New leaf- and litter-dwelling species of the genus Pholcus from Southeast Asia (Araneae, Pholcidae)

. We describe eight new species of the genus Pholcus , and document their microhabitats. Four species are assigned to the previously described Pholcus ethagala group: P. tanahrata Huber sp. nov., P. uludong Huber sp. nov., and P. bukittimah Huber sp. nov. from the Malay Peninsula, and P. barisan Huber sp. nov. from Sumatra. These species are all litter-dwellers that build domed sheet webs on the undersides of large dead leaves on the ground. The other four species are assigned to newly created species groups: the P. tambunan group with two species from northern Borneo: P. tambunan Huber sp. nov. and P. bario Huber sp. nov.; and the P. domingo group with two species from the Philippines, Mindanao: P. domingo Huber sp. nov. and P. matutum Huber sp. nov. These latter four species are leaf-dwellers that build barely visible silk platforms tightly attached to the undersides of live leaves. The main rationale for this paper is to provide part of the taxonomic and natural history background for upcoming phylogenetic and evolutionary (microhabitat shifts) analyses.


Introduction
Pholcidae occupy a wide range of microhabitats, particularly in tropical forests.Some prefer leaf litter, some live in larger sheltered spaces among rocks and logs and in caves, some build their webs freely among vegetation, and some are found on the undersides of live leaves up to several meters above ground.While some genera are homogeneous in this respect (e.g., all Ninetinae seem to be ground-dwellers; all Calapnita Simon, 1892 seem to be leaf-dwellers), others include representatives from various different microhabitats (e.g., Belisana Thorell, 1898;Mesabolivar González-Sponga, 1998;Metagonia Simon, 1893;Modisimus Simon, 1893; Smeringopus Simon 1890; see Huber 2005, 2012, 2015and Huber et al. 2005, 2010).This implies that evolutionary microhabitat shifts have occurred, both among and within genera.Several such shifts have recently been documented in some detail (e.g., Dimitrov et al. 2013;Huber et al. 2015), but our general understanding of the patterns is still rudimentary because it requires good data about both microhabitat and phylogeny, and at least one of them is often missing or incomplete.European Journal of Taxonomy 200: 1-45 (2016) Of particular interest in this respect is the genus Pholcus Walckenaer, 1805.Almost every microhabitat accessible to Pholcidae in general is occupied by specific representatives of this large genus.Most species groups in Pholcus that appear well supported are homogeneous with respect to their microhabitat, but relationships among groups do not seem to correlate with microhabitat preferences and in some cases even closely related species occupy different microhabitats (Dimitrov et al. 2013;Huber & Dimitrov 2014).This suggests multiple convergent shifts of microhabitat in Pholcus alone.Preliminary phylogenetic data suggest that the story may be even more complicated.Several genera currently considered close to Pholcus, such as Calapnita Simon, 1892;Leptopholcus Simon, 1893;Micropholcus Deeleman-reinhold & Prinsen, 1987;Micromerys Bradley, 1877;Panjange Deeleman-reinhold & Deeleman, 1983;etc.(the Pholcus group of genera in Huber 2011) may actually be nested within a paraphyletic Pholcus (Huber 2011;A. Valdez-Mondragón, B.A. Huber & D. Dimitrov, unpublished data).If this proves to be correct, then these mostly leaf-dwelling taxa may represent numerous additional microhabitat shifts within this large taxonomic group (currently comprising some 400 described species).
The present paper focuses on Pholcus in Southeast Asia (Fig. 1) and is one of several recent contributions providing the taxonomic and natural history background for an upcoming analysis of microhabitat shifts in Pholcidae.Of particular interest in this respect is the Pholcus ethagala group because previous cladistic analyses (Huber 2011) have suggested a close relationship between this group (whose Southeast Asian representatives turn out to be litter-dwelling; e.g., Figs 2-9) and the leaf-dwelling genus Panjange.The other two species groups treated below are also 'untypical' Pholcus (e.g., they both lack an uncus on the male genital bulb) and might previously have been considered to represent new genera, but preliminary analyses of molecular data (A. Valdez-Mondragón, B.A. Huber & D. Dimitrov, unpublished data) strongly support their inclusion in Pholcus, i.e., a close relationship with more 'typical' Pholcus.We deliberately do not dwell on phylogenetic aspects in this paper as all species treated below will be included in the comprehensive molecular phylogeny to be published in 2017.

