Identify caterpillars and, if possible, parasitoid (Sabah, north Borneo)

Someone on Twitter posted some photos yesterday of a lump of mud that, when knocked off a chair by accident, revealed a number of caterpillars:

I've uploaded one of the photos below. The assumption (which seems reasonable, but I suppose isn't certain) is that they were left there by a parasitoid.

Can anyone identify the caterpillars, confirm whether they are likely to have been left by a parasitoid and if so identify a likely culprit? The sample was from a forested area in Sabah, northern Borneo; I've requested a size from the person who posted the photos.

These are wasp larvae, but likely not parasitoids. They are the larvae of predatory Sphecid or Crabronid wasps, both of which build mud nests, generally provisioned with spiders or other arthropods that have been killed (and by some wasp species, dismembered) or paralyzed by the female (mother) wasp. It looks like one or two of them may be pupae (closer to eclosing--"hatching"--as adults) instead of larvae. If kept carefully they can be raised to adulthood for identification (at least by an expert familiar with the aculeate, or stinging, wasp fauna of that part of the world).

Effect of habitat transformation from grassland to Acacia mangium plantation on dung beetle assemblage in East Kalimantan, Indonesia

Clean Development Mechanism afforestation often involves the creation of fast growing tree plantations on non-forest lands. To estimate the possible impacts of afforestation on the biodiversity of local species, we compared the diversity of dung beetles collected using baited pitfall traps placed in grasslands, plantations of Acacia mangium, and intact natural forests in East Kalimantan, Indonesia. The species richness in plantations was higher than that on grasslands but lower than that in intact natural forests. Ordination analysis revealed that the structures of beetle assemblages in plantations were intermediate between intact natural forests and grasslands. However, the indicator species for the intact natural forests were never or rarely seen in the plantations. These results suggest that afforestation increases the local native diversity of dung beetles but that plantations are not readily colonized by the indicators of intact natural forests. Conversely, it is suggested that afforestation decreases the abundances of two grassland specialists.

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Penyerpihan habitat semulajadi boleh memudaratkan sesuatu spesies jika ianya gagal menyeberangi sempadan bukan habitat bagi mencapai lokasi baharu yang seterusnya boleh mengurangkan fungsi habitat semulajadi hutan yang bersambungan. Habitat yang bersambung adalah penting bagi meningkatkan julat pergerakan spesies ketika perubahan iklim dan oleh itu, adalah amat penting untuk mengetahui faktor-faktor yang mungkin menghalang pergerakan mereka dalam landskap yang telah diubahsuai oleh manusia. Di kawasan tropika, perkembangan pesat pertanian telah menyebabkan banyak hutan hujan semulajadi menjadi habitat yang terserpih, dan ia akan menyebabkan potensi sesuatu spesies yang mempunyai kepelbagaian biologi yang tinggi dan amat bergantung terhadap habitat hutan hujan tersebut menjadi terasing. Kebarangkalian untuk menyeberangi sempadan bukan habitat adalah penting dalam menentukan penyebaran spesies di kawasan landskap hutan hujan yang terserpih dan oleh itu, penyelidikan tentang pergerakan spesies di kawasan hutan hujan dan sempadan ladang kelapa sawit di Borneo dengan menggunakan taksa kupu-kupu famili Nymphalidae sebagai model organisma akan dikaji. Sebanyak 1666 individu dari 65 spesies kupu-kupu telah ditandai dan sebanyak 19 peratus (100/527) adalah individu yang telah melintasi sempadan bukan habitat. Sesetengah spesies kerap melintasi sempadan bukan habitat dan pergerakan mereka adalah dari kawasan hutan hujan ke ladang kelapa sawit. Walaubagaimanapun, penyeberangan sempadan bukan habitat dari hutan ke ladang kelapa sawit bagi spesies kupu-kupu yang berjaya ditangkap semula yang telah dikesan adalah kurang daripada 50 peratus (12/28) dan ianya dikuasai oleh spesies kupu-kupu bersaiz kecil yang mempunyai tumbuhan perumah larva di kawasan ladang kelapa sawit. Secara amnya, ladang kelapa sawit boleh bertindak sebagai penyekat kepada pergerakan sesuatu spesies yang bergantung penuh terhadap habitat hutan hujan (cthnya spesies yang mempunyai keperluan khusus kepada habitat hutan hujan untuk pembiakan) dan kepentingan untuk memastikan habitat hutan terus bersambung bagi memulihara spesies-spesies hutan hujan wajar ditonjolkan.

