b) T. caligophilus Maczey, 2010 (Bhutan) (Maczey et al., 

2010). Images courtesy of Norbert Maczey. Scale = 5 mm.


Fig. 6. Distribution records for Thitarodes (Bang-Haas, 1939; Chu & Wang, 1985; Liang et al., 1988; Wang, 

1990; Yang, 1993, 1994; Yang & Jiang, 1995; Yang & Yang, 1995; Yang et al., 1992, 1993; Robinson et al., 

1995; Ueda, 1996, 2000; Tu et al., 2009; Maczey et al., 2010; Zou et al., 2011). 

Thitarodes has a northern limit about 

50º N in central Asia and the Far East, and a 

westernmost limit about 85ºW in Nepal and 

Southern China (Fig. 6). There is a considerable 

species concentration in southeast China and the 

Himalayas, including 16 species being recorded 

from the province of Yunnan alone. With many 

species only superficially described and lacking 

comparative diagnoses with respect to the rest of 

the genus, it is possible that some of this 

diversity may be reduced by future synonymy. 

There is a notable lack of confirmed records 

from eastern China, resulting in a current 

geographic disjunction between the center of 

diversity for this group and species occurring in 

Taiwan, Japan, and the Russian Far East. 

4) Palpifer Hampson, [1893] 

The few published records for Palpifer (Fig. 7) 

are geographically scattered between Japan, 

northwestern India and Sri Lanka, and Java (Fig. 

8). Palpifer comprises nine species (Nielsen et 

al., 2000) and is in need of taxonomic revision 

(Robinson et al., 1995). Monophyly for some or 

all species may be indicated by the uniquely 

shared presence of a small dark spot along the 

posterior forewing margin basal to vein CuA2, a 

white spot at the base of the forewing discal cell 

on or near vein M3, and an orange-brown fringe 

on some or much of the outer and posterior 

hindwing margins. Larvae are root borers 

(Kalshoven, 1965; Robinson et al., 1995). 

Fig. 7. Palpifer murinus, West Pahang, Malasia. 

(BMNH(E) #953899). Scale = 5 mm. Image courtesy 

Carlos Mielke. Copyright: Trustees of the Natural 

History Museum, London, used with permission 

5) Endoclita Felder, 1874 

This large genus of 60 species (Nielsen et al., 

2000) ranges between the Russian Far East, 

southern India, Sri Lanka, and Southeast Asia 

(Fig. 9). Some species are widespread in India 

and China while most are known from one 

locality. The map (Fig. 9) includes an 

unconfirmed record by Smetacek (1997) from 

northwestern India. Monophyly of the genus has 

yet to be corroborated. Larvae are wood borers 

of trees and shrubs (Grehan, 1989).


Fig. 8. Distribution records for Palpifer (Butler, 1879; Moore, 1887, 1879; Cotes & Swinhoe, 1887; Piepers & 

Snellen, 1900; Swinhoe, 1905; Pfitzner, 1914; Matsumura, 1931; Daniel, 1940; Viette, 1968; Robinson et al., 

1995; Utsumi, personal communication). 

Fig. 9. Distribution of Endoclita (Van Eecke, 1915; Daniel, 1940; Tindale, 1941, 1942, 1958; Viette, 1950b; 

Robinson et al., 1995; Tshistjakov, 1996; Jeon et al., 2000; Utsumi & Ohgushi, 2007; Utsumi, personal 

communication).


BIOGEOGRAPHY 

Generic categories are a proxy for one or more 

characters that group two or more species 

together as being more closely related to each 

other than to other species. Genera, like any 

other taxonomic category, often represent one or 

more features that are relatively unambiguous 

for grouping species together, whereas features 

that identify relationships between those groups 

may be more difficult to recognize – if they are 

evident at all. This is the current situation for the 

Hepialidae where many genera have long been 

recognized or well defined while inter-generic 

relationships remain uncertain. 

As clusters of related species, there is 

no necessary relationship between the generic 

distributions and any particular geographic area 

such as ‘Eastern Asia’ or any of the artificial 

biogeographic classifications that create 

geographic regions such as ‘Indo-Malaysia’ and 

‘Australasia’ (e.g. Olsen et al., 2001). 

Instead, it is the geographic and 

phylogenetic relationships of taxa that are 

biogeographically informative. Within the 

current constraints of limited phylogenetic 

resolution for inter-generic relationships, the 

uncertainty of species identification within some 

genera and the paucity of collecting records in 

many areas, biogeographic interpretation of 

eastern Asian Hepialidae is limited here to 

geographic and phylogenetic examples 

involving distributions that are geographically 

vicariant or non-overlapping (and often referred 

to as allopatric). 