Material and methods
Most of the material studied herein was collected during recent expeditions to the Philippines (Feb.-Mar.2014), northern Borneo (Jul.-Aug.2014), and the Malay Peninsula (Feb.-Mar. 2015).This material is currently deposited at the following institutions: Mindanao State university -Iligan Institute of Technology, Iligan (MSu-IIT), Philippines; Museum of Zoology, university of Malaya, Kuala Lumpur (MZuM), Malaysia; Princess Maha Chakri Sirinthorn Natural History Museum, Prince of Songkla university, Hat Yai (PSuZC), Thailand; Sarawak Museum, Kuching (SMK), Malaysia; and Zoologisches Forschungsmuseum Alexander Koenig, Bonn (ZFMK), Germany.Further material came from the American Museum of Natural History, New York (AMNH); and the Netherlands Centre for Biodiversity Naturalis, Leiden (rMNH).
Methods and terminology used are as in Huber (2011).Measurements are in mm unless otherwise specified.Eye measurements are approx.± 5 µm.Epigyna were cleared in warm NaOH solution and stained with chlorazol black.For SeM photos, specimens were dried in hexamethyldisilazane (HMDS) (Brown 1993), and photographed with a Hitachi S-2460 scanning electron microscope.SeM data are presented within the descriptions but are not based on the holotype specimens described.Locality coordinates are in round brackets when copied from labels and original publications or when received directly from collectors, in square brackets when originating from some other source (such as online gazetteers, Google earth, etc.).The distribution map was generated with ArcMap 10.0.

Natural history
The seven species newly observed in the field (Malay Peninsula and Sumatra) were mostly found on the undersides of large dead leaves on the ground.This is in contrast to the two Sri Lankan species that supposedly live on live leaves (at least P. ethagala; Huber 2011).Very few specimens of the newly observed species were also found under logs (P.ledang) and in bamboo sheaths and under rocks (P.gombak).The availability of suitable large leaves on the forest floor strongly influenced spider abundance.Webs consisted of small domed sheets closely attached to the leaf surface.When disturbed, the spiders barely reacted; they vibrated only for a short time at low amplitude or were not seen to vibrate at all.Two egg-sacs (of two species) were parasitized by Idris Foerster, 1856 (Scelionidae, Baeini) wasps 62).For further information see individual descriptions below.

Distribution
The P. ethagala group is known from Sri Lanka (see Huber 2011; not treated herein and not shown in Fig. 1) and from Southeast Asia (Malay Peninsula, Sumatra; Fig. 1).Huber, 2011 Figs 2-3

Description -amendments
Tibia 1 in three newly collected males: 9.

Etymology
The species is named for Tanah rata, the town close to the type locality; noun in apposition.
Color.Carapace ochre-yellow with light brown triangular mark posteriorly, ocular area, eye stalks and clypeus light brown, some irregular back pigment in AMe area; sternum whitish, labium brown; legs ochre-yellow with darker brown patellae and tibia-metatarsus joints; abdomen ochre-gray with some dark marks dorsally and laterally, monochromous ventrally.
CheliCerae.As in Fig. 12, with single pair of frontal proximal apophyses directed towards median; without distal modification; without stridulatory ridges.
PalPs.As in Figs 10-11; coxa unmodified; trochanter with very long curved retrolatero-ventral apophysis and small weakly sclerotized retrolateral process; femur with indistinct retrolatero-dorsal hump proximally and ventral hump more distally; tarsus with large rounded elongation; procursus complex, proximal part with large ventral 'knee' and complex dorsal and prolateral processes; hinged distal part with distinctive flat dorsal process and whitish areas retrolaterally and prolaterally, transparent processes prolaterally; bulb with very large appendix, long slender embolus, with small uncus-like process partly sclerotized as continuation of proximal bulbal sclerite (hidden by bulb in Fig. 10).Male (variation) Tibia 1 in other male: 9.7; this second male died while molting and is thus very pale; genitalia identical to holotype.

Natural history
All specimens were found under large dead leaves on the ground.Such leaves were very rare at the type locality, which explains the low number of specimens despite several hours of intensive search.

Distribution
Known from type locality in Malaysia only (Fig. 1).

Etymology
The species name is derived from the type locality; noun in apposition.

