Review of the millipede genus Orthomorpha Bollman, 1893 (Diplopoda, Polydesmida, Paradoxosomatidae) in Vietnam, with several new records and descriptions of two new species

Abstract The genus Orthomorpha is shown to currently be represented in Vietnam by ten species or varieties, including new records of O. arboricola (Attems, 1937), O. coarctata (de Saussure, 1860), O. rotundicollis (Attems, 1937) and O. scabra Jeekel, 1964, and two new species, O. caramelsp. nov. and O. vietnamicasp. nov. A key to all eight Orthomorpha species and two varieties known to occur in Vietnam is provided. Although the morphological characters that have been traditionally used for Orthomorpha taxonomy are here considered superior to molecular ones, molecular-based phylogenetic relationships and taxon assignments within the tribe Orthomorphini are provisionally analyzed using fragments of the cytochrome c oxidase subunit I (COI) mitochondrial gene. The preferred phylograms, both rooted and unrooted, demonstrate the monophyly of the tribe Orthomorphini, but due to the special, uncertain or even controversial position of O. coarctata, which occurs closer to the genera Antheromorpha and Hylomus, the genus Orthomorpha in current usage appears to be polyphyletic. However, if O. coarctata is to be treated within the monotypic genus Asiomorpha, the monophyly of Orthomorpha becomes manifest. On the other hand, a cautious approach is followed to avoid descriptions of suspicious new taxa/species. Thus, solely because the average genetic distance between O. rodundicollis subrotundicollisvar. nov. and O. rodundicollis, as well as that between O. scabra grandisvar. nov. and O. scabra, are both found to be negligibly small, the statuses of the sympatric and closest yet morphologically different varieties are treated only as such, i.e., infrasubspecific categories. The apparent discord observed between morphological and molecular data is obviously due to only partial and single-gene topologies used, possibly also to hybridization already known to occur in some closely related and sympatric paradoxosomatid species or even genera.


Introduction
The Southeast Asian millipede genus Orthomorpha Bollman, 1893 is the largest in the tribe Orthomorphini, family Paradoxosomatidae (Nguyen and Sierwald 2013). This genus has recently been reviewed and shown to comprise as many as 54 species ranging from northern Myanmar and Thailand in the northwest to Lombok Island, Indonesia in the southeast (Likhitrakarn et al. 2011(Likhitrakarn et al. , 2014. Only one species, O. coarctata (de Saussure, 1860), has attained a pantropical distribution due to anthropochory.
Only six recognized species of Orthomorpha have hitherto been known to occur in Vietnam. Among them, O. hydrobiologica (Attems, 1937), O. rotundicollis (Attems, 1937), and O. scabra Jeekel, 1964 are widespread also in Laos and Cambodia (Likhitrakarn et al. 2011(Likhitrakarn et al. , 2013(Likhitrakarn et al. , 2014. This paper is devoted to descriptions of two new species from southern Vietnam, based on the material collected by one of us (IS) in the Cat Tien, Kon Ka Kinh, Hon Ba and Bidoup Nui Ba national parks, all being priority zones for the protection of biodiversity in Vietnam (e.g., Polet and Ling 2004). In addition, we provide new records of three previously described species and a key to all eight Vietnamese species of the genus, with their distributions mapped.
Molecular analyses were performed to obtain a phylogram in order to visualize the phylogenetic relationships. The topology from MrBayes was consistent with the present-day morphology-based taxonomy/classification of the tribe Orthomorphini (Nguyen and Sierwald 2013).

Materials and methods
The specimens were collected as part of IS' research project on the diversity, biology and ecology of millipedes in Vietnam. Field-work by IS, including material collection, was conducted in accordance with Agreement 37/HD for the scientific cooperation between the Cat Tien National Park and the Joint Russian-Vietnamese Tropical Center; that in the Kon Ka Kinh National Park according to Agreements 432/TCLN-BTTN and 142/SNgV-VP; and that in the Bidoup Nui Ba National Park and Hon Ba Nature Reserve according to Agreement 774/TCLN-DDPH. The Animal Care and Use Protocol Review No. 1723018 was strictly followed. Coordinates and elevations were recorded by Garmin eTrex 30 using the WGS84 datum and subsequently doublechecked with Google Earth.