In most biogeographic approaches, 

vicariant distributions are interpreted as the 

result of an imagined dispersal from an 

imagined center of origin, where the ancestral 

form it theorized to have occupied a geographic 

area smaller than the combined geographic area 

of its vicariant descendants. This approach goes 

back to Darwin’s theory of evolution and has 

remained a predominant theory of evolution to 

this day. 

While very popular, the center of origin 

theory relies more on intuition than evidence for 

proposed dispersal events to explain the 

vicariant differentiation of descendant taxa 

(species or any other taxonomic level). It also 

generates internal contradictions where a barrier 

is supposed to explain the isolation and 

differentiation of vicariant taxa while at the 

same time being just permeable enough to allow 

either a one-off or a few chance dispersals over 

the barrier to reach the other side where there is 

sufficient isolation to allow divergence (Craw et 

al., 1999). 

The center of origin/dispersal theory is 

most often represented by the location of the 

most primitive lineage being at or nearer the 

center of origin, with more derived groups 

representing successively recent colonization 

(Heads, 2009a, b). But this phylogenetic pattern 

could be just as well explained as the sequence 

of differentiation of a widespread ancestor (and 

without generating contradictions between 

theorized dispersal and actual distribution). In 

the following sections the presence of vicariant 

patterns is proposed as evidence of a formerly 

widespread ancestor with a geographic range 

that spanned all or much of the combined 

distributions of the vicariant taxa involved. 

1. Geographic relationships 

All of the eastern Asian genera occur in the 

Himalayas and southwestern China/northern 

South East Asia, and only Bipectilus is restricted 

to the continental mainland. The other four 

genera geographically vary with respect to their 

presence on adjacent island archipelagos and the 

Indian subcontinent. Thitarodes is particularly 

diverse in species in western China/Himalayas 

and also occurs in Taiwan and at least southern 

Japan (each with an endemic species). The 

Hepialiscus group is represented by very 

scattered mainland records as well as occurring 

on Taiwan and northern Borneo. The mainland 

Asian distribution of Palpifer is limited to 

scattered localities in the Himalayas and south to 

Borneo and the Malaysian Peninsula with 

disjunct records from eastern China, Taiwan, 

Japan, Java and Sri Lanka. Endoclita is the most 

widespread genus, being present across the 

Indian subcontinent and Sri Lanka as well as all 

major island archipelagos south east to the 

Moluccas. 

The northern distributions of Endoclita 

and Thitarodes overlap with three genera that


also occur beyond eastern Asia. These occur in 

the region of China, the Russian Far East, and 

Japan. Pharmacis Hübner, [1820], with eight 

species in Europe, including P. fusconebulosa 

(De Geer, 1778) that ranges east to the Amur 

River region of the Russian Far East and Japan 

(Fig. 10). A similar overlap occurs with 

Gazoryctra and Phymatopus (Figs 11-12), both 

of which occur in western Asia and also North 

American north of Mexico. 

Fig. 10. Distribution range for Pharmacis. Dotted line indicates uncertainty about the precise geographic limits 

(De Freina & Witt, 1990; Staudinger, 1887; Utsumi, personal communication). 

The Eurasian distribution of 

Gazoryctra (Fig. 11) involves one species 

confined to Europe, two restricted to the Russian 

Far East and Japan, and G. fuscoargenteus 

(Bang-Haas, 1927) ranging across northern 

Eurasia. The distributions of the nine species of 

North America north of Mexico are poorly 

documented. About five of these appear to be 

confined to the west, and two range across 

Canada and parts of northern United States. 

There appears to be only one eastern endemic 

which is found in the southern Appalachians. 

In contrast to Gazoryctra, the Eurasian 

range of Phymatopus is represented by only a 

single species, and in North America the genus 

is represented by only three species that are also 

geographically restricted to the coastal region of 

the western United States (Fig. 12). 

Geographic overlap of the Eurasian and 

North American genera with the northern range 

of Endoclita and Thitarodes suggests that the 

Asian northeast is an important biogeographic 

center for understanding the origin and evolution 

of Eurasian Hepialidae. Biogeographically, 

northeastern Asia represents a center of 

differentiation between northern Eurasian/North 

American genera, and those of central/southern 

Asia that are otherwise widespread across 

eastern Asia. This pattern may represent 

ancestral distributions that occupied different 

geographic sectors other than their current 

overlapping boundary, or the marginal 

geographic overlap is the result of subsequent 

range expansion between the two vicariant 

groups. 

The overlapping boundary between the 

two Asian distributional patterns contrasts with 

the apparent lack of geographic overlap between 

the eastern Asian genera and the diverse 

Australasian hepialid fauna. This apparent 

absence of overlap may, however, be an artifact 

of current taxonomy and further consideration of 

phylogenetic relationships may show a closer 

relationship for at least some groups as 

discussed in the next section on phylogeny. 