Natural history
All specimens were found under large dead leaves on the ground.They barely reacted to disturbance.

Diagnosis
easily distinguished from most similar known relatives (other species of the P. ethagala group on the Malay Peninsula) by morphology of male palps (figs 819-820 in Huber 2011; short ventral trochanter apophysis; procursus proximal part with prolateral rather than dorsal process, with large retrolateroventral whitish area, with distinctive complex tip), and by details of female internal genitalia (figs 821-822 in Huber 2011; pore plates widening anteriorly).

Natural history
Specimens were found under tree logs and under dead leaves on the ground.They barely moved when disturbed, other than a short burst of low-amplitude vibration.

Etymology
The species is derived from the type locality; noun in apposition.
Color.Carapace pale ochre-yellow with light brown median V-mark posteriorly, clypeus light brown, ocular area not darkened; sternum whitish; legs ochre-yellow with darker brown patellae and tibiametatarsus joints; abdomen gray with some black marks dorsally and laterally.
PalPs.As in Figs 67-68; coxa unmodified; trochanter with very long curved retrolatero-ventral apophysis and small weakly sclerotized retrolateral process; femur with indistinct ventral humps proximally and distally; tarsus with large rounded elongation; procursus complex, proximal part with distinctive dorsal process; hinged distal part with large whitish area retrolaterally and transparent fringed processes prolaterally; bulb with very large appendix, long slender embolus, and small uncus-like process partly sclerotized as continuation of proximal bulbal sclerite.legs.Without spines and curved hairs; few vertical hairs; retrolateral trichobothrium on tibia 1 at 2%; prolateral trichobothrium absent on tibia 1, present on other tibiae; tarsus 1 pseudosegments very indistinct, apparently irregular.

Female
In general similar to male (Fig. 62) but eye triads not on stalks and much closer together than in male (PME-PME distance: 200 µm); clypeus and ocular area frontally dark brown, with pair of small black marks in area of AMe (but without lenses).Tibia 1 in 1 female: 5.8.epigynum weakly sclerotized (Fig. 87), only posterior rim slightly more sclerotized, with small dark 'knob' (Fig. 70

Natural history
All specimens were found in an area of the forest where large dead leaves on the ground were abundant.They built their domed webs closely attached to the undersides of the leaves.A few eggs in an egg-sac of one female were parasitized by Idris wasps (Fig. 62).

Distribution
Known from type locality in Singapore only (Fig. 1).relative straight male palpal trochanter apophysis similar to P. uludong Huber sp.nov.and P. phui but shorter; shape of relatively small appendix), by modified male clypeus (pair of small dark processes; Fig. 80), by male eye stalks directed more towards lateral (Fig. 80), and by details of female internal genitalia (Figs 75, 76; anterior undulating sclerite; large posterior semicircular structure).

Etymology
The species name is derived from the type locality; noun in apposition.
Color.Carapace pale ochre-yellow with large dark brown mark posteriorly, dark median line, ocular area, stalks and clypeus also dark brown; irregular black marks in place of AMe; sternum whitish; legs ochre-yellow with darker brown to black patellae and tibia-metatarsus joints; abdomen ochre-gray with some darker marks dorsally and laterally.

Female
In general similar to male (Figs 65-66) but eye triads not on stalks and much closer together than in male (PME-PME distance: 240 µm); dark posterior mark on carapace smaller than in male, without median dark line on carapace; with pair of small black marks in area of AMe (but without lenses); clypeus unmodified and only in upper third dark brown.Tibia 1 in 6 females: 6.0-6.9 (mean 6.4).Some females (and juveniles) with laterally strongly widened abdomen (Fig. 65).epigynum weakly

Natural history
All specimens were found under large dead leaves on the ground.They hung in domed sheet webs closely attached to the leaf surface.They barely moved when disturbed and were not seen to vibrate.

Distribution
Known from type locality in Sumatra only (Fig. 1).

Description
See individual descriptions of the two very similar species below.

Composition
Only the two species described below.

Natural history
Both species were found on the undersides of live leaves where they built very fine webs tightly attached to the leaf surface.Both species were frequently seen to share their webs with cecidomyiid flies.For further information, see descriptions below.

Distribution
The P. tambunan group is restricted to northern Borneo (Fig. 1).
CheliCerae.As in Fig. 103, with distinctive pair of frontal apophyses (large flat plates, distally drawn out into small tip; Fig. 108) and rounded lateral humps; without modified hairs; without stridulatory ridges.