Live animals were photographed in their habitats using a Canon PowerShot A4000IS 16.0 MP Digital Camera and Panasonic DMC-TZ80 -LUMIX Digital Camera. Specimens were preserved in 96% ethanol, and morphological characters were studied in the laboratory using an Olympus stereo microscope. Scanning electron micrographs (SEM) of coated gonopod specimens were taken using a JEOL, JSM-5410 LV microscope. Specimens were also photographed in the laboratory as digital images assembly using automontage software techniques, while gonopods were also illustrated. All material, including the holotypes and voucher specimens for molecular analyses, is housed in the collection of the Zoological Museum, State University of Moscow (ZMUM), Russia.
In the catalogue sections, D stands for the original description, subsequent descriptive notes or appearance in a key, R for a subsequent record or records, N for giving a new name while M for a mere mention and Di stands for genetic distances in the molecular analyses.
DNA was isolated from specimens fixed in 96% ethanol using QiaAMP DNA Mini Kit (Qiagen). Sequences of the cytochrome oxidase subunit I (COI) gene were amplified using an EncycloPlus PCR kit (Evrogen, Russia) with the primer set COI-1F20 (5'-ACT CTA CTA ATC ATA AGG AT-3') and COI-1R19 (5'-TAA ACC TCC GGG TGA CCAA-3') derived from Nguyen et al. (2017). Polymerase chain reaction (PCR) products were visualized with the help of gel electrophoresis, excised, and cleaned using a SV Gel and PCR CleanUp System kit (Evrogen, Russia). The sequences were combined and aligned using ClustalX software after the addition of sequences from the GenBank (Thompson et al. 1997). MEGA 6.06 (Tamura et al. 2013) was used to calculate the genetic distances between samples. Of the trees in the post burnin posterior distribution inferred with MrBayes version 3.2.3, a consensus tree was constructed using sumt (Huelsenbeck and Ronquist 2001). The MrBayes analysis was conducted in the CIPRES server with the evolutionary model selected based on the results of the analysis in jModelTest2 (Darriba et al. 2015). The sequences have been deposited in NCBI GenBank (Table 1). Unfortunately, as there were fixation problems with the DNA, only around a third of the COI gene became available for analyses.  (Attems 1937(Attems , 1938. New material examined. 2 ♂ (ZMUM), Vietnam, Khanh Hoa Province, Hon Ba Nature Reserve, 12°07'02"N, 108°56'45"E, 1,550 m a.s.l., mixed mossy forest on mountain ridge, on forest floor, night time, 27.VI.2018, I. Semenyuk leg.
Live colouration (Fig. 1A) and colouration in alcohol, after one year of preservation, similar, only slightly faded in fixed material, yellow-brown to brown ( Fig. 1B-F), paraterga, most of mid-dorsal parts of metaterga and tip of epiproct orange-yellow to light yellow, legs and sterna light dark brown to light yellowish, antennae blackish (Fig. 1A, B).
Remarks. This species is known not only from type material from southern Vietnam (Dalat and Peak Lang Biang, at both sites found to coexist with O. rotundicollis (Attems, 1937)), but also from the new samples cited above. The types have recently been redescribed by Likhitrakarn et al. (2011) (Attems, 1937), ♂ from Hon Ba National Park A habitus and live colouration B, C anterior part of body, dorsal and lateral views, respectively D, E segments 10 and 11, dorsal and lateral views, respectively F-H posterior part of body, dorsal, ventral and lateral views, respectively I, J sternal cone between coxae 4, subcaudal and sublateral views, respectively.  Attems, 1927: 63 (D). Synonymized by Jeekel (1968 Remarks. This pantropical anthropochore species has been redescribed several times and recently revised (Likhitrakarn et al. 2011). It is often assigned to the monotypic genus Asiomorpha Verhoeff, 1939 (e.g., Nguyen andSierwald 2013), but we follow Jeekel (1968) who treated it in Orthomorpha. Molecular evidence, however, shows a very strong genetic isolation of coarctata from the other formal congeners,  (Attems, 1937), left gonopod A, B mesal and lateral views, respectively C tip of right gonopod, lateral view. Scale bars: 0.5 mm. thus favouring the recognition of Asiomorpha as a separate monobasic genus (Fig 20). This species has already been recorded from Vietnam, on sea shore north of Nhatrang (Golovatch 2017 Attems, 1927, from Ambon, Indonesia, has long been synonymized under O. c. coarctata (see Jeekel 1968Likhitrakarn et al. 2011).