2. Phylogenetic relationships 

The phylogenetic relationships of eastern Asian 

genera have yet to be explored in detail. Nielsen 

(1988) suggested Bipectilus represented a basal 

lineage within the Hepialidae, but was unable to 

draw any definitive conclusions. Among some 

of the key questions for the future will be the 

relationships of eastern Asian genera with each 

other, and with Hepialidae in other parts of 

Eurasia and northern America, southern Africa, 

and Latin America. At this time there is some


phylogenetic evidence in support of a close 

relationship between the Hepialiscus group and 

the Australasian Oxycanus, and between 

Endoclita and the Australasian Aenetus. 

Fig. 11. Distribution localities for Gazoryctra (Edwards, 1886; Neumoegen & Dyar, 1894; Wagner & Tindale, 

1988; De Freina & Witt, 1990; Handfield, 1999; Tshistjakov, 1997; Grehan & Rawlins, 2003; Utsumi, personal 

communication). 

Fig. 12. Distribution of Phymatopus. Solid-dotted line – uncertain eastern limits of the only western Asian 

species Phymatopus hecta (Linnaeus, 1758) (De Freina & Witt, 1990; Grehan & Rawlins, 2003; Utsumi, personal 

communication). 

(a) Hepialiscus-Oxycanus: Ueda (1988) 

illustrated subanal sclerites (or paramedial 

sclerites) for Hepialiscus (Fig. 13a-d), and 

suggested these were also present in 

Parahepialiscus borneensis (Pfitzner, 1933) 

based on Viette’s (1950) illustration of a 

“garniture latérale, sclérifeé en forme d’Y, au 

pénis” (Fig. 13e) and Xhoaphryx lemeei (Fig. 

13f) as they were ‘vaguely figured’ by Viette 

(1953). He also drew attention to the presence of 

subanal sclerites in Oxycanus goldfinchi Tindale 

1935 (Fig. 13g) and suggested this shared 

similarity represented evidence of a close 

phylogenetic relationship. 

Comparison of 44 of the 57 recognized 

hepialid genera (listed in Grehan, 2010 and 

subsequently Andeabatis chilensis [Ureta, 

1951]), resulted in subanal sclerites being found 

only in Oxycanus (Fig. 13h-i) (the Hepialiscus 

group was not available). Subanal sclerites are 

also absent from other hepialoid genera in the 

Exoporia and its sistergroup Heteroneura (cf. 

genitalic descriptions in Kristensen, 1999). 

The subanal sclerite represents a 

uniquely shared feature for Parahepialiscus, 

Xhoaphryx and some or all Hepialiscus and 

Oxycanus species. Excluded from this clade are 

the other ‘oxycanine’ genera in Australia: 

(Jeana - Fig. 14a, Elhamma - Fig. 14b) and 

New Zealand (Cladoxycanus - Fig. 14c, 

Dioxycanus - Fig. 14d, Dumbletonius - Fig. 

14e), Heloxycanus - Fig. 14f, Wiseana - Fig. 

14g) (see also Dugdale, 1994) or the North 

African Neohepialiscus algeriensis (Fig. 14h). 

Genitalic illustration of Napialus hunanensis by 

Chu & Wang (1985) lacks the detail to confirm 

subanal sclerite presence or absence (Fig. 14i). 

A phylogenetic affinity between 

Hepialiscus and Oxycanus suggests a broad 

ancestral range encompassing the now vicariant 

distributions of Parahepialiscus, Xhoaphryx,


Hepialiscus and Oxycanus between south the disjunction between Borneo and New 

eastern Asia and Australasia (Fig. 15). Whether Guinea represents an actual gap or is the result 

a b c 

d e f 

g h i 

Fig. 13. Subanal sclerite (marked in yellow): (a) Hepialiscus nepalensis (Walker, 1956) (Ueda, 1988:fig. 6); (b) 

Hepialiscus taiwanus Ueda, 1988 (Ueda, 1988: fig. 10); (c) Hepialiscus robinsoni Ueda, 1988 (Ueda, 1988: fig. 

9); (d) Hepialiscus monticolis Ueda, 1988 (Ueda, 1988: fig. 11) (e) Parahepialiscus borneensis (Pftzner, 1933) 

(Viette, 1950a: fig. 2); (f) Xhoaphyrx lemeei Viette, 1950 (Viette, 1953: fig 1); (g) Oxycanus goldfinchi Tindale, 

1935 (Ueda, 1988: fig. 7); (h) Oxycanus dirempta (Walker, 1865), Australia (M219, CMNH); (i) Oxycanus 

‘lyelli’ Tindale, 1935, Australia (M218, CMNH).


a b c 

d e f 

g h i 

Fig. 14. Absence of the subanal plate in various oxycanine genera: (a) Jeana sp., Australia (M194, JRG), (b) 

Elhamma australasiae (Walker, 1856), Australia (M181, NZAC), (c) Cladoxycanus minos (Hudson, 1905), New 

Zealand (M137, JRG), (d) Dioxycanus fusca (Philpott, 1914), New Zealand (M172, JRG), (e) Dumbletonius 

unimaculata (Salmon, 1948), New Zealand (M209, JRG), (f) Heloxycanus patricki Dugdale, 1994, New Zealand 

(M151, JRG), (g) Wiseana copularis (Meyrick, 1912), New Zealand (M216, JRG); (h) Neohepialiscus 

algeriensis (Viette, 1948: fig 5); (i) Napialus hunanensis (Chu & Wang, 1985: fig. 36).