Female
In general similar to male (Fig. 96) but carapace mark only in posterior half; ocular area dark as in male but clypeus light.eye triads much closer together than in male (Fig. 107; PMe-PMe distance: 265 µm).Dark marks on abdomen less distinct or even absent.Tibia 1 in 18 females: 6.2-7.2 (mean 6.7).epigynum weakly sclerotized (Fig. 125), wide transversal plate with scape directed towards anterior (Fig. 104); dark internal arch and lateral posterior structures visible through cuticle; internal genitalia as in Figs 105, 126, 127, with pair of highly distinctive three-layered telescopic tubes.ALS with one large widened, one pointed, and six smaller cylindrically-shaped spigots (Fig. 118).The female from Sepilok is assigned tentatively because no male is known from this locality.

Natural history
The spiders were found on the undersides of live leaves about 50-200 cm above the ground.They built barely visible silk platforms directly attached to the leaf surface rather than the 'usual' pholcid domes.Several webs were occupied (in addition to the spider) by large numbers of Diptera (probably Cecidomyiidae, not collected).

Distribution
Known from three localities in northeastern Borneo (Sabah) (Fig. 1; but note that the single specimen from Sepilok is assigned tentatively).

Etymology
The species name is derived from the type locality; noun in apposition.
Color.Carapace whitish with dark median mark widening anteriorly and including ocular area and clypeus in upper part and laterally; sternum whitish; legs pale ochre-yellow with dark brown patellae and tibia-metatarsus joints; abdomen pale ochre-gray with black marks dorsally and laterally, monochromous ventrally.
CheliCerae.As in Fig. 120, with distinctive pair of frontal apophyses (large flat curved plates) and small rounded lateral humps; without modified hairs; without stridulatory ridges.
PalPs.As in Figs 121-122; coxa unmodified; trochanter with strong ventral apophysis; femur cylindrical, with two small retrolateral humps proximally and small ventral hump at half length; procursus with strong ventral 'knee', with distinctive small prolateral sclerotized process at 1/3 length (arrow in Fig. 121), distal half with large whitish area retrolaterally, tip complex with distinctive sclerites; bulb elongated, long appendix with distinctive prolateral process provided with small teeth, further teeth along ventral rim, proximal processes of appendix 'guiding' embolus between them; embolus long and weakly sclerotized.legs.Without spines and curved hairs; few vertical hairs; retrolateral trichobothrium on tibia 1 at 2%; prolateral trichobothrium absent on tibia 1, present on other tibiae; tarsus 1 pseudosegments not seen in dissecting microscope.

Female
In general similar to male (Fig. 100) but carapace mark only in posterior half; ocular area dark as in male but clypeus light.Eye triads closer together than in male (PME-PME distance: 230 µm).Dark marks on abdomen less distinct.Tibia 1 in 2 females: 6.6 (both).epigynum weakly sclerotized (Fig. 128), wide transversal plate with long scape directed towards anterior (Fig. 123); dark internal arch and lateral posterior structures visible through cuticle; internal genitalia as in Figs 124, 129, 130, with pair of highly distinctive three-layered telescopic tubes.

Natural history
All specimens were found in one small section of forest along a small river.No further specimens were found in two days of intensive search.They built barely visible webs on the undersides of live leaves, and often shared webs with large numbers of Diptera (Cecidomyiidae, in ZFMK).

Distribution
Known from type locality in northern Borneo (Sarawak) only (Fig. 1).

Description
See individual descriptions of the two very similar species below.

Composition
Only the two species described below.

Natural history
Both species were found on the undersides of live leaves where they built very fine webs tightly attached to the leaf surface.For further information, see description of P. domingo Huber sp.nov.below.

Distribution
The P. domingo group is restricted to Mindanao (Fig. 1).

Diagnosis
Distinguished from most similar known relative (P.matutum Huber sp.nov.) by morphology of male palps (Figs 140-141; longer procursus with shorter dorsal process; appendix with long rod-shaped main branch); females of these two species are difficult to separate (internal genitalia relatively shorter in P. matutum Huber sp.nov.; compare Figs 144 and 160).

Etymology
The species name is derived from the type locality; noun in apposition.