Remarks. This species has recently been redescribed (Likhitrakarn et al. 2011), based both on type and non-type material. The species is widespread, probably due to anthropochory, especially along the coastal areas ranging from northern Vietnam down to southern Cambodia.
Colouration in alcohol, after one year of preservation, blackish to black-brown ( Fig. 3A-F), paraterga and epiproct orange-yellow to light yellow; legs and sterna light yellow to light brown; antennae yellowish to dark brownish distally (Fig. 3A). Colour polymorphism evident, colouration of paraterga generally ranging from white or yellow to red in one and the same population (see also below under Remarks).
Remarks. This species has recently been redescribed (Likhitrakarn et al. 2011), based on type material, and it is widespread from Luang Prabang Province, northern Laos to Cat Tien National Park, southern Vietnam (Fig. 19). The fresh specimens agree nearly fully with the available descriptions, except for the ♂ tarsal brushes being present until legs 10-14 vs. legs 5, and the pleurosternal carinae represented by small teeth gradually reduced until segment 15. These variations, as well as colour morphs, are certainly not more than infraspecific (cf. Likhitrakarn et al. 2011). One variety, subrotundicollis var. nov., is more disjunct morphologically, but not genetically, and is treated separately below.
The biology and behaviour of this species, referred to as Orthomorpha sp., has recently been described in detail in Cat Tien National Park (Semenyuk and Tiunov 2018). Millipedes are abundant almost throughout the year with a slight decline in the dry season (winter and early spring). Juveniles start swarming in the rainy season, mainly on logs or other decaying wood debris, also in suspended soil in tree holes. Swarms are active mostly in the night but can often be seen also in the daytime. Juveniles of later instars do not swarm, but still tend to group.
Colouration of live animals (Fig. 6A) black-brownish with contrasting dark yellow to orange paraterga and epiproct, head and antennae brownish, legs pale brownish; colouration in alcohol, after one year of preservation, faded to black-brown ( Fig. 6B-H), paraterga and epiproct pale whitish yellow or pale brown, legs whitish to pale brown distally.
Remarks. This species has recently been redescribed (Likhitrakarn et al. 2011), based on type material. The new specimens agree nearly fully with the available descriptions except for the ♂ tarsal brushes being present until legs 7 or 8 vs. legs 5. One variety, grandis var. nov., is more disjunct morphologically, but not genetically, being treated separately below.
This species seems to be widespread, occurring in Vietnam and northern Laos (Fig. 19), and in two places it was found coexisting with O. rotundicollis (Attems, 1937) (see also above).
According to IS' field observations, the millipedes occupy a wide range of habitats, including riparian forests with flooding occurring every year and non-flooding mixed forests on slopes. They also appear in a forestless farm area, usually being found on the forest floor and at bases of tree trunks. They occupy many microhabitats in lower forest strata. The activity is mainly restricted to the night time, but some individuals keep it up in daylight as well.
Orthomorpha rotundicollis (Attems, 1937), subrotundicollis var. nov. Name. To emphasize the strong similarity to the typical O. rotundicollis (Attems, 1937). Normally, no Latin names are to be applied to varieties as infrasubspecific categories, but because this new variety had first been qualified and described as a new species based on purely morphological grounds before the molecular evidence showed it to be the same as O. rotundicollis, we allot it the previously chosen name subrotundicollis and treat it separately in our analyses, key, and map.
Diagnosis. Using the latest key (Likhitrakarn et al. 2011), this new variety keys out and seems to be especially similar to the typical O. rotundicollis, but it differs in the smaller size (up to 23 mm (♂) or 29 mm (♀) long and 2.9-3.2 mm (♂) or 3.3-3.9 mm (♀) wide, respectively), and the pleurosternal carinae being complete crests until segment 7 (♂) or 5 (♀), each crest supplied with an evident sharp denticle caudally, Figure 9. Orthomorpha rotundicollis (Attems, 1937), subrotundicollis var. nov., ♂ A habitus and live colouration B, C anterior part of body, dorsal and lateral views, respectively D, E segments 10 and 11, dorsal and lateral views, respectively F-H posterior part of body, dorsal, ventral and lateral views, respectively I, J sternal cones between coxae 4, subcaudal and sublateral views, respectively. thereafter increasingly strongly reduced until segment 16 (♂, ♀), coupled with tarsal brushes being traced until ♂ legs 11.