Fig. 15. Distribution of Hepialiscus group (red circles) and Oxycanus (blue circles). 

a 

b 

Fig. 16. Vicariant relationship between Endoclita (red circles) and Aenetus (blue circles). (a) Total distribution of 

both genera; (b) Distribution records of Endoclita and Aenetus at their shared biogeographic boundary in the 

Moluccas archipelago. 

of inadequate collecting remains to be 

determined. 

b) Endoclita. The adjacent vicariant 

distributions of Endoclita and Aenetus Herrich- 

Schäffer, 1855 (Fig. 16a) (Grehan, 1988; Craw 

et al., 1999) along with specialized similarities 

in larval morphology and similar larval wood 

boring feeding habits (Grehan, 1988; Craw et 

al., 1999; Grehan & Rawlins, 2003) suggests


these genera are closely related, if not sister 

taxa. Unlike Hepialiscus and Oxycanus, the 

vicariism of Endoclita and Aenetus is separated 

by only 80 km between the Endoclita locality of 

Bacan (next to Halmahera) and those of Aenetus 

in New Guinea and the nearby islands of Misool 

and Ceram (Fig. 16b). 

As with Hepialiscus/Oxycanus, this 

vicariant relationship may be understood as the 

result of a broad ancestral range encompassing 

Asia and Australasia that preceded the 

differentiation of each genus. The region of 

differentiation between Endoclita and Aenetus 

approximates the triple plate junction between 

the Asian, Philippine and Australian plates, 

suggesting that this tectonic convergence may 

have promoted the differentiation of the two 

genera. 

Even if a close phylogenetic affinity 

between Endoclita and Aenetus was not 

supported by future analysis, the vicariant 

distributions would still indicate that their 

ancestral ranges were vicariant and only brought 

into contact through tectonic convergence in the 

region of their respective distributions coming 

into contact. The evolutionary origin of a 

common ancestral range for Endoclita and 

Aenetus may also involve the ancestral range of 

related genera, such as Phassus Walker, 1856, in 

Latin America (Grehan & Rawlins, 2003). 

CONLCUSIONS 

The geographic and phylogenetic patterns 

described here are attributed to the eastern Asian 

genera having originated from ancestral 

distributions that ranged between the Russian 

Far East/Northern Japan, South East Asia, and 

India. The ancestral ranges may have also 

represented a part of larger ancestral ranges that 

included these genera and their closest relatives 

outside what is now eastern Asia. This 

possibility is suggested for the vicariant 

relationship of the Hepialiscus group with 

Oxycanus, and Endoclita with Aenetus, that each 

originated from an ancestral distribution that 

formerly extended between Asia and 

Australasia. In the case of Endoclita and 

Aenetus, the vicariant relationship remains 

geographically close. 

The Asian and Australasian regions are 

often treated as distinct biogeographic entities 

(Olsen et al., 2001), but they are 

biogeographically indistinguishable with respect 

to the multitude of phylogenetic groups that 

have differentiated across both areas and with 

distributions that are spatially correlated with 

Mesozoic and Tertiary geology (Heads, 2005). 

The spatial proximity of the 

northwestern boundary of Oxycanus, the 

northwestern boundary of Aenetus, and the 

southeastern boundary of Endoclita to the triple 

junction between the Asian, Australian and 

Philippine tectonic plates suggests a historical 

relationship between the differentiation of these 

genera and the geology of the region. 

Given the biodiversity prominence of 

the Lesser Sunda, the Celebes and nearby 

islands (Myers et al., 2000; Olson et al., 2001), 

it is ironic that this region is so poorly known for 

the presence or absence of Hepialidae, 

particularly for the lack of published records of 

Endoclita between Java/Borneo and Halmahera. 

ACKNOWLEGEMENTS 

I am very grateful to John Dugdale for feedback 

and insights on hepialid phylogeny, to Carlos 

Mielke and Thomas Simonsen for comments on 

the manuscript, to Michael Heads for discussion 

on the biogeography, and to Shunsuki Utsumi 

for information on Japanese Hepialidae. I am 

also grateful to the Natural History Museum 

(London) for the permission to reproduce 

specimen images. 

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