Male (variation)
Dorsal process of procursus varies considerably among localities: in comparison to males from type locality (Santo Domingo; Fig. 146), it is slightly more slender in male from Malagos; considerably stronger and longer in male from Mt. Mupo (Fig. 147; males from CeDAr intermediate between males from type locality and male from Mt. Mupo); almost absent in males from Dimapatoy (Fig. 145).Male from Mt. Mupo also with slightly different appendix (distal part of sclerotized main branch slightly longer).Males from Dimapatoy with slightly shorter trochanter apophysis.All specimens other than those from Santo Domingo are therefore assigned tentatively to this species.Sternum in most males examined either monochromous whitish or whitish with a few darker marks posteriorly; only one male from Malangob, male from Malagos, and male from Mt. Mupo like holotype; male from Mt. Mupo with more complex carapace pattern (Fig. 137; similar to females from other localities).Tibia 1 in 15 other males: 7.4-9.0(mean 8.2).

Female
In general similar to male but sternum always dark brown and carapace pattern different: in most females posterior half with light brown to orange pattern (Fig. 136), one female from Dimapatoy with pattern like male holotype.Eye triads slightly closer together than in males (PME-PME distance: 200 µm).Tibia 1 in 19 females: 6.0-7.0 (mean 6.6).epigynum very simple, weakly sclerotized (Fig. 161), with transverse ridges and finger-shaped posterior 'knob' (Figs 143,156); internal genitalia as in Figs 144, 162, 163.ALS with one large widened, one pointed, and six smaller cylindrically-shaped spigots (Fig. 155).The single female specimen from epol Falls is assigned tentatively because no males are known from this locality.

Natural history
At Santo Domingo and CEDAR, the spiders were found in very fine, barely visible webs tightly attached to the leaf surface of live leaves about 50-200 cm above the ground.The same kinds of leaves were also occupied by two species of Calapnita, and sometimes a Pholcus and a Calapnita were seen sharing a leaf.When disturbed, the Pholcus just remained still, with their bodies pressed against the leaf; when disturbed with more insistence, they ran away on the leaf or even let themselves down on a silk line.In the egg-sac of one female from the type locality, most eggs were parasitized by a parasitic wasp (Fig. 132).

Distribution
Known from several localities in Mindanao (Fig. 1

Etymology
The species name is derived from the type locality; noun in apposition.
PalPs.As in Figs 157-158; coxa unmodified; trochanter with very long curved ventral apophysis; femur with ventral and retrolatero-dorsal processes (slightly different shapes than in P. domingo Huber sp.nov.); tibia very large (relative to femur); procursus rather simple, with strong ventral 'knee' and distinctive long dorsal process distally; bulb with long semitransparent embolus and large sclerotized process (putative appendix) densely covered with scales on retrolateral surface, with lighter prolateral branch proximally (appendix with more pointed tip than in P. domingo Huber sp.nov.; prolateral branch with more rounded tip).

Female
In general similar to male (Fig. 139), carapace pattern different: most females with two pairs of light brown to orange marks, one pair near median line posteriorly, one pair laterally; one female with same carapace pattern as P. domingo Huber sp.nov.(posterior half of carapace largely covered by light brown to orange pattern).Eye triads slightly closer together than in males (PME-PME distance: 230 µm).Tibia 1 in 4 females: 6.6, 6.9, 6.9, 7.2.Sternum whitish as is males (not brown as in P. domingo Huber sp.nov.females).epigynum very simple, weakly sclerotized (Fig. 164), with transverse ridges and finger-shaped posterior 'knob' (Fig. 159); internal genitalia as in Figs 160, 165, 166.

Natural history
The spiders were found in very fine, barely visible webs tightly attached to the leaf surface of live leaves.

Distribution
Known from two localities in southern Mindanao (Fig. 1).The single female specimen from epol Falls assigned tentatively to P. domingo Huber sp.nov.might actually belong to this species.