Colouration of live animals blackish (Fig. 9A), paraterga and epiproct contrasting yellow-orange, head and antennae dark brownish, legs pale brownish; colouration in alcohol, after one year of preservation, blackish or faded to black-brown ( Fig. 9B-H), paraterga and epiproct pale whitish yellow to light yellow, legs and sterna light yellow to pale brown, antennae light yellow to dark brownish distally.  (Attems, 1937), subrotundicollis var. nov., ♂, left gonopod A, B mesal and lateral views, respectively C distal part, submesal view. Scale bars: 0.2 mm.
Remarks. Based on morphological characters alone (see Diagnosis), this variety had first been qualified and distinguished as a full new species before the molecular analyses unequivocally showed it to be genetically the same as O. rotundicollis. Indeed, the average genetic distance (Di) of subrotundicollis from the typical, morphologically closest and sympatric O. rodundicollis is null (Table 2). In one of the phylograms (Fig. 20), subrotundicollis falls out between the two individuals of rotundicollis, and overall the genetic variation is strikingly scant. This is one of the direct consequences of the molecular analyses accepted in our study.
Adults of this variety were found only in May during a short expedition to Kon Ka Kinh National Park, as part of IS' research on the diversity, biology, and ecology of millipedes in Vietnam. The previous trip to the same area in 2016 failed in finding this species. In 2017, millipedes were very abundant in many types of forest ranging from native to highly disturbed ones, up to even forestless hills with farmed crops. The activity was mainly noted in the night time in forest, but walking millipedes appeared even in daylight in open areas. Mating was recorded in May.

Table 2.
Average genetic distances (Di) among the available taxa, the matrix constructed using Kimura's two-parametric/pairwise model of nucleotide replacements that suggests a considerable prevalence of transitions over transversions in mitochondrial DNA. Old information is available in Nguyen et al. (2017Nguyen et al. ( , 2018 and GenBank, the new one is contained in Table 1. Name. A noun in apposition, to emphasize the general caramel colouration of the animals. Diagnosis. Using the latest key (Likhitrakarn et al. 2011), distinguished from all known congeners by the tip of the solenophore being very faintly bifid, with a nearly smooth terminal lobe bearing a minute lobule at the base; in the gonopod structure it is similar to O. tuberculifera Likhitrakarn, Golovatch & Panha, 2011, but differs in the rather smooth and shining dorsal tegument devoid of tubercles, coupled with the particular colouration.
Colouration of live animals dark chocolate brown (Fig. 12A); metaterga, paraterga and epiproct caramel in colour; head, antennae and legs dark brownish; colouration in alcohol after one year of preservation chocolate brown or faded to light brownish ( Fig. 12B-H); metaterga, paraterga and epiproct caramel to light brownish in colour; legs and sterna light brown to pale yellow; head and antennae dark brownish to brown.
Paraterga very strongly developed (Fig. 12B-H), mostly upturned, all lying below dorsum, set at ca. upper 1/4 of midbody height, nearly level with dorsum on segments 15-18, caudal corner always spiniform and narrowly rounded, extending beyond rear tergal margin; in lateral view, paraterga thinner in poreless segments and modestly enlarged in pore-bearing ones.
Sterna sparsely setose, without modifications except for two rather large and long, fully separated, sternal cones between ♂ coxae 4 (Fig. 12I, J). A paramedian pair of evident tubercles in front of gonopod aperture. Legs long and slender, midbody ones ca. 1.2-1.4 times as long as body height, prefemora without modifications, ♂ tarsal brushes present until legs 15.

Remarks.
The biology and behaviour of this species are very similar to those of O. vietnamica sp. nov. During field observations in May 2017, millipedes occurred mainly on tree trunks. Mating was also recorded, but no females were collected. Paratype: 1 ♀ (ZMUM Rd 4200), same locality, together with holotype. Name. Adjective to refer to the country of origin. Diagnosis. This species seems to be especially similar to O. caramel sp. nov., but differs from all congeners (Likhitrakarn et al. 2011) in the presence of a conspicuous, Figure 15. Orthomorpha vietnamica sp. nov., ♂ holotype A, B anterior part of body, dorsal and lateral views, respectively C, D segments 10 and 11, dorsal and lateral views, respectively E-G posterior part of body, lateral, dorsal and ventral views, respectively H, I sternal cones between coxae 4, subcaudal and sublateral views, respectively. densely setose, rounded tubercle (Fig. 16, cxp) on the gonocoxa and the evident trifid tip of the solenophore, as well as the ♂ tarsal brushes showing until segment 16, coupled with the pleurosternal carinae present as complete crests with a sharp caudal tooth on segments 2-4.