Across the globe , natural habitats are being fragmented by human activities with detrimental consequences for biodiversity (Canale et al. 2012 , Melo et al. 2013 , Almeida-Gomes et al. 2016 ). Habitat connectivity is important for population persistence (Hanski 1999 ), and species are predicted to shift their ranges in response to climate change (Chen et al. 2011 ), making it important to understand the permeability of fragmented landscapes (Hodgson et al. 2011 ) and to maintain landscape connectivity (Martensen et al. 2008 ). Loss of connectivity is of particular concern in tropical regions (Wade et al. 2003 ) because rain forests are global hotspots for biodiversity but have already experienced extensive deforestation (Gibbs et al. 2010 ). For example, in parts of Southeast Asia, fragmentation of lowland forest is primarily due to the expansion of large-scale oil palm plantations (Elaeis guineensis Jacq.) (Gaveau et al. 2014 ), which can lead to the isolation of populations of forest-dependent species in the remaining areas of forest within these landscapes (Scriven et al. 2015 ).

The ability of species to move between habitat patches depends on species dispersal ability, a complex process that integrates the physical costs of movement through preferred habitat (Bonte et al. 2012 ), the response of species to habitat boundaries (Kallioniemi et al. 2014 ), and the permeability of the matrix (Perfecto & Vandermeer 2002 ). For tropical forest species to disperse successfully through fragmented habitats, they need to cross forest–non-forest edges, which are frequently avoided by forest specialists (e.g., Laurance 2004 , Watson et al. 2004 ). Thus, an important component of dispersal involves species behavior upon reaching the forest edge, and responses to habitat boundaries affect emigration rates from suitable habitat (Ries & Debinski 2001 ). Boundary crossing by individuals (e.g., butterflies) may be part of a random walk or movement (e.g., see Schultz et al. 2012 ), although it is also likely that crossing may represent an active decision by an individual to leave areas of suitable habitat, and so the likelihood of crossing an edge may be an indicator of dispersal ability. However, leaving areas of suitable habitat may not always indicate longer distance dispersal (see review by Stevens et al. 2010 ), but boundary crossing is a prerequisite for individuals moving through highly fragmented landscapes.

While some tropical forest species avoid forest edges (Hansbauer et al. 2008 ), there is little information on the variation in boundary crossing among species. In temperate regions, species have been shown to recognize boundaries between suitable and unsuitable habitat and can actively control their rate of boundary crossing (Conradt & Roper 2006 ) and modify their movement behavior in response to boundaries (e.g., birds: Rodríguez et al. 2001 , butterflies: Schultz & Crone 2001 , bush crickets: Berggren et al. 2002 , and salamanders: Rittenhouse & Semlitsch 2006 ). Several temperate studies of butterflies have also reported species-specific differences in boundary-crossing ability (e.g., Haddad 1999 , Ries & Debinski 2001 , Kallioniemi et al. 2014 ), and differences among species in their overall levels of activity can also affect rates of boundary crossing (Mair et al. 2015 ). Thus, current evidence implies that tropical species may vary in their sensitivity to habitat boundaries, and hence to rain forest fragmentation effects, but data quantifying movement of species across rain forest boundaries and how ecological traits influence edge-crossing behavior are lacking.

The movement of individuals across a habitat boundary is predicted to follow productivity (Rand et al. 2006 ) and population source-sink (Pulliam 1998 , Tscharntke et al. 2005 ) gradients. In both tropical (e.g., Lucey & Hill 2012 ) and temperate (e.g., González et al. 2015 ) regions, there is evidence of spillover from natural habitats into managed systems, although spillover can also occur in the opposite direction (Barcelos et al. 2015 ). Studying net movement of individuals across rain forest-agricultural boundaries is important for understanding species diversity and ecosystem functioning for example, if forest pests move into plantations and reduce crop yields or if crop-dwelling predators move into forests and reduce biodiversity (Rand et al. 2006 ).

Conversion of rain forest to oil palm agriculture reduces tropical biodiversity (Fitzherbert et al. 2008 ) and remaining tracts of rain forest become isolated within agricultural landscapes (Scriven et al. 2015 ). In order to develop effective conservation management, there is a pressing need to determine the permeability of forest-oil palm plantation boundaries to forest-dependent species (i.e., species that are dependent on forest habitat to breed). If forest species are unable to cross forest boundaries, then plantations will form barriers to the movement of individuals among forest patches, thereby reducing habitat connectivity for these species. We investigated the movement of species at forest-oil palm plantation boundaries and tested the hypotheses that net flow of individuals is from forest into plantations, and that plantations are barriers to movement of many forest-dependent species hence, we predicted fewer overall movements of species from forest into plantations compared with movements within forest. In addition, we predicted that plantations will be less of a barrier to species whose larval host plants occur within the plantation, and we also examined whether other species traits (forewing length, larval host plant specificity, and geographic range size) affected boundary crossing. We selected these traits for study because they have previously been shown to affect the sensitivity of tropical butterfly species to forest fragmentation (Benedick et al. 2006 ). Our study taxon was nymphalid butterflies, which are diverse (Benedick et al. 2006 ), relatively mobile (Marchant et al. 2015 ), and many species are dependent on closed-canopy forest (Hill et al. 2001 ). Butterfly distributions have also been shown to correlate well with observed patterns in other taxa (Schulze et al. 2004 , Thomas 2005 , Gardner et al. 2008 ), and so butterflies are considered sensitive ecological indicators of environmental changes (Cleary 2004 ).