Discussion
None of the species described above is a 'typical' Pholcus, i.e., none of them belongs to what has been described as the "core group" of Pholcus (Huber 2011).Most representatives of this core group are characterized by the combination of two characters: a heavily sclerotized epigynal plate, and male chelicerae with proximal frontal apophyses.All species described above have very weakly sclerotized epigyna, and they lack proximal frontal apophyses on the male chelicerae (the apophyses in representatives of the P. ethagala group are considered homologues of the proximal lateral apophyses, a synapomorphy of Pholcinae; Huber 2011).This requires some justification for why we assign these species to Pholcus.
A first cladistic analysis of Pholcus (Huber 2011) has brought some structure into this large taxon (at that time 254 species; now almost 350 species), and it provided strong support for a group of genera close to Pholcus (the Pholcus group of ten genera).At the same time, it also highlighted some fundamental problems with Pholcus and its closest relatives.It suggested that Pholcus is a paraphyletic group and that at least some of the closely related genera like Panjange, Leptopholcus, Micropholcus, Micromerys, Calapnita, and Uthina Simon, 1893 may in fact be nested within Pholcus, more closely related to some 'basal' groups of Pholcus than to the core group.Taxonomic consequences (synonymization of existing genera and/or creation of new genera) were not implemented because the cladistic analysis seemed too preliminary to justify such formal decisions.The same reasoning lies behind our decision to place the new species in Pholcus, especially the two new species groups (the Pholcus ethagala group was already described in Huber 2011): these new taxa do clearly not belong to any of the nine genera closely related to Pholcus, and creating new genera for them might unnecessarily complicate future taxonomic decisions without adding to our current understanding of relationships.At some point a split of Pholcus will probably be necessary, but in our view this should be based on a large phylogenetic analysis, ideally including molecular data, all species groups of Pholcus, and a large sample of the nine closely related genera.Such an analysis is currently in progress (A. Valdez-Mondragón, B.A. Huber & D. Dimitrov, unpublished data).
Interestingly, the taxonomic problem with Pholcus is largely an Indomalayan problem, and it seems to be closely related to evolutionary shifts of microhabitat.All Afrotropical species groups, the single Nearctic species group, and all Palaearctic species groups of Pholcus are quite unambiguous representatives of the core group of Pholcus (Huber 2011).Within this core group, the large majority of species occupy large sheltered spaces among rocks, logs, buttresses, etc.; very few species occupy the leaf litter and ground substrate (e.g., P. kribi Huber, 2011 in Cameroon; some members of the P. opilionoides group in North Africa, Huber 2011) or live leaves (e.g., the P. taarab group in east Africa and the P. debilis group in Central Africa; Huber 2011).even these ecologically exceptional species and species groups are unambiguous representatives of the core group of Pholcus.
By contrast, the nine species groups assigned tentatively to Pholcus are all Indomalayan: P. andulau group, P. buatong group, P. domingo group, P. ethagala group, P. halabala group, P. kerinci group, P. krabi group, P. minang group, P. tambunan group (Huber 2011;Huber et al. 2016;herein).Most of the species included in these species groups occupy either the leaf litter of live leaves; exceptions are a few cave-and rock-dwelling species such as P. hurau Huber, 2011;P. diopsis Simon, 1901;P. kuhapimuk Huber, 2016;P. khaolek Huber, 2016;P. sudhami Huber, 2011;andP. buatong Huber, 2016 (Huber 2011;Huber et al. 2016).Three of the problematic genera possibly nested within Pholcus are also leaf-dwelling and Indomalayan (partly reaching Australia): Calapnita, Panjange, and Micromerys.A possible explanation to be tested in future phylogenetic analyses is that Pholcus in Southeast Asia has a longer history of leaf-dwelling than in tropical Africa, resulting in more conspicuous deviations from the 'typical' morphology of Pholcus, which in turn obscured relationships and prompted researchers to

Fig. 1 .
Fig. 1.Known distributions of the Pholcus ethagala group (Malay Peninsula and Sumatra; Sri Lankan species not shown), of the P. tambunan group (Borneo) and of the P. domingo group (Philippines).
HUBER B.A. et al., Southeast Asian leaf-and litter-dwelling Pholcus create possibly unjustified new genera.Future large scale analyses of Pholcus should thus continue to focus on Indomalayan taxa.

Figs 2-9. Live specimens. 2-3. Pholcus phui
Southeast Asian leaf-and litter-dwelling PholcusAll specimens were collected within a small swampy area in the forest that was characterized by numerous large dead leaves on the ground.Almost each of the largest leaves had an adult Pholcus phui on its lower surface, while the more numerous smaller leaves were mostly either unoccupied or occupied by juveniles.No further specimens were found during three days of collecting in various parts of the forest.