Colouration of alcohol material after one year of preservation dark brown (Fig. 10B-H); metaterga brown to light brownish; paraterga and epiproct light yellow to yellowish; legs and sterna light brown to pale yellow; head, collum, and antennae dark brownish to brown. Bases of paraterga marbled (Fig. 15C).
Axial line visible on collum and both on following pro-and metazonae. Transverse sulcus usually distinct (Fig. 15C, E, H), complete on metaterga 5-18, incomplete and nearly wanting on segments 4 and 19, nearly line-shape, rather shallow, almost reaching the bases of paraterga, at most very faintly beaded at bottom.
Remarks. We assume the apicodorsal lobule of the solenophore to be broken off, considering the structure of the solenophore tip in the other Orthomorpha spp.
The ecology of this species is very similar to that of O. caramel sp. nov. These two species are syntopic and share the same microhabitats.   Jeekel, 1964, grandis var. nov., ♂ A habitus and live colouration B, C anterior part of body, dorsal and lateral views, respectively D, E segments 10 and 11, dorsal and lateral views, respectively F-H posterior part of body, dorsal, ventral and lateral views, respectively I, J sternal cones between coxae 4, subcaudal and sublateral views, respectively. Name. Adjective to emphasize the unusually large size of the animals. Normally, no Latin names are to be applied to varieties as infrasubspecific categories, but because this new variety had first been qualified and described as a new species based on purely morphological grounds before the molecular evidence showed it to be the same as O. scabra, we allot it the previously chosen name grandis and treat it separately in our analyses, key, and map. This is also one of the direct consequences of the molecular analyses accepted in our study.
Colouration in alcohol after ten months of preservation dark brown (Fig. 17A-G); metaterga, paraterga and epiproct red-brown to dark brown; legs, antennae and sterna light brown to light yellow; head brownish to brown.
Following paraterga very strongly developed (Fig. 17A-F), upturned, lying below dorsum until segment 4, following ones above dorsum, caudal corner ranging from obtuse-angular to subrectangular, always and increasingly extending behind rear tergal margin ( Fig. 17A-C), from segment 15 on spiniform, long and pointed; in lateral view, paraterga thinner in poreless segments and modestly enlarged in pore-bearing ones.
Sterna sparsely setose, without modifications except for two rather large and long, fully separated, sternal cones between ♂ coxae 4 (Fig. 17H, I). A paramedian pair of small tubercles in front of gonopod aperture. Legs long and slender, midbody ones ca. 1.3-1.4 times as long as body height, prefemora without modifications, ♂ tarsal brushes present until legs 7.
Remark. This species was found only on Mount Bidoup in the summit area, while none of the inspected adjacent forests, even those located at the same altitudes, including a very similar forest on Mount Hon Giao, failed to reveal these millipedes. On Mount Bidoup, this species was quite abundant, occurring mainly on the forest floor, on logs and at bases of tree trunks in the night time, as well as under logs in the daytime.  A single sternal cone between ♂ coxae 4 ( Fig. 1H, I). Distal part of gonopod evidently curved and tip bifid (Fig. 2) (Fig. 15B, D, E). Tarsal brushes present until segment 16 (♂). Tip of gonopod evidently trifid and coxa with a conspicuous, densely setose, rounded, distoventral tubercle (cxp) (Fig. 16A, B) Larger (29-38.5 (♂) or 31-40.5 mm (♀) long and 3.5-4.3 (♂) or 3.6-4.9 mm (♀) wide, respectively). Pleurosternal carinae complete crests on segments 2-4 (♂, ♀), each with an evident sharp denticle caudally, thereafter increasingly strongly reduced until segment 10 (♂). Tarsal

Conclusions
The phylogenetic analyses performed here are considered as supporting to the traditional morphological taxonomy since it is a single-gene topology. While it should provide some resolution to disentangling Orthomorpha evolutionary history, the limitations of using just one gene are evident. The more so as, due to fixation problems with the DNA, we have only been able to get approximately one-third of the COI gene. In addition, not only sympatric and closely related species, but even certain genera of Paradoxosomatidae sometimes seem to show hybridization (Decker 2018). As our study shows (Tables 1, 2, Fig. 20), within the tribe Orthomorphini, almost all particularly closely related species belong to the genus Orthomorpha. Only O. coarctata is set apart and appears to be closer to other genera (Table 2). This is clearly seen also in Figure 20. Again only O. coarctata shows a special position clustering closer to Antheromorpha Jeekel, 1968 andHylomus Cook &Loomis, 1924 than to other formal congeners. This is hardly surprising though, because that species is often treated as the sole constituent member of a separate genus, Asiomorpha Verhoeff, 1939 (cf. Nguyen andSierwald 2013). However, we prefer to stick here to the traditional, purely morphologically based concept of the Orthomorphini and Orthomorpha, taking most of our and all previous molecular evidence as only subordinate, provisional, and auxiliary.