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The greatest part of the material examined for this study is preserved in the collection of the Natural History Museum, London (BMNH), with significant studied material also in the Biosystematics Laboratory, Kyushu University, Fukuoka (BLKU). During this work, more than 8500 museum specimens were examined, and approximately 200 genitalia dissections prepared. In addition, research was undertaken on the extensive type material of these butterflies held in the BMNH collections to ensure that, as far as possible, the species group names applied are typified correctly and appropriate to employ.

Other material examined is located in the Entomological Laboratory, Faculty of Agriculture, Kyushu University (AGKU), Muséum National d’Histoire Naturelle, Paris (MNHN), Oxford University Museum of Natural History (OUMNH) and the Bishop Museum, Honolulu (BPBM).

Genitalia preparation and terminology

For the preparation of genitalia, either the entire abdomen or posterior half of the abdomen was removed, macerated in 10% aqueous KOH, and dissected in water using a binocular microscope. Except where noted, genitalia drawings were executed using a camera lucida from the entire genitalia or single parts submerged in a Petri dish of water, without any compression by glass slide and cover slip. For better contrast, some preparations were stained with Chlorazol Black. Terminology for male genitalia is based on Shirôzu's (1960: 1–10) extensive account, except that we use the term phallus instead of the more frequent ‘aedeagus’, as endorsed by Kristensen (2003) . Terminology for female genitalia mainly follows van Son (1949) , with some additions from Kusnezov (1915) and Yamauchi & Yata (2004) .

Wing venation terminology

The Comstock–Needham wing-vein and cell nomenclature adopted in the descriptions is based on Nielsen & Common (1991) and Smith & Vane-Wright (2001) . This terminology, together with the numerical system employed by Yata (1981) and many other lepidopterists (e.g. Corbet & Pendlebury, 1992 ), is illustrated in Fig. 1.

Wing venation of Appias (Catophaga) paulina, showing both the Comstock–Needham terminology and the numerical system (small ciphers) for the long veins. The short cross-veins closing the discal are notated according to common lepidopterological practice: upper, middle and lower discocelluar veins (udc, mdc, ldc). The cells are notated using the Comstock–Needham system only. Dotted lines in the discal cells indicate ‘folds’ (probable courses of proximal parts of veins M1–M3), and in CuA2 the lost vein CuB (which supposedly appears during early development but is later resorbed). Based in part on Smith & Vane-Wright (2001: 513 , fig. 7).


This study suggests that vertical sampling is important to obtain accurate measurements of the insect fauna in managed forest stands. The vertical distribution of insects was significantly different among the three management types studied. A greater proportion of the insect fauna was recovered close to the ground in clearcut stands than in selection and shelterwood stands. There was a greater decline in insect richness with increasing trap height in the clearcut stands than in the selection and shelterwood stands ( Table 1 Fig. 2) and traps above 5 m recovered a greater number of additional morphs in shelterwood and selection stands than in clearcut stands ( Table 2 Fig. 3). Furthermore, the individual insect morphs that showed differing height associations among the management systems and associations for the top traps in this study provide further evidence that sampling along a vertical gradient is important (Tables 3 and 4).

The detected variability in the vertical distribution of insects may reflect trap placement in relation to the canopy of the stands. Despite variation in canopy height the greatest abundance and diversity of insects has been found within and just above the canopy of the forest ( Sutton and Hudson 1980, Sutton 1983, Sutton et al. 1983, Kato et al. 1995). In our study, the top traps were above the canopy in clearcut stands, within the canopy in shelterwood stands, and just into the bottom edge of the canopy in selection stands. In clearcut stands, the paucity of additional morphs in the top traps and sharper decline in insect richness with increasing trap height suggests that the lower traps adequately sampled the canopy fauna. Conversely, the greater proportion of additional insect morphs collected in the top traps of shelterwood stands indicates the top traps were sampling the relatively taller shelterwood canopy. In the selection stands the top traps sampled only the bottom edge of the canopy. To compare the insect communities of forest stands with differences in stand structure vertical sampling must be done within comparable physiognomic classes of vegetation.

We thank J. Brisette, D. Debinski, F. Drummond, W. Haltemann, J. N. Jaros, and A. White for thoughtful reviews that significantly improved the manuscript. Considerable technical assistance was provided by the staff at the Penobscot Experimental Forest. Voucher and reference collections are stored at The University of Maine Insect Museum. This research was funded by USDA National Research Initiatives award no 96-35106-3725, USDA Forest Service cooperative agreement no. 23-938, and the University of Maine's Forest Ecosystem Research Project, Department of Biological Sciences, and Department of Wildlife Ecology. Maine Agriculture and forestry experiment station publication No. 2458.

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