Figure 20.
A rooted phylogram based on COI sequences obtained from the relevant literature on Paradoxosomatidae (Nguyen et al. 2017(Nguyen et al. , 2018 and available in GenBank (all given in black, yet with several name updates according to modern taxonomy), combined with new haplotypes (six species and two varieties, these latter put in quotation marks and all highlighted in red). Tree topology was reconstructed using MrBayes software. The values of statistical support are given above the branches only when these exceed 65%.
We present here only a phylogram rooted in Apheloria virginiensis (Drury, 1770) (Xystodesmidae) as perhaps the closest outgroup currently available (Fig. 20). Even though the presence or absence of a root can drastically change tree topology, because our unrooted trees completely failed to alter the picture, they have been omitted. The same applies to a number of other rooted phylograms.
However, as noted above, two important taxonomic changes have been accepted following solely our molecular analyses and trees. We simply follow a cautious approach to avoid descriptions of suspicious new taxa/species. These changes concern the admittance of two morphologically distinct (especially so in body size), but genetically unsupported varieties, O. rotundicollis subrotundicollis var. nov. and O. scabra grandis var. nov. Because neither is distinguished from their typical forms by average genetic distances (Table 2), they have been allotted no taxonomic status. The other results, however, clearly suggest that more molecular and morphology-based taxonomic work should be paid to Orthomorpha in the future. The animals are large-sized, often vividly coloured and readily exposed, thus being easy to spot and collect. In addition, Orthomorpha enjoys a recent, colourful, open-access, monographic treatment with an identification key to species (Likhitrakarn et al. 2011).
In any event, although the trees both in Nguyen et al. (2017Nguyen et al. ( , 2018 and our present research must only be taken as provisional, they provide a basis for future phylogenetic analyses implementing additional gene regions. Further efforts, both morphology-and molecular-based, are required to reveal and refine the relationships between Orthomorphini (and not only) genera and species. At least for the time being, the Orthomorphini seems to be a monophyletic tribe of Paradoxosomatidae, whereas Orthomorpha remains polyphyletic and Antheromorpha is paraphyletic (Fig. 20).
A total of eight species and two morphologically distinct varieties of Orthomorpha is currently recognized in the fauna of Vietnam, including two new congeners: O. caramel sp. nov. and O. vietnamica sp. nov. Both new species come from national parks and surrounding areas, thus priority zones for the protection of biodiversity in Vietnam. Several, often two or three species co-exist in one place and are strictly syntopic, this having been observed, e.g., in the pristine tropical forest of the Kon Ka Kinh National Park (Fig. 19).
Most of the Vietnamese species tend to be restricted to the southern parts of Vietnam, except for the likely anthropochore O. hydrobiologica which ranges from Hong Gai, Quảng Ninh Province, northern Vietnam to southern Cambodia. The range of O. coarctata in Vietnam must be even greater, covering most of the country in man-made habitats. No native species seem to exist in northern Vietnam. A similar distribution pattern concerns Laos where no native Orthomorpha seem to populate the northern parts of the country (Likhitrakarn et al. 2014).
There can be little doubt that further new Orthomorpha species and/or records await discovery in Indochina, including Vietnam. This seems particularly topical in the still persisting forested areas of that huge region.